Fossils in

Earth Sciences

Anis Kumar Ray

 Fossils in
Earth Sciences

Anis Kumar Ray

Formerly, Reader in Geology
Presidency College
Kolkata

New Delhi-110001
2008
FOSSILS IN EARTH SCIENCES
Anis Kumar Ray

© 2008 by PHI Learning Private Limited, New Delhi. All rights reserved. No part of this book may be
reproduced in any form, by mimeograph or any other means, without permission in writing from the
publisher.

ISBN-978-81-203-3432-8

The export rights of this book are vested solely with the publisher.

Published by Asoke K. Ghosh, PHI Learning Private Limited, M-97, Connaught Circus,
New Delhi-110001 and Printed by Jay Print Pack Private Limited, New Delhi-110015.
 Contents
Preface ................................................................................................................................................ xiii

PART ONE PRINCIPLES

1. INTRODUCTION ............................................................................................................... 3–12
1.1 Defining Fossil 3
1.2 Defining Branches 6

2. TAPHONOMY ................................................................................................................... 13–31

2.1 Introducing Taphonomy 13
2.2 Coming into Being of Fossils: Organisms of Biosphere Turn into Fossil
in Lithosphere 19
2.3 Fossil Lagerstätten 27
2.4 To Read Out the Taphonomic History 28

3. PALAEOECOLOGY ........................................................................................................ 32–51

3.1 Introducing Palaeoecology 32
3.2 Essential Terms and Concepts 33
3.2.1 Palaeoautecology and palaeosynecology 33
3.2.2 Habits and habitats 35
3.2.3 Limiting factors 37
3.2.4 Adaptation 38
3.2.5 Echinoid adaptation 40
3.2.6 Adaptation and functional morphology 40
3.2.7 Ancient community to fossil assemblage 43
3.2.8 Palaeoecology redefines its course 44

iii
iv Contents

3.3 On Methodology of Palaeoecology 45

3.3.1 Palaeoecology works on simple premise of principle of
uniformitarianism 45
3.3.2 Some major methods 47
3.4 Appendix: Marine and Terrestrial Environments Contrasted 50

4. SYSTEMATICS AND BIOSTRATIGRAPHY ............................................................... 52–72

4.1 Systematics: Introduction 52
4.2 Types 54
4.3 Naming Species 54
4.4 Species Concept and Species Problem 57
4.4.1 Dual problem 57
4.4.2 Typological species concept 58
4.4.3 Biological species concept 60
4.4.4 Evolutionary species concept 61
4.4.5 Phylogenetic species concept 62
4.5 Speciation or Origin of Species 65
4.5.1 Phyletic gradualism 65
4.5.2 Punctuated equilibria 65
4.6 Biostratigraphy 66
4.6.1 Background material 66
4.6.2 Introducing biostratigraphy 66
4.6.3 Essence of biostratigraphy: index or guide fossils 68
4.6.4 Relationship with other stratigraphic units and correlation 71

5. EVOLUTION OF ORGANISMS ................................................................................... 73–112

5.1 Introduction: Evolution 73
5.2 Organic Evolution 74
5.2.1 Two modes of evolution 74
5.2.2 Rates of evolution 74
5.2.3 Rates of evolution, appearance and extermination in evolution 75
5.2.4 Extinction 78
5.2.5 Divergence and convergence 79
5.3 Palaeontology in the Study of Organic Evolution 79
5.4 Introduction: Proboscidea, Equidae and Hominidae 80
5.5 Revised Views on Vertebrates 81
5.5.1 Phylogenetic systematics and molecular phylogeny 81
5.5.2 Some revisions 81
5.6 Order Proboscidea 82
5.6.1 Introduction 82
5.6.2 First proboscideans 82
5.6.3 Phenotypic characteristics and trends 82
5.6.4 Proboscidean radiation 83
5.6.5 Trends in elephantid evolution 84
5.6.6 Mammoths and extinction of the order 84
 Contents v

5.7 Family Equidae 85

5.7.1 Introduction 85
5.7.2 Ancestry 86
5.7.3 General observation 86
5.7.4 Phenotypic trends 86
5.7.5 Migration and evolution of equidae 90
5.8 Family Hominidae 91
5.8.1 Classical view on humans in the organic world 91
5.8.2 Tertiary apes 91
5.8.3 Changed view on phylogeny 92
5.8.4 Different genera and species of ‘hominids’ 92
5.8.5 Evolutionary stages: Are there any? 94
5.8.6 Species of Australopithecus and Homo contrasted and compared 95
5.8.7 Differences between apes and man: Phenotypic distinction of humans 97
5.8.8 Human ancestry: Revisited 103
5.8.9 A probable scenario 107
5.8.10 Appendix: On man’s uniqueness 110
5.9 General Comments 111

6. MAJOR EVENTS OF HISTORY OF LIFE ............................................................... 113–122

6.1 Stages in the History of Life 113
6.2 Some General Difficulties 114
6.3 Origin of Organic World and Early Stages of Its Evolution 114
6.3.1 Two fundamental characteristics of life 114
6.3.2 Experimental Verification of pre-biotic happenings 114
6.3.3 Cells and organic evolution 115
6.3.4 Key role of environment 116
6.3.5 Evidence of early life 116
6.3.6 Chemotrophs, autotrophs or heterotrophs 117
6.3.7 Internal and external environment fortified
the organic world and opened new vistas as well 117
6.4 Diversification Ensues and Continues 118
6.4.1 Multicellularity and biomineralization 118
6.4.2 Diversification in water: Vertebrates reign supreme 119
6.4.3 Adaptive radiation, extinction and advent on
land shaped the organic world through Palaeozoic 119
6.4.4 Mesozoic ushered in modernization 120
6.5 Major Mass Extinctions: Important Component in the History of Life 121

PART TWO MAJOR INVERTEBRATE GROUPS

7. PHYLUM CNIDARIA .................................................................................................. 125–139
7.1 Introduction 125
7.2 Cnidaria in Palaeontology: Importance of Polyp and Medusa 125
vi Contents

7.3 A Framework for Morphology 127

7.3.1 Shape of corallites 128
7.3.2 Colony and shape, and arrangement of corallites 128
7.3.3 Economy of space and closer interconnection
control arrangement in colonies 130
7.3.4 Wall 132
7.3.5 Internal features 132
7.3.6 Septa 132
7.3.7 Other structures 134
7.3.8 Septa and classification of Anthozoa 135
7.4 Geological Importance of Anthozoa 137

8. COILED SHELLS: AN INTRODUCTION ................................................................ 140–148

8.1 Introduction 140
8.2 Molluscan Body Plan and Variation 141
8.3 Shell Growth and Its Computer Simulation Model 143
8.4 Univalved and Bivalved Shell 144
8.5 Orientation and Symmetry of Bivalved Shells 145

9. BRACHIOPODA ........................................................................................................... 149–161

9.1 Introduction 149
9.2 The Valves 149
9.3 Appearance and Measures 150
9.3.1 Orientation 150
9.4 Brachiopod Hinge 153
9.5 Pedicle and Its Opening 154
9.6 Internal Features 154
9.6.1 Brachiopod musculature 154
9.6.2 Lophophore and brachidium 156
9.7 Mineralogy and Microstructure of Shells 156
9.8 Punctation of Shells 157
9.9 Surface Features 157
9.10 Additional Information Beyond Morphology 159
9.10.1 Ecology and palaeoecology 159
9.10.2 Classification 160
9.10.3 Affinity and brief history 161

10. BIVALVIA (MOLLUSCA) ............................................................................................ 162–175

10.1 Introduction 162
10.2 Morphological Variation: Adaptation, the Cause 162
10.3 Shape, Dimensions and Orientation 162
10.4 Umbo and Beak 163
10.5 Hinge and Dentition 163
10.6 Hinge Plate and Hinge Area 165
 Contents vii

10.7 Variations in Dentition 165

10.8 Adductor Muscles and Ligament 168
10.8.1 Opening and closing of shell 168
10.8.2 Variations in adductor muscles 169
10.8.3 Variations in ligament 169
10.8.4 Pallial line and sinus 170
10.8.5 Significance of pallial sinus and pedal scar 170
10.8.6 Surface ornaments 171
10.9 Bivalve Adaptation: Functional Morphology of Shells 171
10.9.1 Recent eco-morphotypes: Endobenthic types 172
10.9.2 Epibenthic types 173
10.9.3 Evolution of adaptation 174
10.10 Some Other Aspects 174

11. GASTROPODA (MOLLUSCA) ................................................................................... 176–185

11.1 Introduction 176
11.2 Two Important Characteristics of Body 177
11.3 Morphology: Coiled Shells 177
11.4 Compactness of Coiling 179
11.5 Orientation, Dimension and Shape 179
11.6 Functional Morphology of Gastropod Aperture 180
11.7 Internal Structures 183
11.8 Surface Ornaments 184

 12. CEPHALOPODA (MOLLUSCA) ................................................................................ 186–209

12.1 Introduction 186
12.2 Cephalopod Groups and Their Shells 186
12.3 Shape of Shells 186
12.4 Aperture, Columella, Umbilicus and Ornaments 188
12.5 Internal Structures 193
12.6 Septa, Suture, Camera 193
12.7 Siphuncle Draws the Main and Fundamental Difference with Gastropods 193
12.8 Predatory Habit and Developed Brain 194
12.9 Movement Control in Cephalopods 194
12.10 Contradiction between Weight and Buoyancy:
Stable Posture of Swimming Animals 195
12.11 Pressure at Depth and Strength of Shell 195
12.12 Shell Shape and Posture 195
12.13 Case of Ammonoids 195
12.14 Ammonoid Suture and Heteromorphy 196
12.15 Dimorphism in Ammonoids 199
12.16 Ammonoid Palaeobiogeography 199
12.17 Ammonoid Palaeoecology 201
12.18 Indian Case Studies 202
viii Contents

12.19 Evolution of Ammonoidea 203

12.19.1 Biostratigraphic and evolutionary significance 203
12.19.2 Ancestry of ammonoids 203
12.19.3 Phylogeny 203
12.19.4 Phenotypic trends 204
12.19.5 Indian ammonoids 205
12.19.6 General comments 206

13. ECHINOIDEA (ECHINODERMATA) ........................................................................... 210–230

13.1 Introduction: Characteristics of the Phylum 210
13.2 Subdivisions and Brief History 212
13.3 Introduction: Echinoidea 212
13.4 Adaptation and Symmetry 215
13.5 Water-Vascular System 215
13.6 A Format for Morphology 217
13.7 Shape-Size-Symmetry, etc. of Test 217
13.8 Corona: Ambs and Interambs 221
13.9 Interamb, Tubercles and Fascioles 224
13.10 Peristomial System, Aristotle’s Lantern 226
13.11 Periproct and Apical Disc 227
13.12 Mode of Living and the Plate Systems 227
13.13 Brief Phylogeny 229
13.14 Appendix: Echinoidea 229
13.14.1 Characters of a few genera and suggesting
functional morphology on them 229

14. TRILOBITA (ARTHROPODA) ................................................................................... 231–241

14.1 Introduction: Arthropoda 231
14.2 Introduction: Trilobita 232
14.3 Three Lobes and Segments 233
14.4 Cephalon 234
14.5 Thorax and Pygidium 236
14.6 Chemical Composition of Carapace 239
14.7 Palaeobiogeographic and Stratigraphic Use of Trilobites 239
14.8 Indian Record 240

PART THREE MISCELLANEOUS

15. MICROFOSSILS ........................................................................................................... 245–256
15.1 Introduction: Definition 245
15.2 Basic Varieties 246
15.3 Use of Microfossils: Systematics and Taxonomy 247
15.4 Use: Biostratigraphy 250
15.5 Use: Palaeoecology and Microfacies 250
 Contents ix

15.6 Use: Hydrocarbon Prospecting and Exploration 253

15.7 Microfossils and Lithogenesis 253
15.8 Microfossils and Hydrocarbon Generation 255

16. MICROFOSSILS: FORAMINIFERA ........................................................................ 257–263

16.1 Introduction 257
16.2 Foraminiferal Ecology 258
16.3 Some Morphological Details 259
16.4 Brief History 263

17. MISCELLANEOUS FOSSIL GROUPS ..................................................................... 264–278

17.1 Introduction 264
17.2 Porifera and Bryozoa 264
17.3 Pteropoda 266
17.4 Sessile Echinoderms 266
17.5 Chelicerata 267
17.6 Crustacea 268
17.7 Graptolites 268
17.7.1 Introductory remarks 268
17.7.2 Stratigraphical use 270
17.7.3 How did graptolites live? 271
17.8 Trace Fossils 272
17.9 Early Life and Stromatolites 274
17.9.1 Stromatolites: what, where and how 275
17.9.2 Indian examples 277

18. VERTEBRATA OF CHORDATA ................................................................................ 279–320

18.1 Introduction 279
18.2 Debate Starts from “Where to Start” 280
18.3 A Necessary Digression to Clinch the Issue of Vertebrate Origin 280
18.4 Vertebrates Through Ages 282
18.5 Basic and Major Features of Vertebrate Body and Skeleton 284
18.6 Early Innovations 285
18.7 Advent of Tetrapods 285
18.7.1 Tetrapods and Amphibians 285
18.7.2 Problems of living on land: Support 285
18.7.3 Problems: Locomotion 286
18.7.4 Feeding and respiration 286
18.7.5 Sensory systems and water balance 286
18.7.6 Reproduction 286
18.7.7 Fossil evidences 286
18.7.8 Carboniferous scenario 287
18.7.9 Summary narrative on origin of tetrapods 287
18.7.10 Hypotheses on transition 288
x Contents

18.8 Early Amniotes 288

18.9 Importance of Triassic in Tetrapod Evolution 289
18.10 Dinosaurs 289
18.10.1 Introducing the terrible lizards 289
18.10.2 Origin of dinosaurs 291
18.10.3 End triassic dinosaurian radiation
sparks off a macroevolutionary debate 291
18.10.4 Unusual fossils 297
18.11 Extinction of Dinosauria: Reality of a Myth 297
18.11.1 Gradual or catastrophic 297
18.11.2 Bolide impact theory 299
18.11.3 Volcanism theory 300
18.11.4 Debate changes to ‘how catastrophic the event was’ 300
18.11.5 New facts and views 300
18.12 Origin of Mammals 303
18.13 Vertebrates and Palaeogeography-Palaeobiogeography 303
18.14 Indian Records 305
18.14.1 Non-mammalian vertebrates 305
18.14.2 Gondwana vertebrates 305
18.14.3 Indian dinosaurs 311
18.14.4 Vertebrates from Lameta Beds and Deccan Intertrappeans 312
18.14.5 Important mammalian fauna of India 312
18.14.6 Palaeoclimatic interpretations of Siwalik vertebrates 314
18.14.7 Bugti bone beds of Pakistan 316
18.14.8 Extinction of Siwalik mammals 317

19. PLANTA ......................................................................................................................... 321–349

19.1 Introduction 321
19.2 Appreciation of Plant Fossil Record 322
19.3 Scope of the Chapter 324
19.4 Indian Record of Land Plants 324
19.5 Some Relevant Questions on Gondwana Stratigraphy 341
19.6 Brief Critical Appraisal of Indian Record 344
19.7 Calcareous Algae 348

PART FOUR SOME SYNTHESIS

20. FOSSILS AND CLIMATE, GEOGRAPHY, BIOGEOGRAPHY,

AND ECOLOGY OF THE PAST ................................................................................ 353–367
20.1 Introduction 353
20.2 Palaeogeography 354
20.2.1 Brief history from continental drift to plate tectonics 354
 Contents xi

20.3 Palaeobiogeography 355

20.3.1 Modern biogeographic divisions 355
20.3.2 Two models of biogeographic changes 357
20.4 Palaeoclimatology 369
20.4.1 Some facts about climate 360
20.4.2 Methods and examples 360
20.4.3 Indian examples 362
20.4.4 Some recent studies 362
20.4.5 Stable isotope studies in palaeoclimatology 364

PART FIVE APPENDICES

1 Fossil Lagerstätten ................................................................................................... 371–387

2 Lab Exercises ............................................................................................................ 388–402
3 Indian Stratigraphy for Palaeontologists ................................................................ 403–408

Epilogue .................................................................................................................................. 409

References ....................................................................................................................... 411–421
Index ............................................................................................................................... 423–429
 Preface
The title of the book conveys the purpose. It is a textbook, dealing with various aspects of the subject of
palaeontology, written mainly for undergraduate students of geology. The span and scope of the book
have been set largely in accordance with the demands and limits of the curricula generally adopted in
Indian universities. Yet both have often been freely transgressed in the book with a view to accommodate
updated information and concepts. At the same time, I feel that palaeontology students in India basically
face a twofold problem. On the one hand, there is a need for a convenient compilation of information,
particularly on Indian fossils. This is perhaps easier to address. The second part of the problem is more
latent, yet vital. How is the subject to be learnt and, for that matter, taught? Should palaeontology be
treated as mainly a wealth or a junk of biological names and terms, as it is often looked at? Obviously
the answer is in the negative. The subject needs to come out of the ‘glossary’ identity, or ‘fossils can
also be used for these purposes’ sort of narratives. I have made an attempt in these two regards. But how
successful this attempt has been can be judged from students taking interest in the book, even if their
number is few.
The book is divided into five parts and is meant to cater the subject to the readers in a way that does
not hamper their interest. Part One deals with the principles and various classification of fossils.
Part Two with major invertebrate groups while Part Three with other non-invertebrate groups.
In Part Four synthesis is done and Part Five includes the appendices. Background information have been
presented in factsheets or appendixes chapters or sections, without adding extra load on the main
discussion. As it is demanded from the community of target readers in different Indian academic institution,
in the book emphasis is given on morphology of major invertebrate fossil groups. Non-invertebrate
organisms are treated partially on essential points without details. Principles and methods are dealt with
both separately and intermingled in the discussion of morphology. The topics are discussed with a view
to make students ask questions as far as possible. Answers are there in the text, not for the readers to
cram, but search and frame for themselves. In addition, I have left the materials included and treated
sometimes at a level set slightly above, yet reachable, for a student-reader with a hope that it may
generate a drive in him or her to arrive at it. This is a process that really helps students rise from where
they start, and to be set in students by their teachers as I have realized from my nearly four decade long
teaching career. For obvious reasons, the book, however, cannot be used for research-references.

xiii
xiv Preface

I would like to acknowledge the inspiration, encouragement, grooming, including assistance that I
received from my teachers, colleagues, friends and, more particularly, my students during the preparation
and production of the book. Mentioning any name, in particular, may be unfair and, thus, unwarranted.
Two names are, however, unavoidable for different reasons. The first is of (late) Professor H. C. Dasgupta,
who set the tradition of and kindled the urge for enriching teaching-learning with research in Presidency
College and Calcutta University nearly a century back; his prime interests were in palaeontology and
stratigraphy. I owe to the latent the urge to follow that tradition. The second is of (late) Professor
S. Ray, a worthy student of Professor Dasgupta, my teacher-mentor. His exceptional acumen in teaching
prepared, created and, finally, strongly founded the system of thinking-reasoning-expressing for teaching
with continuous vigilance and constantly improve it for the development and progress of the science on
the basis of feedback from students. If the book is of any worth, it is built on the edifice erected by these
two giant teachers; wherever it falls short of its mark, the responsibility certainly devolves upon me.
The book is written largely on the financial assistance from UTILISATION OF SCIENTIFIC
EXPERTISE OF RETIRED SCIENTISTS (USERS) Scheme (HR/UR/29/2003 dt. 08-07-2004)
of the Department of Science and Technology, Government of India.
A few of the figures of the book are acquired from different publications and are incorporated in
modified, redrawn and compiled form. Permissions from the publishers are thankfully acknowledged.

Anis Kumar Ray

 Part one
Principles
 1
Introduction
1.1 Defining Fossil
Palaeontology is the branch of science that deals
with life and organisms of the geological past and
the environments the latter lived in. The geological
past has a long span. So, palaeontology also covers
how life, organisms and their environments
changed in an interrelated and interacting way,
through the vast span of time, on what factors,
towards which direction, and so on.
Fossils are the tools of palaeontological
studies. They are well-known to commoners as well
as to knowledgables (Factsheet 1.1). But they are
variously defined. Fossils can be defined as
(i) objects occurring in or dug out of the earth’s
crust that (ii) were embedded or buried and
preserved by natural agencies in geologically
ancient time (Factsheet 1.2), and that (iii) give
some idea about the biological organization and
morphology, activity, habit, etc. of the organism
in question (see Shrock and Twenhofel 1953). But
expanding knowledge and idea about fossils
demand more than this simple definition of fossils.
We may then try defining fossils in a different way.
Thus, we may argue: a fossil is an inanimate
object found in, and/or recovered from the earth’s
crust, and that once in the geological past it was
the whole or part of a living organism or was made
3
by it in or on the sediment (or the enclosing
medium, if it is not sediment). It is, thus, the
evidence of existence of an organism conveyed
through its remains or traces of its activities made
during its lifetime.
So, a fossil conveys two kinds of information,
one biological and the other geological, covering
ancient time and space. One may lay more
importance on one or the other, depending on the
purpose of the study. A palaeontologist, who is
interested to know what are the different biological
groups (see Factsheets 1.3 and 1.4 for the following
terms of Linnean hierarchy, viz. phyla, class, order,
family, genera, species, etc. See Chapter 4 for
details) present in the fossil record that he or she
is working upon will be more biological in purpose
and method. Another palaeontologist who would
want to correlate two fossiliferous units, will be
performing more of a geological exercise than a
biological. The two kinds of information are,
however, closely interlinked and interdependent,
a fact that is brought out in the growth and
development of a branch of palaeontology, called
taphonomy.
But before we get into any more details about
taphonomy, we should digress to take a look at
certain other aspects. Fossils definitely make one
of the most useful tools in reconstruction of the
4 Part One: Principles

FACTSHEET 1.1
Palaeontology and ‘Fossils’ in History:
Important Events in their Studies and the Scientists Contributing

Both derived from the Greek Palaeontology; palaios (ancient), ontos (being), logos (study); fossilis (things dug
out of the earth).
Fossils: ‘half-life’ with a look of living things but without life (Greeks’ idea). However, Greek scholars like
Xenophanes and Herodotus (a few century BC) observed fossils of marine shells on the hill slopes inland and
imagined the area to have been under the sea.
Interest in fossils dwindled during the Roman empire.
Medieval scholars, too ignored fossils as ‘caprice or sport of nature’. Domination of the church between 12th
and 14th centuries controlled ideas. Albertus Magnus (a 13th century Bishop), Ristoro d’ Arezzo (a monk),
Boccaccio (a 14th century poet) and a few others, however, showed interest.
Fossils as ancient organisms preserved in the earth was suggested by:
(i) Chu Hsi in the Chinese analects of 1227 (Haq and Boersma 1978); or
(ii) Leonardo da Vinci (1452-1519), the Renaissance illuminary.
Agricola coined the word fossil in 1546, but could not give the correct idea about it.
Biblical thoughts, for example the deluge or the flood, were revived in the 17th and 18th centuries.
Definitive understanding of the true nature of fossils was reached in later parts of 18th century.
Meanwhile, Nicolas Steno (Dane) in 1669, recognized superposition of strata and significance of unconformities
and, thus, established the principles of modern stratigraphy.
Linnaeus (Swedish) in 1758 suggested method and system of naming animals.
Lomonosov (Russian) and Giraud-Soulavie (French) threw light on relative age significance of fossils.
Palaeontology grew rapidly towards the end of the 19th century:
Sowerby, Goldfuss, Munster, Cuvier, d’Orbigny, Agassiz were among the stalwarts.
The main debate hinged on between the creationist school with scientists like Cuvier, d’Orbigny and Brongniart
and others favouring divine hand in origin of organisms and the contending evolutionist school of Lamarck,
Saint-Hilaire and others, advocating natural process. The question was decided in favour of the materialist
explanation of life, the organic world and the organic evolution with the publication of The Origin of The
Species by Charles Darwin. Palaeontology emerged as the modern science and ramified into its various branches,
viz., micropalaeontology, palaeoecology, palaeobiogeography, etc.
History will remain incomplete without the mention of the frauds, it came to be infested with. A few of them are
worth mentioning:
1. Beringer in 1726 reported some fantastic fossils from Germany. Soon it was clear that those objects were
pranks played by some students; Beringer spent the rest of his life in buying back the copies of his book.
2. In 1913, a primate fossil was described from England, as a so-called ‘missing link’ between apes and man.
Detailed studies proved in 1953 that it was a fake and fraud , in which a skull of modern man was set with
the jaw of an orang-utan. The case is known as that of ‘Piltdown Man’.
3. Graptolites and certain other fossils described from the Himalayas turned out to be planted objects.
4. Protoavis reported from Antarctica as the first bird was also suspected to be a fraud.
 Chapter 1 Introduction 5

FACTSHEET 1.2
The Geological Time Scale

Number against each period/epoch represents the age of its lower boundary.
Eon Era Period Epoch
10,000 years Recent/Holocene 10,000 years
Quaternary
1.6 mil. yrs. Pleistocene 1.6 mil. yrs.
Pliocene 5
Neogene
Cenozoic 23 Miocene 23
Tertiary
Oligocene 34
Palaeogene Eocene 57
65 Palaeocene 65
Cretaceous 146
Mesozoic Jurassic 208
Triassic 245
Phanerozoic Permian 290
Pennsylvanian 323
Carboniferou
Mississippian 363
Palaeozoic Devonian 409
Silurian 439
Ordovician 510
Cambrian 570 million years
Proterozoic 2.5 billion years
Precambrian
Archean 4.6 billion years
NB: A new period, Ediacaran, has been added below Cambrian, spanning 635 to 543 million years, on the strength of its
typical biota and its boundaries being defined by Global Stratigraphic Section and Point (GSSP) method following
the International Commission on Stratigraphy (Knoll et al., 2006).

earth’s history. They help in understanding different multifarious use of fossils. They are introduced
aspects of that history, particularly in relation to here in two ways: one diagrammatically and the
the biosphere and its evolution. Different branches other by defining branches as answers to some
of palaeontology have developed based on this basic questions that the subject tries to deal with.
6 Part One: Principles

FACTSHEET 1.3 Thus, every fossil has a taphonomic history,

including its burial to enable fossilization and
Taxonomic Hierarchy preservation. So, taphonomy and biostratinomy
(study of burial processes and conditions) surround
(Linnean hierarchy modified)
the initial living organism in circles. Then again,
Superkingdom Eucaryota before any and every study on fossils, each of them
Kingdom Animalia
must be recognized, identified and named. To begin
Superphylum
Phylum Echinodermata
with, palaeontologists take help of morphology of
Subphylum Echinozoa fossils to these ends. So, morphology, taxonomy
Class Echinoidea (principles on which organisms or their fossils are
Subclass Euechinoidea classified), systematics (the entire study of
Order Cassiduloida systematization, i.e., covering identification and
Suborder classification) and nomenclature (naming each
Superfamily (-na/-ceae) identified and classified kind or unit for all future
Family (-dae) references) come next in circles around those of
Subfamily (-nae) taphonomy and biostratinomy.
Genus Stygmatopygus At the next level, the three sides of the triangle,
Subgenus
on the other hand, relate to three different aspects
Species S.elatus
Variety/subspecies
of fossils. Fossils of different organisms vary in
their biological characteristics; related branches
occupy one side of the triangle. Fossils may also
FACTSHEET 1.4 be of different time or of organisms living in
Taxonomic Categories For different places or environments on the surface of
the earth. So, branches dealing with questions of
Homo sapiens time and space aspects occupy the other two sides
Superkingdom Eucaryota 1900.0 of the triangle.
Kingdom Animalia 1000.0 All said and done, fossils of different biology,
Phylum Chordata 0500.0 time and space represent the ancient organic world,
Class Mammalia 0250.0 evolution of which (phylogeny being the group
Order Primate 0030.0 history of any kind of organisms during this evolution
Family Hominidae 0005.0 of theirs) and taxonomy and systematics on the basis
Genus Homo 0002.0
of evolutionary or phylogenetic relationships, that
Species sapiens 0000.1
Variety sapiens
is, the natural relationship of organisms, appear as
the final circle. (Similar morphology may emerge in
Figures refer to approximate age of the first
different genetically unrelated groups on account of
appearance on the earth in million years.
similar kind of adaptation; hence, classification on
morphology may lead to artificial grouping together
1.2 Defining Branches of those unrelated forms. This point will be further
treated later.) The diagram, thus, highlights the
Figure 1.1 presents the branches of palaeontology trichotomy in characteristics of fossils, at the same
diagrammatically. The ancient living organism is at time pointing to the basic relations among the
the centre of all exercises, at the focus. Circles in branches.
the diagram represent those branches which pertain Factsheet 1.5 presents the basic questions, in
to one and all fossils. Each side of the triangles, on answers to which different uses of fossils and the
the other hand, represents one kind of study. different branches developed respectively, on these
 Chapter 1 Introduction 7

uses are defined. It is self-evident. It may only be by sedimentological or other physical controls.
added that in the case of biostratinomy and While in all other cases, biological and other
diagenesis, the two branches of taphonomy, fossils, characteristics, and relations of ancient organisms
rather the remains of organisms, are controlled during their lifetime remain the issues of the
more by their dead or inanimate nature and, thus, branches to study.

Biological aspects : Org = Env.

Palaeobiology
Functional Growth and Form
Palaeoecology Morphology (Ontogeny) Ichnology

SYSTEMAT
E— IC
S
R —
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Fossils biological aspects

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* Ancient Living Organism

Fig. 1.1 Core branches of palaeontology and their interrelationships.

8 Part One: Principles

(a) (b)

(c )

Fig 1.2
(d)

Fig. 1.2 Major types of growth in invertebrates.

(a) Bivalve shell grows by accretion, (b) Cephalopod shell grows by accretion of shells and periodic
addition of septa, (c) Plates in ambs (or interambs) of echinoids grow by accretion as well as new plates are
added, (d) Trilobites grow by moulting, where each ontogenetic stage may leave an independent fossil.

FACTSHEET 1.5
Basic Questions on an Organism Answered in Palaeontology and Uses of
Fossils Defined on Them (Modified after Goldring 1999)

Basic Questions Category of Information Branches of Palaeontology

l What type of organism is it?
l When did it live on the earth?
l Where did it live?
l How did it live?
l From what (whence) and to
what (whither) did it evolve?
l With what organisms was it
associated during its lifetime?
l How and in which conditions
was it fossilised?
l In what modes and ways was it
preserved and changed?
l Morphological-systematic
l Stratigraphical-temporal
l Biogeographical-ecological
l Physiological-functional
l Evolutionary
l Community structure
l Hydraulic and stratinomic
l Diagenetic
l Taxonomy-Systematics Classification
l Biostratigraphy- Chronostratigraphy
l Palaeobiogeography-Palaeoecology
l Palaeobiology-Palaeophysiology
l Evolution-Phylogeny
l Community analysis
l Biostratinomy
l Diagenesis
 Chapter 1 Introduction 9

It may now be argued that since fossils as the
‘evidence of ancient life’ serve different purposes,
and thus have different uses in geological studies,
we may have to differentiate various kinds of
fossils. Factsheet 1.6 presents such a scheme.
Obviously the kind of fossils defined is on the
criterion we choose which, in turn, is determined
by what we wish to study about them. The most
commonly used is the biological kind
(Factsheet 1.7). It includes five kingdoms of the
organic world (Whittaker 1969), erected on the
basis of levels of organization and on three
principal means of nutrition or feeding, viz.
photosynthesis, absorption and ingestion. On these
bases, the kingdoms are Monera (Prokaryote with
anucleate cells; unicellular), Protoctista (Eukaryote
with nucleate cells; unicellular; also called
Protista), Planta (Eukaryote; multicellular,
autotrophic, i.e. feeding by photosynthesis),
Animalia (Eukaryote; multicellular, heterotrophic,

FACTSHEET 1.6
How FOSSILS may be Defined on Purpose of Study
FOSSILS Evidence of Ancient Life
On Type of Evidence On Size
Evidence of form and structure of hard Nanofossils
and/or soft part constituents of body Microfossils s.s.
BODY FOSSILS (sensu stricto)
(including moulds and casts) Microfossils s.l.
Evidence of activity made during lifetime (sensu lato)
TRACE FOSSILS Macrofossils
Evidence of chemical constituents of body
Megafossils
CHEMICAL FOSSILS
On Habit and Habitat On Biology
sessile/attached/fixosessile Monera = Procaryota
Epibenthic attached by root-like processes/rhizosessile Protoctista ü
free lying/liberosessile Planta ï
ý Eucaryota
Benthic vagrant/vagile Animalia ï
ï
boring (through hard shells, rocks or wood) Fungi þ
Endobenthic burrowing (digging in soft sediments)
nestling (living in holes/crevasses)
On Stratigraphy Evolution
Aquatic
holoplanktic Facies fossils
Zone fossils
Planktic meroplanktic
Index/Guide fossils
pseudoplanktic
Living fossils
Pelagic Neritopelagic/Euphotic: living in neritic water
Nektic Oceanopelagic: living in oceanic water
Epipelagic/Euphotic: in water < 200 m On Preservation History
Mesopelagic/Disphotic: in water < 1000 m Biocoenosis
Bathypelagic/Aphotic: in water < 4000 m Thanatocoenosis
Abyssopelagic
Ichnocoenosis
Terrestrial Taphocoenosis
10 Part One: Principles

FACTSHEET 1.7

Major Groups of Organisms: Traditional Classification of the Organic World

Taxonomic hierarchy from Superkingdom to Phyla/Divisions with a differentiation on trophic relationship

(food habit) superimposed on it; all phyla or divisions not shown: also not shown are the divisions or phyla in
Protoctista intermediate between Fungi and Animalia or questionably between Planta and Fungi (based on
Whittaker 1969; Clarkson 1998).

ORGANIC WORLD
Procaryota Eucaryota
(Anucleate) (Nucleate)
Unicellular Multicellular
Feeding by Monera (a) Protoctista (a, b) Planta (a) Animalia (b) Fungi (c) Kingdom
photosynthesis / Protista
(a) Autotrophic Protophyta :
Protozoa Sub-kingdom/
Cyanophyta Pyrrophyta Rhodophyta Phyla or
Schizomy- Dinoflagellata Chlorophyta Division
cophyta Chrysophyta Pheophyta
Eugienophyta Bryophyta
Tracheophyta

Feeding by Sarcodina Porifera

ingestion Ciliophora Cnidaria
(b) Hetero- Mollusca
trophic Brachiopoda
Echinodermata
Annelida
Hemichordata
Chordata

Feeding by Oomycota
absorption Arthropoda Basidiomy-
(c) Hetero- Trilobita cota
trophic Crustacea
Uniramia

i.e. feeding by ingestion) and Fungi (Eukaryote;
multicellular, heterotrophic, feeding by
absorption). The other kinds include those on size
(whereupon depends the method of study whether
using electron or optical microscope or not), on
types of evidence or on preservational history
(related to taphonomy), on habit and habitat (related
to ecology) and on stratigraphy and evolution. They
will be treated in more details in proper contexts.
A few more aspects may need a brief
discussion in this introductory chapter. As
mentioned, phylogeny is the group history of any
kind of organism during its evolution. On the other
hand, ontogeny is the life history of an individual,
which is marked by separate stages from the larval
to the senile or old stage. Individuals vary in
morphology through these stages and often they
are quite significant in understanding biology and

evolution of the group to which it may belong. In
fact, Haeckel’s Law or Theory of Recapitulation
was long considered an important understanding
to explain evolution. It states, “Ontogeny
recapitulates phylogeny”, meaning that different
ontogenetic stages of individuals of a group may
mark the characters (morphological, etc.) of the
successive preceding stages of evolution in the
lineage in which the concerned individuals
evolved. Thus, if A, B and C be the successive
species in an evolutionary lineage, by the said
theory, the latest descendant C will have the
ontogenetic stages 1, 2 and 3, where stage 1 will
bear the characters of species A, stage 2 of species
B and stage 3, the mature stage, will have the
typical characters of species C. Later studies have,
however, proved that the Law is not valid for at
least most of the cases and the phenomenon may
be exactly the reverse. (see Ammonoid evolution
for more details.) Factsheet 1.8 shows the terms
for different ontogenetic stages; accompanying
them are also the terms, used for the ontogenetic
stages of trilobites, rather arthropods.
FACTSHEET 1.8
Ontogenetic Stages
Different ontogenetic stages of an individual are
termed:
Embryonic/larval
Nepionic
Neanic
Ephebic/adult
Gerontic/senile
In shells or skeletons that grow by accretion with or
without addition, the different ontogenetic stages are
preserved and can be studied from serial sections of
the hard parts.
Under exceptional conditions ontogenetic stages of
trilobites from the larval stages may be preserved
(see Factsheet 14.2 for more details) as separate
moults. They include:
Protaspis (larval)
Meraspis
Holaspis (adult)
Chapter 1 Introduction 11
The reason for this use of different sets of
terms lies in the difference in the mode of growth
of many ‘invertebrate’ animals from that of
trilobites.
Broadly speaking, the growth of hard parts in
‘invertebrate’ animals are as shown in
Factsheet 1.9.
At each stage of accretionary growth, as the
body grows further, new shell material is added
along the boundary of the existing hard part. Thus,
normally, this break in growth of hard parts is
marked by a groove along the margin of the
previous stage. It is referred as the growth line. In
this type of growth, the shape and outline of the
existing hard part is essentially maintained and,
hence, the growth lines are often described as
parallel. Brachiopod and mollusc shells are largely
marked by this type of growth.
However, as in cephalopods, a very few
gastropods and in corals in addition to the main
shell or skeleton (in corals) growing by accretion,
newer hard parts are added. Septa of cephalopods
or the very few gastropods, and the internal features
within the corallite of a coral (viz. septa, tabulae,
axial structure and dissepiments among the major
ones) are evidences of such growth by addition. In
echinoids (or rather echinoderms), where the hard
part is made of innumerable smaller plates, it grows
FACTSHEET 1.9
Type of Growth in ‘Invertebrates’
l Accretion: Hard part grows through accretion
about the immediately preceding stage. For
example, bivalve or brachiopod shells.
l Addition: New hard parts are added to the
older shell or skeleton. For example, septa are
added to a cephalopod shell or phragmocone
which itself grows by accretion; echinoid tests
grow by addition of newer plates and accretion
of older ones.
l Moulting/ecdysis: Hard part of each stage,
inadequate for the growing body is shed apart
and a new hard part develops on the body grown
larger. For example, trilobites.
12 Part One: Principles

by both accretion and addition. As the body grows
and the test becomes inadequate for it in size, newer,
that is more plates are added at different points. At
the same time, existing plates grow to some extent
by accretion along their border, thereby
compensating the total increase in volume,
necessary for the growing tests.
In ecdysis, hard part of each stage, inadequate
for the growing body is shed apart and a new hard
part develops on the body grown larger, as it is in
trilobites, or, for that matter, in arthropods. Thus,
in between the shedding of older moult and
formation of newer one, the animal exists without
any hard part, that is, unprotected and vulnerable
to attack. It may count on their preservation.
In all cases of growth by accretion, the hard
part of an individual will bear in one single
specimen (read fossil) all the ontogenetic stages.
On the contrary, growth by moulting/ecdysis
leaves a number of moults or their fosssils for one
single individual. Unless it is recognized properly,
those different fossils of one and the same
individual may be confused as belonging to
different individuals or even different species.
We come to the end of this introductory chapter
with Factsheet 1.10, which helps in having an idea
of how ancient fossils are. To be succinct, fossils,
at least life is virtually as old as the earth itself is.
Details of the different aspects mentioned here
are incorporated in later chapters.

FACTSHEET 1.10
How Ancient Fossils Are? (modified after Lehmann and Hillmer 1980)

Geochronologic Million Stratigraphic Chemical Prokaryotes Stromatolites Eukaryotes Metazoa

unit years unit and locality fossils

Cambrian (Base) 570 y y y y y

P Late Ediacara y y y y y
Vendian (Australia)
R Early 680
E Late Bitter Springs y y y y
(Australia)
C Riphean
A Middle Belt Supergr y y y y
(USA)
M Riphean
B Early 1600 Botswana (SW y y y y
Africa)
R 2000 Gunflint Fmn y y y y
(Canada)
I Pre-Riphean Witwatersrand y y y
(S Africa)
A --------------- 2500
N Archean Fig Tree Fmn y y
(S Africa )
3400 Onverwacht y y
(S Africa)
NB: Cambrian/Precambrian boundary at 570 million years; within Precambrian, Proterozoic/Archean boundary at 2500
million years
y-signifies presence
For more details see Chapter 6
 2
2.1 Introducing Taphonomy
Past decades have seen immense changes in
palaeontology. Volumes of facts and findings have
accumulated; understanding of different aspects of
the ancient world has been enriched; but what has
been added, in particular, is the appraisal of how
the record of that organic world is preserved in the
earth and how the history of preservation stands
important in palaeontology, rather geology. It is
now appreciated more and more that palaeontology
misses a lot of information, if it does not consider
how the organism could become a fossil and how
the latter could continue to exist till discovery and
may be, collection. The processes involved are
known as fossilization and preservation. While
the topic was a part of the ‘Introduction’ chapter
in textbooks of the middle of the last century, it
has now attained the status of a developing
important branch of palaeontology, viz.
Taphonomy. Though Efremov introduced the term
in 1940 and Lawrence (1968) widened its scope,
this branch of palaeontology was revived from
oblivion only some two or three decades back.
Before entering into the topic, we may refer a
case which shows how fossils may tell a story that
may be apparent, but not true. It is taphonomy
which helps bring out the real scene.
13
Taphonomy
There is a Pleistocene cave deposit in South
Africa. There an underground cave complex acted
as a catch basin for creatures which fell in it or
whose remains were washed into (it is a
‘concentration trap’ type of deposit).
The fauna included many Australopithecus
(a primate, precursor of man; discussed in details
later) remains, of which one partial juvenile
cranium fossil had two holes about 6 mm in
diameter, 33 mm apart. These were earlier
interpreted as mark of blows from a pointed
weapon, suggesting even tool-using and tool-
making habit of the genus and cannibalism.
But similar holes were found in skulls of
baboons killed by leopards preserved in the same
cave. The holes also matched in diameter with the
lower canines of a leopard jaw.
Hence, it is now interpreted as some activities
of leopards. It also means that the animals including
juvenile Australopithecus did not fell in, but were
either thrown or brought into.
Thus, taphonomy is the branch of
palaeontology (refer also Factsheet 2.1), which
studies how a fossil came into being, more
succinctly, the organism-fossil pathway from the
death of an organism to its being discovered and
collected. The entire pathway has additional stages:
death succeeded by burial in a medium
(generally sediment), diagenesis playing its role
14 Part One: Principles

on the buried material (when the sediment is
transformed into rock) tectonics and/or weathering
affecting the fossil, discovery and collection of the
fossil if it is not totally weathered or eroded out or
deformed beyond recognition or situated in yet
inaccessible localities.
So put in a different way, a fossil owes its
existence to a break in the natural cycle
(Ziegler 1983). An organism with its body,
including its soft and/or hard parts, and the vital
processes of its life, works as an integrated whole
in close interaction with its environment. As soon
as the organism dies, its fight to maintain the
integrity of its body and life stops. In lieu of the
biological processes such as growth, reproduction,
feeding, locomotion (in animals) and such others,
controlled by biological laws, the remains of the
organism is then acted upon by other processes,
viz. various physico-chemical-biological processes
of the atmosphere-lithosphere-biosphere subject to
laws of sedimentation, diagenesis, weathering, etc.
Life processes are stopped, soft parts decay, hard
parts disaggregate and disintegrate. Fossilization
processes disrupt these changes at some stage and
maintain the status quo at that stage as long as the
enclosing medium, commonly sediment, remains

FACTSHEET 2.1
Taphonomy : At a Glance

I. Taphonomy defined
Taphonomy comprises the history of fossils encompassing how the concerned organism was related in
its living to the site where it is preserved, what were its death conditions, what were the pre-burial
chemistry and sedimentology of the organism and its fossilization environment, what were the burial
conditions and processes, nature of necrolysis and diagenetic chemical sedimentology of the dead
remains. It is thus correlated to environmental setting (living and burial) and ambient sedimentary
conditions.
II. Fossilization process involves three stages:
1. Mortality or death converting once-living organisms to non-living materials
2. Burial (biostratinomy) placing the remains/traces in the lithosphere.
3. Diagenesis converting biosphere materials to lithosphere materials.
Taphonomy covers all the three.
III. Knowledge of taphonomy separates a palaeontologist from a biologist
l All organisms are made up of a variety of parts and materials.

l Materials vary in their ability to withstand physical or chemical changes, viz. degradation,
dissolution, breakage, etc. and so to stand preservation.
l Parts also vary in their ability to withstand disaggregation, dissociation, compaction and
entrainment, and transport by hydraulic or wind action and so also to stand preservation.
l The eventual mode of preservation depends further on the nature of the sediment in which the
organism is entombed, its softness to take impressions, its permeability and cementation potential.
l The preservation potential of the sediment, and the basinal and tectonic histories of the site, are
also important.
l Thus, there are the intrinsic and the extrinsic factors.

l The former relate to the organism, the latter to the sedimentary environment and to its geological
history.
l It is this understanding and appreciation of taphonomy that separates a palaeontologist from a
biologist.
 Chapter 2 Taphonomy 15

intact; they thus prevent any further destruction. A
substance, an organism that was once a part of the
biosphere is thus transferred to the lithosphere. All
this is relevant, even to beginners, to appreciate
one basic point. To understand and know a fossil
well, one must have an adequate knowledge of its
biology and taphonomic history as well. It also
draws attention to another basic tenet of science
that identifies mutually interactive relationship of
different material processes and things; in this case
between the biosphere and the lithosphere
(see Factsheet 2.2).
Let us discuss a few concrete cases. In the several
thousand metre thick succession of the Gondwana
Supergroup in India, we have only one ‘plant’ fossil,
viz. Williamsonia sewardiana; the rest of the rich flora
consists of leaf, stem or other separate parts of trees
or plants. Even the separate leaves, stems, etc. are

FACTSHEET 2.2
Fossilization Preservation: Processes and Conditions

Fossilization preservation i.e. organism fossil pathway is removal of an object (organism) from biosphere to
lithosphere (as fossil).
Fossils to form and be preserved require two basic conditions:
1. Body of an organism (animal or plant) contains soft tissues with or without hard mineralized (in animals)
or cuticular (plants) parts. Presence of preservable durable parts or traces is the first basic condition of
fossilization. Here the controlling factor is primarily pre–mortem, biological, i.e. as it was before the death
of the organism.
2. Cut off from destructive agencies or processes. Here controlling factor is primarily post-mortem,
taphonomic, i.e. as it happened after death during fossilization-preservation.
In fact, it is required that organic remains or traces being fossilized are protected from mechanical
pressures or impact, from chemical reactors including oxidation, hydration, etc. from bacterial
decomposition and from scavenging action or predation by other organisms.
Specific conditions of fossilization-preservation may be listed as follows:
1. Mineralized (and/or organic) skeletal parts or covers to add to durability. Here control is primarily
biological as different organisms have different minerals in hard parts.
(a) Chemically stable mineral like calcite is more durable than aragonite or hydrous silica.
(b) Phosphatic minerals generally more stable than calcareous minerals.
(c) Stability also depends on how strong or weak, robust or delicate, the skeleton is.
2. Burial or entrapment cuts off the material to be fossilized from air and/or water and bacteria, often also
from scavengers, deposit feeders or predators. Here control is primarily non-biological.
(a) Sediments form the most common medium, hence preservation more likely in areas of sedimentation,
viz. shallow sea, lakes, etc. and less so in areas of erosion. For example, on land, particularly
mountains, etc.
(b) Other media include resin secreted by trees, ice, natural asphalt or pitch.
3. Areas (basins) of low energy cause less mechanical breakage: in this case control is post-mortem,
biostratinomic.
4. Areas of rapid sedimentation ensures quick burial: control post-mortem, biostratinomic.
5. Fine sediment cover gives way to impermeable rock as host environment helping less action of diagenetic
fluid: control post-mortem, diagenetic.
6. Early cementation of host sediments may cause less compaction: diagenetic control.
7. Weathering and tectonic deformation may bring about damage; partial or complete.
8. Accessibility as a secondary factor may leave fossils of yet inaccessible areas still to be discovered.
16 Part One: Principles

preserved in most cases, not as actual remains of the
plant concerned, but as impressions or compressions,
carbonized or not, and rarely as silicified remains. It
is always difficult to ascertain which leaf fits with
which stem or what is the plant organism to which a
leaf, a stem or any other fossil may belong. Only in
the case of Williamsonia sewardiana, we know that
its leaf is Ptilophyllum,stem Bucklandia and cone
Williamsonia, all found together in one case.
Obviously, a pertinent question to ask would be:
why is it so that virtually all the Gondwana plants are
preserved as separate parts and that too not as their
actual remains? The answer is to be looked in
taphonomy.
Our second example pertains to the Gondwana
vertebrate fossils of India. Six major taphonomic
types of skeletal assemblages of Gondwana
vertebrates, viz. reptiles, amphibians and fishes have
been identified (Bandyopadhyay, Roy Chowdhury
and Sengupta 2002) (see Factsheet 2.3). They
include complete articulated or semi-articulated
skeletons disarticulated skeletons of a large number
of individuals of the same species; disassociated
bones of individuals of the same species along with
large tree trunks; disarticulated, yet not dispersed
bone assemblage of mixed individuals; isolated
remains of single individuals and isolated
fragments. Each of these types can be successfully
correlated to this or that type of environmental
condition in which they were fossilized, highlighting
the fact that these land vertebrates that lived in the
Gondwana time did not have their body fossilized
in the same manner. Again, taphonomy answers the
question: why?

FACTSHEET 2.3
Taphonomy of Vertebrate Skeletal Assemblages from Indian Gondwanas
(Based on Bandyopadhyay, Roy Chowdhury and Sengupta 2002)

Type A: Complete articulated or semi-articulated skeleton/ Triassic / Panchet, Yerrapalli and Maleri Formations
of DVB and PGVB: Life-like posture of autochthonous remains: quick burial: trapped-bogged down
in thick mud of flood plains with rapid and heavy sedimentation.
Type B: Disarticulated skeletons of many individuals of the same species / Mid Trias Yerrapalli and Up. Trias
Maleri Formation of PGVB: Mass-mortality in a flood-like catastrophe.
Type C: Disassociated bones of individuals of one species in a ‘log-jam’ condition/ Lr. Jurassic Kota Formation
of PGVB: Mass-mortality in a flood-like catastrophe.
Type D: Disarticulated bones of mixed individuals/Mid Trias Yerrapalli and Denwa Formations of PGVB and
SB: Small juvenile reptiles trapped by fast sedimentation and preserved near the site of death.
Type E: Isolated bones of single individuals/in all horizons of all basins: Disarticulated lighter bones-brought
by thin sheets of flowing water to a flood plain covered up by not-too-fast, yet not-too-slow
sedimentation, allowing scavenger action and compaction.
Type F: Isolated fragmentary bones in all horizons of all basins: Long post-mortem pre-burial history, i.e.
transported carcasses ‘de-fleshed’ and disarticulated subaerially by scavengers or in other ways and
further transported to be buried in channel sediments
NB: 1. Damodar Valley Basin (DVB), Satpura Basin (SB), Pranhita-Godavari Valley Basin (PGVB)
2. Ingested bones of small Malerisaurus robinsonae found inside the rib cage of two individuals of phytosaur reptile
Parasuchus hislopi
3. Wadiasaurus indicus, a common lowland dicynodont reptile in Yerrapalli Formation (Type B) has skull and maxillae
tuskless. But from a dozen other localities with one or two individuals represented, W.indicus shows tusked maxillae,
quite robust nasals and maxillae. The former may be females and juveniles living in herds, latter isolated males
(Bandyopadhyay 1999).
 Chapter 2 Taphonomy 17

One answer to the above questions is: land
plants and vertebrates are relatively larger organisms
that have many skeletal elements in their body;
hence, they are easily disarticulated during
fossilization and behave differently during
fossilization (see Factsheet 2.4). Even smaller
invertebrates, more so the microfossils among them
that stand far greater chance of preservation with
the whole body (skeleton normally) intact, present
interesting taphonomic characteristics and effects.
In Kutch (Kachchh is a more recent variant of
spelling and used from here onwards), Tertiary
rocks (or elsewhere too), fossiliferous limestones
often have prolific foraminifera (one-celled

FACTSHEET 2.4
Plant-Vertebrate Taphonomy Contrasted

They stand alike on:

Both land plants and vertebrates are generally relatively large bodied organisms, with numerous parts in the
body, whether hard or soft.
Individual fossils are usually one or the other part of the body (e.g., leaf, stem, bone, jaw, teeth).
These organisms lived on land and hence their remains were needed to be transported to nearby basins (lakes,
rivers channels or flood plains, etc.), where they may be buried under sediments for fossilization.
During this process, their body, including the hard parts in them, are commonly disarticulated (parts separated
from each other), disaggregated (separated parts are strewn apart) and even disfigured.
They differ on:
Parts of extinct plants often cannot be readily and definitely assigned to the actual organism. This is because
many of the major groups of ancient plants are no longer living. Thus, a leaf is identified, for example, as
Glossopteris or Schizoneura (in the second case associated with or without the stem) solely depending on their
respective morphology without being sure of its botanical affinity. They are then regarded as form genera.
Vertebrate anatomy is better comprehended and so the parts may be correlated and assigned to precise genera or
species, i.e. the animal itself. Their zoological affinity thus ascertained, such parts are not form genera.
Plants are generally preserved as impression, compression, anatomical preservation or a combination of them
(Goldring 1999). Impression is left on the enclosing sediment surface without any original organic material, and
by a thin film of carbon or, say, iron oxide.
Anatomical preservation has cellular structure in three dimensions preserved in part or in full. Cells may be
preserved in the form of charcoal (partially burnt rapidly in oxygen-deficient condition by say lightning strike,
wild fire, volcanic heat, etc.), calcified, silicified, pyritized, etc. with cell walls having been replaced (petrifaction)
or intracellular voids filled in (permineralization). Both petrifaction and permineralization require rapid burial
and no decay to preserve the cell structures.
In vertebrate fossils, generally the skeletal material and the anatomy are both preserved; the former may be
mechanically affected or chemically replaced.
Plant roots, under suitable conditions, or trunks may be autochthonous and preserved in situ. Vertebrate fossils
may or may not be autochthonous, depending on habit and size of the animals.
NB: A few commonly used terms in regard to plant preservation are impression, compression (type of preservation in
which plant remain is represented by the negative imprint); cast (type of preservation that forms in a mould); mould
(type of preservation in which organic material is represented by a distinctly and strongly three-dimensional negative
imprint); petrifaction/ permineralization (type of preservation in which extraneous mineral material has permeated
the cells and interstices soon after deposition). Authors are, however, often at variance in defining the terms. Here
Taylor (1981) and Diessel (1992) are followed.
18 Part One: Principles

organism) fossils. The latter organisms lived on the parallel to the bedding; they rather showed different
seafloor and had thin disc-like tests (hard skeletal sets of orientation and even sorting on size of
part). Yet they were preserved not necessarily all individuals of the same species (Figure 2.1).

(a) (b)

(c) (d)

(e) (f)

(g) (h) (i)

Fig. 2.1 Bioclastic grain fabric representing depositional-cum-burial conditions from (a) – (d); Varied effects
of diagenesis (micritization) on foraminiferal tests depending on initial morphology and nature of
diagenesis from (e) – (i).
(a) Rock sample of a foraminiferal limestone, (b) and (c) Thin section of two samples of foraminiferal
limestone, (d) Sketch of the fabric drawn from another thin section, (e) Partially micritized test of Assilina;
solid lines show actually observed parts; dashed ones original parts, now lost, but reconstructed from
symmetry, etc., (f) Original parallel-sided test of the same genus with semi-involute spiral lamina;
reconstructed from the observed parts in (e), (g) Same such test reduced to a biconcave one with apparently
evolute spiral laminae; black portion shows test material lost (along the margin) or added (inside chambers)
during diagenesis; thus, the biconcave test, characteristic of a species of Assilina does not represent the
actual form; it is an effect of diagenesis on a test of a completely different morphology pertaining to a
different species, (h) and (i) A Discocyclina test, partially drawn; (h) and a Nummulites test (i); neither
will lose its diagnostic characters even if the portions beyond the dashed lines are removed during
diagenesis, pointing to the role of initial morphology.
 Chapter 2 Taphonomy 19

It meant, they were either moved from their life
position, deposited in different orientations as and
when the current varied and finally buried in such
positions, or they were disturbed from their buried
positions during diagenesis of the host carbonates,
an interesting point to settle from other evidences.
2.2 Coming into Being of Fossils:
Organism of Biosphere Turn
into Fossil in Lithosphere
What exactly happens during fossilization and
preservation? Taphonomic processes produce three
basic types of fossils: body fossils, trace fossils
and chemical fossils. (See Factsheet 2.5 for
definitions and Figure 2.2.) Moulds, casts,
impressions or such other evidences of organisms
(see Part B of Factsheet 2.5) made through
abiogenic chemical or physical interactions
between the organic remains and sediments around
them, are not results of processes during the
lifetime of organisms and, hence, are not trace
fossils. They are also not strictly part or whole of
the body. Yet, so long as from them one obtains
some idea about form and/or structure of the body
of the organism, they are regarded with the body
fossils themselves. Trace fossils, otherwise also
called lebenspurren or ichnofossils, are often
classified as behavioural/functional, nutritional,

FACTSHEET 2.5
Taphonomic Types of Fossils
(Based on Babin 1980, Frey and Pemberton 1985, Nield and Tucker 1985, Clarkson 1998)

A. TAPHONOMIC TYPE
BODY FOSSILS
...in which whole or parts of
the body, including hard
and/or soft parts are pre-
served, or from which some
idea about their form and/
or structure may be made.
Unaltered Non-oxidizing decay preventing condition and/or
soft parts. medium. For example, ice, bogs, wax, tar, resin, silica.
Unaltered Calcareous: calcitic/aragonitic Oganic: chitinous/scleroprotein,
hard parts phosphatic, siliceous, etc. Soft parts oxidized or decomposed.
Altered Soft parts lost; hard part alteration by changes isochemical
hard parts Recrystallization, e.g. aragonite-calcite by changes involving:
l Addition, e.g. permineralization
l Removal and addition, e.g. replacement
l Removal, e.g. distillation, compression

TRACE FOSSILS Behavioural Resting traces Grazing traces

...which preserve traces of /functional Dwelling traces Crawling traces
activities made during life Nutritional Escape traces Feeding traces
time of the organism. Reproductive
Associational

CHEMICAL FOSSILS Palaeoproteins

...which preserve chemical Palaeoamino acids
compounds that were part Palaleosugars
or constituent of the Lipids
body during lifetime. Bacteria

TYPES VARIATIONS PROCESSES

(Cont...)
20 Part One: Principles

FACTSHEET 2.5 (Cont...)

Taphonomic Types of Fossils
(Based on Babin 1980, Frey and Pemberton 1985, Nield and Tucker 1985, Clarkson 1998)

B. TAPHONOMICALLY CONTROLLED FEATURES

SEDIMENT-BIOTA interaction and products
ABIOGENIC passive interaction
Remains of dead organism interact passively with the enclosing sediments.
Sediments around impressed upon
l by internal/external surfaces of hard parts

For example, carbonized impression (plant, graptolites), compression (do), impression (any kind of organism)
l by soft tissues

For example, death marks (jellyfish)

Sediment filling bound
l by internal surface, e.g. internal mould (gastropods, cephalopods)

l by both internal and external surfaces, e.g. cast (bivalves)

l by external surface, e.g. external mould (bivalves)

BIOGENIC active interaction

Live organisms react actively with sediments, in the process leaving (for traces)/not leaving (for non-traces)
indication of functional anatomy or habit
I. Biosedimentary Structures
Bioturbation Biostratification Biodeposition
Disrupts bedding, etc. builds bedding, etc. produces material for;
For example, traces like resting For example, non-traces like For example, traces like
marks, burrows, trails, tracks, footprints stromatolites, biogenic graded beds faecal pellets, coprolites
II. Bioerosion
Excavates solid structures, including bedding in rocks.
For example, borings, gnawings, scrapings, bitings
III. Other Evidences
For example, eggs (also considered body fossils); tools

reproductive and associational; the first includes
more known traces of burrows, which may be of
different kinds, viz. resting, grazing, dwelling,
crawling, escape and feeding traces. Trails
(for crawling organisms, e.g. trilobites), tracks
(for instance of birds) and footprints (dinosaur or
human) are other examples of behavioural traces.
Grasses in the mouth of Pleistocene Siberian
mammoths or undigested food material in dinosaur
coprolites are among the nutritional traces; fossil
eggs of dinosaurs or other reptiles are reproductive
traces. There is, however, a different opinion on
fossil eggs, that tend to consider them as body
fossils of the earliest stage of life.
Chemical fossils are rarer, though
paradoxically, not the youngest (see Factsheet 2.6).
In fact, delicate chemical compounds stood better
chances of preservation in sediments deposited in
the non-oxidizing or oxygen-depleted primitive
atmosphere.
Body fossils are by far the most frequent. To
become a body fossil, the body of an organism
(animal or plant) undergoes a number of
overlapping, yet consecutive processes
(Factsheet 2.7). It contains soft tissues or fleshy parts
which are mostly made of organic compounds such
as proteins, fats and carbohydrates with or without
hard mineralized (in animals) or cuticular (plants)
 Chapter 2 Taphonomy 21

(d)
(c)

(a) (b)
(f) (g)
(e)

(j)
(i) (k)

(h)

(l)

(m) (n)
(o) (p)

Fig. 2.2 Types of fossils.

Trace fossils (a) Burrow, (b) Burrow, (c) Exposure of a possible crab burrow from Oligocene of Kachchh,
(d) Shark teeth, (e) Juvenile echinoid, (f) Juvenile bivalve, (g) Alveolina, a larger foraminifera (microfossil
s.s.), (h) Echinoid spine (microfossil s.l.), (i) Handful of a river sand from Kachchh showing foraminiferal
tests of different size, (j) Half of a Nummulites test, a larger foraminifera (microfossil s.s.), (k) Fossil of a
bone (vertebra), (l) A single molar tooth of a mammal (pig), (m) Dinosaur bones (marked X) exposed in a
section (note the size as compared to the adult human figure, marked Y and to the microfossils), (n) Internal
mould of a cephalopod with suture, (o) External mould of a bivalve, (p) Internal mould of a gastropod.
22 Part One: Principles

FACTSHEET 2.6 decomposition or decay of soft tissues with or

Chemical Fossils: Aminostratigraphy
An interesting recently developed use of chemical
fossils may be found in Aminostratigraphy which
deals with distribution of fossils with similar amino
acid characteristics and organization of strata into
units on the basis of the fossils they contain. Thus,
an aminozone is a regional aminostratigraphic unit
used for relative age dating and correlation.
Authors (Bowen 2000) even suggest that for
Pleistocene or similar successions where terrestrial
deposits show frag-mented record and there are
insufficient paleon-tological first-appearance and
extinction events because of the relatively short time
duration, aminostratigraphy may help produce more
refined and efficient tools.
substances, which house these softer parts. The
mineral matters may be calcareous, calcium
phosphatic, siliceous or of other chemical
composition (Factsheets 2.8, 2.9). After death
overcomes the organism, there is a gradual
without hard parts being broken, disarticulated or
separated from the body (just as bivalve shells fall
apart or bones of a vertebrate skeleton are
separated). Soft tissues are destroyed, i.e.
decomposed or decayed, (the process referred as
necrosis) in different ways under different
conditions. Changes are basically chemical,
affected through bacterial or chemical causes.
Thus, tissues may be affected by either aerobic
bacteria in the presence of water and oxygen or
by anaerobic bacteria where there is no oxygen.
In the former case, soft tissues react with oxygen
and water to break down into carbon dioxide, water
and other gaseous substances that are released to
the surroundings. In the absence of oxygen,
anaerobic bacteria use up the tissues of the host
organism in their own metabolism, thereby causing
those tissues to break down into hydrocarbons. The
latter may ultimately, under increased suitable
temperature and pressure, change into hydrocarbon
fuels, viz. bitumen oil; natural gas, methane, etc.

FACTSHEET 2.7
Stages and Processes in Formation of Body Fossils
Any body fossil has undergone a series of consecutive yet overlapping processes
l Death or entrapment of an organism.

l Decay or decomposition of soft parts + separation of hard parts.

l Burial or biostratinomic processes including preburial sedimentation history with sorting, attrition, reworking,
etc.
l Final incorporation into the sediment/other media.

l Diagenesis including compaction and chemical changes.

l Tectonic deformation and/or weathering.

l Discovery with/without collection.

In anaerobic condition, i.e. in the absence of oxygen, they are acted upon by anaerobic bacteria.
l Products are hydrocarbon of high molecular weight plus materials used in metabolism of anaerobic bacteria;
closed, stagnant condition;
l Soft parts lost leaving behind bituminous mud, even hydrocarbon fuels or bituminous under suitable P-T
conditions.
In aerobic condition in the presence of oxygen and water, acted upon by aerobic bacteria, products are:
l CO , H O or other gaseous compounds that volatilize;
2 2
l Condition open, aerated;
l Soft parts are lost without residue.
 Chapter 2 Taphonomy 23

FACTSHEET 2.8
Relative Importance of Organic Groups as Fossils with Hard Parts

As fossils Common A few Uncertain Unknown

Hard parts
Common Rhizopoda Porifera
Cnidaria Mollusca
Arthropoda Tentaculata
Echinodermata Vertebrata
In some Ciliata
Annelida
Flagellata
Branchiotremata
Absent Tunicata Mesozoa
Nemertini Sporozoa
Sipunculida Ctenophora
Onychopora Protociliata
Chaetognatha +8 groups
Nemathelminthes

FACTSHEET 2.9
Hard Part Constituent in Major Organic Groups
(Based on Tasch 1973, Leeder 1983, Black 1988, Clarkson 1998, Goldring 1999)

Hard Part Siliceous Calcium Calcareous Organic

Composition Opaline Phospatic Calcitic Aragonitic
(Vateritic)
Organic group
Porifera G/H M G Spongin
Archaeocyatha D/H
Cnidaria M G G Chitin/ protein
Bryozoa D M M Chitin
Brachiopoda Mil G Gim/Gil Chitin/
Protein
Mollusca
Bivalvia Gil Gil (M) Conchiolin
Gastropoda Gil Dil Mil
Cephalopoda Dil Mim/Mil
Arthropoda D/H Gil Chitin
Trilobita D/L Undefined
Echinodermata D/H M Protein
Annelida G/H G Protein
Hemichordata Dil Protein
Foraminifera G D/H M (M) Tectin
(Cont...)
24 Part One: Principles

FACTSHEET 2.9 (Cont...)

Hard Part Constituent in Major Organic Groups
Based on Tasch 1973, Leeder 1983, Black 1988, Clarkson 1998, Goldring 1999

Hard Part Siliceous Calcium Calcareous Aragonitic Organic

Composition Opaline Phospatic Calcitic (Vateritic) (Variety)

Radiolaria D M Protein
Diatom D
Silicoflagellate D
Dinoflagellate D Cellulose
Coccolithophore D
Acritarch D
Taphonomic Generally and relatively Generally and relatively
stability more stable less stable
Explanation: (M) minor (G) in some groups (D) dominant
(il) interlayered/ (im) intermixed
(L) low Mg calcite (H) high Mg calcite

Preservation of soft tissues is caused by loose or disarticulated. If they are delicate or

stagnation or airtight condition where bacterial
decomposition, oxidation or hydration or such
other changes are prevented. It may be either, in
the absence of water (in aridity) or in the presence
of hygroscopic material, or from freezing or due
to incorporation in decay-inhibiting material such
as resin trapping insects, natural asphalt or pitch
or bogs drowning organisms that fell into, or fine
grained impermeable sediments quickly con-
solidated by early diagenesis as it happens in
lithographic limestone. Since these are not usual
situations, soft parts of organisms are rarely
preserved. Yet those rare instances stand out, as
they provide important information, normally not
available, about ancient organisms, their body,
habit or environment in which they lived.
(see Factsheet 2.10 and Appendix 1.)
Commonly, however, soft parts of the body
are lost by early necrosis. Hard parts made of
mineral matters do not react largely in these
processes. Rather on the decay of soft parts, hard
parts that are held together by soft tissues are let
loosely held, they may be mechanically broken into
pieces, particularly if acted upon by high energy
running water or wind, or by load pressure of any
sort. In some cases, they are preserved without
any alteration in chemical or mineralogical
composition. But in most cases, hard part mineral
matter is partially or totally altered mineralogically
or even chemically. It depends on their chemical-
mineralogical composition and their susceptibility
to change. For example, aragonite of, say gastropod
shells, is more susceptible to alteration than calcite
of which, say echinoid tests are made up. Different
groups of organisms have hard parts of different
types, physical–structural or chemical-
mineralogical. Hence, they vary in durability and
stability.
These chemical changes of hard parts may be
isochemical (aragonitic mineral matter
recrystallizing into calcitic matter) or may involve
addition of material mostly in interstices
(permineralization) or removal, normally of
volatile substances (distillation, compression) or
 Chapter 2 Taphonomy 25

FACTSHEET 2.10
Instances of Extraordinary Preservation and Uncommon Fossil Types

Unaltered Soft Parts:

In ice: Mammoths from Siberia with 40 specimens, 4 intact
In paraffin: Mammoths and rhinos from Galicia
In bogs: Irish elk and humans. Humans in the Tollund Fen, Denmark (both Subrecent)
In asphalt: Mammals of Ukraine pitch and California asphalt (both Pleistocene)
In silica: Protoplasms of protoctists sensitive to cytological staining.
Soft Tissues as Imprints:
Entire soft body: Late Precambrian annelids, Cambrian medusae, Pennsylvanian hydroids
Organ: Liassic/Cretaceous moulds of molluscan intestines; Jurassic Archaeopteryx feather from
Solenhofen Lst.; Wings of insects from Carboniferous shales; Leaves from travertine of
Sezanne
Uncommon Fossils:
Excrement: Selachian coprolites; Tomaculump—excreta of trilobites or other invertebrates as tiny
ellipsoidal pellets in condensed layers or arranged along sinuous surface markings.
Feeding group: Cephalopod shells of Orthoceras around an accumulation of organic debris.
Feeding habit: Jeholornis prima, early Cretaceous bird of the size of a large crow from China (also Spain,
Madagascar) shows its gut full of seeds — the first direct evidence of seed eating in a
bird (Zhou and Zhang 2002); Grasses in the mouth of frozen Siberian mammoths.
Reproductive Traces:
Ordovician organic vesicles (cephalopod eggs); Oligocene birds’ eggs; Cretaceous dinosaur eggs from Gujarat;
Two Early Cretaceous fossils of embryo of pterosaur, Mesozoic flying reptile, one from China and another from
Argentina provide insight into the eggshell morphology, early growth and nesting environments of pterosaurs.
(Also see section 18.10.4.)
Commensalism:
Platyceras (gastropod) around anus on the calices of crinoids; Hicetes (worms) with Pleurodictium (tabulate
corals); Phacops in atrypid shells use empty shell for refuge.
Chemical Fossils:
Collagen fibrils from bones (Miocene: 25 Ma); Amino acids from bone fragments (Devonian: 380Ma); Devonian
conodonts and Jurassic pliosaurs analyzed for amino acids and collagens suggest non-affinity from the difference
in composition and combinations.
Predatory scars of different morphology and size, distributed highly non-random in shells of a recent inarticulate
lingulid brachiopod from two intertidal localities in the northern Gulf of California more towards the anterior
shell edge suggest seasonal predation in the late fall and winter months, 25 per cent specimens bear repair scars.
(Kowalewski, Flessa, and Marcot 1997)
Abundant worm borings largely oriented in some brachiopod shells of Ordovician oil shale in North Estonia;
some hosts appear to have made blister-like shell-repair structures in their interior. (Vinn 2005)
NB: More examples to be found in Appendix 1 on Fossil Lagerstätten.

both removal and addition (hence, the original called pyritization, silicification, etc. In the case
material is replaced by another new). Replacements of total replacement of original hard parts, the fossil
are termed by the product material and are, thus, is preserved as a cast (of, say, a bivalve shell) or a
26 Part One: Principles
carbonized impression (of a plant leaf). In fact,
separation of moulds, casts, etc. from body fossils
become difficult on these grounds.
Even hard parts may be lost during burial or
diagenesis. Delicate skeletons may, thus, be
excluded from high energy or chemically active
environments. Not just larger fossils, even
nanofossils may be lost diagenetically from
the fossil record (Taylor in Lord1982). Thus,
apparently unfossiliferous record may really not
indicate the absence of life during its deposition.
An example is found from Pliocene Coralline
Crag where a limited assemblage of nanofossils
consists of solution-resistant forms which are
also of little stratigraphical interest; any delicate
species has been lost (Hamilton and Hojjatzadah
in Lord 1982).
Taphonomic changes may sometimes affect
systematics and other studies. Thus, diagenetic
overgrowth and dissolution may make species
identification difficult, even for nanofossils
(Taylor in Lord, 1982). An example is found in
the case of Micula staurophora and Quadrum
gartneri, where preservational variation is studied
to check whether the two are different species or
not (Crux in Lord 1982).
Another case may be cited from larger benthic
foraminifera, Assilina, in which a complete series
of gradational changes could be noted. In this
genus, diagenetic micritization and, therefrom,
destruction of spiral laminae gave rise to
morphotypes that resembled different species.
Some of these were Lower Eocene, but
diagenetically least altered form resembled a
Middle Eocene genus. Hence, recognition of these
taphonomic effects modify, biostratigraphic
conclusion too (Figure 2.3; Ray, 1988).
Bioturbation and physical reworking also
cause time averaging (temporal mixing) of different
communities and may lead to increased diversity
and variation in morphological features of
several lineages. Temporal mixing often goes
unrecognized in fossil assemblages. Thus, false
FADs and LADs (First Appearance Datum and Last
Appearance Datum, respectively) may result from
bioturbation and reworking. False LADs are more
serious, because bioturbation and reworking
preferentially mix sediments upward (bringing
older sediments up with younger ones; false LAD
becomes extended one).
Part B of Factsheet 2.5 mentions some
taphonomically controlled associated features
formed by sediment–biota interaction. Those
formed by abiogenic processes of passive
interaction between remains of dead organisms and
enclosing sediments, viz. moulds, casts, etc. have
already been discussed. Biogenic processes, in turn,
are those in which live organisms react actively with
sediments, in the process leaving
(for traces)/not leaving (for non-traces) indication
of functional anatomy or habit. Their products
include biosedimentary structures, bioerosion
structures and others. The first includes bioturbation
that disrupts bedding, etc. Examples are to be found
in traces such as resting marks, burrows, trails,
tracks and footprints that speak about different
habits of the concerned organisms, how they made
burrows or how did they move about and so on.
They may also suggest some anatomical
information, for whether they crawled without feet
or what kind of feet did they, have, etc.
Biostratification builds bedding, for example, non-
traces such as stromatolites, biogenic graded beds.
Biodeposition produces materials which are
incorporated in the enclosing sediments such as
traces, e.g., faecal pellets, coprolites. These may
turn out to be helpful in finding out the feeding
habit of the organisms. Bioerosion excavates solid
structures, including bedding in rocks; for example,
borings suggesting its habit, or gnawings, scrapings
or bitings that may indicate the presence of the
necessary organs or appendages. Other evidences
include eggs (also considered body fossils) or tools,
latter particularly in higher forms of animals such
as primates. Most of these biogenic features are
different kinds of trace fossils in a broader sense
of the term. Exception includes stromatolites which
are biosedimentary structures formed by the

interaction of cyanophyte (popularly called blue-
green algae) mats and their surroundings. These
organisms secrete calcareous substance on them
and at the same time trap fine suspended carbonate
particles from the water above. This total bulk of
calcium carbonate form the laminae, which grow
successively as cyanophyte mats on each newly
formed lamina and repeat the same actions and
reactions. The vertically grown columns of
laminae make a stromatolite, a biosedimentary
structure and not a fossil itself. It may contain a
cyanophyte fossil, in the form of thread or bleb or
so, if it happens to be preserved in laminated
growth.
The importance of taphonomy is further
brought out from what are known as fossil
FACTSHEET 2.11
Types of Fossil Lagerstätten
Concentration Deposits
Exceptional number of body remains,
mainly hard parts, entire or broken
accumulate in a deposit as in coquinas,
bone beds, cave deposits, natural traps.
Types of Conservation Deposits
Obrution Stagnation
deposits deposits
Obrution deposits
Cause and environment of preservation
Episodic smothering, i.e. suffocating
death and rapid burial
Communities mainly affected
Mainly benthic communities
2.3 Fossil Lagerstätten
Recent increased interest in taphonomy has led to
thorough and extensive studies of the different
fossil Lagerstätten of the geological column
(Factsheet 2.12). Many of them are known long
since; some are more recently explored or known.
But in all cases there are volumes of fresh vital
information Factsheets A1.1 to A1.15 provided in
Chapter 2 Taphonomy 27
Lagerstätten or taphonomic windows. It will be
clear from the above discussions that fossilization–
preservation depends on a host of factors. Many
information are likely to be lost if these factors do
not work out favourably. Thus, fossil record is
normally far from complete. It has already been
stated above that soft parts are rarely preserved,
though if and when preserved, they provide much
information and insight into many normally
unknown facts and relations among organisms or
between organisms and their environment. Such
records are called fossil Lagerstätten or taphonomic
windows that act as windows or bridges in the fossil
record to fill the gap in reconstruction of the earth’s
history. They may be of a few kinds as shown in
Factsheet 2.11.
Conservation Deposits
Exceptional preservation including
that of soft tissues that provide much
normally not available information.
Conservation Concretions
traps
Stagnation Deposits
Reduced decay due to anoxic conditions
in stagnant or hypersaline bottom waters
Majorily pelagic communities
Appendix 1 present some details of a number of
major fossil Lagerstätten of the geological
column. In the present chapter, Factsheets 2.13
(a, b, c) provide important characteristics of them.
Elaborate discussions are available in, for
instance, Selden and Nudds (2004). A through-
time overview of the organic world in some newer
light may be available from these summary
information.
28 Part One: Principles

FACTSHEET 2.12
Stratigraphical Position of Important Fossil Lagerstätten

Era Million years Period/Epoch Fossil Lagerstätten Location

before present

---- Holocene
Pleistocene RANCHO LA BREA 15 Los Angeles, USA
2.5
Pliocene
Cenozoic Miocene BALTIC AMBER 14 Samland, Russia
23.5 Oligocene
Eocene GRUBE MESSEL 13 Frankfurt, Germany
Palaeocene
-----65.0
Cretaceous SANTANA AND CRATO 12 Santana, Brazil
146.0
SOLNHOFEN LST.11 S. Bavaria, Germany

Jurassic MORRISON FMN. 10 Utah-Colorado, USA

Mesozoic HOLZMADEN SHALE9 Stuttgart, Germany
205.0 Triassic GRÉS À VOLTZIA8 Strasbourg, France
-----230
251 Permian
Pennsylvanian MAZON CREEK6-7 Illinois, USA
320 Mississippian Hot Spring Dep. and Shrimp Scotland
353
Palaeozoic Devonian HUNSRÜCK SLATE 5 Koblenz, Germany
RHYNIE CHERT 4 Aberdeenshire,
Scotland
409
Silurian
439 Ordovician SOOM SHALE 3 Cape Town, S. Africa
510
Cambrian BURGESS SHALE2 British Columbia,
USA Chengjiang,
China
----- 540
Precambrian - 4600 Late EDIACARA 1 Australia
NB: Numbers against each occurrence refer to corresponding factsheet of the series A1.1–A1.5

2.4 To Read Out the
Taphonomic History
Finally, a few words need be added on how to read
out the taphonomic history of a fossil or a fossil
assemblage. Obviously, in keeping with the essence
and coverage of the subject, one requires both
biological and geological knowledge to arrive at
the answer. The former includes being conversant
with the different kinds of morphology, structure
and composition of the body and skeleton or hard
parts found in different groups of organisms,
 Chapter 2 Taphonomy 29

basically biological knowledge, but at the same
time required to ascertain how far they are
potential for preservation, i.e., how durable and
stable they are physically and chemically–
mineralogically. This demands idea about stability
fields of different constituent minerals of hard
parts in respect of Eh (oxidation–reduction
potential: negative value suggesting reducing
condition)– pH (alkalinity–acidity; higher value
suggesting alkalinity)–salinity (36 per cent being
the normal sea water value) and other conditions,
particularly of the area of fossilization. At the same
time, one must be aware of the living condition of
the organisms concerned, including energy
condition, sedimentational characteristics, such as
rate of deposition, grain size of sediments, porosity
and particularly permeability of the sediments,
which in turn will affect lithification–diagenetic
processes prevailing in that part of the basin. The
latter ultimately determines the preservation type
and state of the fossils. All these are concerned
with the geological insight with which a
palaeontologist looks at his or her fossils. The
methods of study will naturally be determined by
these questions and objects involved. In addition
to the traditional descriptive morphological studies
and taxonomy-systematics, qualitative-quantitative
chemical analysis, optical or even finer resolution
microscope studies for petrography–mineralogy,
and others may be taken help of depending on the
depth and span of the work.
In summary, taphonomy will take a
palaeontologist to conclude how the fossils in his
or her collection have come to what they are, as
well as about a host of information on the original
organisms and their life and fossilization
environments. This leads us to initiate some
discussions on organisms and their environments.

FACTSHEET 2.13
Summary of Characteristics of Fossil Lagerstätten Listed in Factsheet 2.12 :
(a) Upto Palaeozoic
1. LAGERST- EDIACARA BURGESS SOOM RHYNIE HUNSRUCK MAZON
ÄTTEN SHALE SHALE CHERT SLATE CREEK
2. Setting Marine Marine warm Marine shallow Hot Spring Marine photic Marine+
photic zone cold water on land zone terrestrial/
freshwater

3. Derivation Autochthonous Autochthonous Allochthonous Allochthonous Both autoch-and Both autoch- and
allochthonous allochthonous

4. Habit of Mainly benthos Mainly benthos Nektons Land plants Benthos, nektons, Varied
Organisms in, or above dominate and animals trace fossils
seabed benthos

5. Preservation Soft part Soft part No soft Soft part Soft part Soft part
Quality preserved preserved part preserved preserved

6. Particularity Pre-hard part Nearly all animal Post-Phane- Early Fauna intermediate From swamp
organic world phyla represented rozoic glaciation terrestrial between Burgess forest, upland to
organic world biota with Shale and Mazon setting fresh, water
diverse plants Creek faunas brackish and
and animals deltaic
environments
(Cont...)
30 Part One: Principles

FACTSHEET 2. 13 (Cont...)
Summary of Characteristics of Fossil Lagerstätten Listed in Factsheet 2.12 :
(a) Upto Palaeozoic
7. Palaeoecology No predators, Entire ecological Nektons likely Preponderant Rapid burial of a Rapid burial in
detritivore, dynamics: predators and/or carnivores, mixed assemblage concretions
suspensivore, predators, scavengers; some in anoxia in before much
planktivore scavengers, benthos detritivores; phases, by turbidity compaction
filterers, filter-feeders/ no herbivores currents caused by preserved
collectors, deposit-feeders tropical storms
swallowers
8. Special Soft bodied Evidence Replacement of Fossils in Mineralized and Siderite
Feature organisms; of Cambrian organic materials silicified particularly soft- concretions
squashed, explosion by clay minerals mode parts pyritized have 3D fossils
yet wealth in cyclic tidal
of data sediments

Fossil Lagerstätten:
(b) Mesozoic

1. GRÉS À HOLZMADEN SOLNHOFEN MORRISON SANTANA AND CRATO

VOLTZIA SHALE LST. FORMN FORMATIONS

2. Semi-arid Marine plus Basin marine, Terrestrial Salinity-stratified Shallow embayment

terrestrial, land biota juxtaposed O2 deficient lake to coast with marine
brackish incursions

3. Different in Allochthonous Mostly allochthonous Allochthonous Allochthonous

different facies

4. Varied Pseudo- and Benthos fossils Varied Crato:aquatic to Santana: fish

nektoplanktons broken land insects, dominant; pterosaurs
and true nektons, excep, cyanobacteria plants, arthropods, and dinosaurs
rare benthos cyanophyte mat

5. Soft Soft part Exquisite Good Good Good

6. 3 facies of Marine commu- Conservation by Rare Jurassic Mass mortality Soft tissues in
a delta nity + allochtho- interplay of land life from salinity and concretion preserved
nous terrestrial sedimentational O2 deficiency; in Caphosphate
elements biostratinomic- burial hinder
diagenetic rapid decay
conditions

7. Wet-dry Trophic web Part trophic web, Acidic condition in Crato Fmn in Santana Fmn in
alternation partially preser- palaeoecology drying water bodies stagnant fresh- shallow embayment
and microbe ved with consu- evident with animals around water lake
help soft part mers to predators facing non-catas-
preservation trophic mass
mortality
(Cont...)

8. Unique feature of
phosphatization
helped by
microbial mat
1. GRUBE MESSEL
2. Crator lake biota and rare
forest biota
3. Allochthonous
4. Mammals, bird, arthropods,
plants of terrestrial and
forest type
5. Occasional soft parts
6. Syn-depositional faulting and
volcanism, crator lake and
forest around control
sediments and burial
7. 3 alternative models:
1. Lake-river system,
2. Large lake,
3. Crator lake; final say
yet undecided
8. Soft part preserved by anoxia
from volcanic gases and by
algae using up all O2
FACTSHEET 2.13 (Cont...)
Fossil Lagerstätten:
(b) Mesozoic
Hostile hyper-saline
basin, sudden burial,
impervious micritic
host help soft part
preservation
Fossil Lagerstätten:
(c) Cenozoic
BALTIC AMBER
Temperate to tropical and wet
‘Baltic Amber Forest’
Allochthonous
Winged insects, arachnids, etc.
varied plants, angiosperms
Good; soft parts preserved
Large scale preservation
in amber
Temperate to tropical change
over time in a wet humid
condition
Soft part preserved in amber
resisting decay
Chapter 2 Taphonomy 31
Vertically paired two limestone for-
mations, (Santana younger) with a third
evaporite in between. Crato compare with
Solenhofen Lst. micritic limestone,
Santana with Mazon Creek in carbonate
nodules, phosphatized fish.
RANCHO LA BREA
Viscous asphalts as natural trap of organisms
Allochthonous
Trapped in summer, solidified in winter and
covered by fluvial deposits
Good; soft parts preserved soft parts preserved
Concentration of rapidly buried and asphalt-
impregnated fossils with organic materials and
bone-details
Terrestrial ecosystem in a cool, glacial climate
Carnivores outnumber herbivores in an inverse
trophic pyramid
 3
Palaeoecology
3.1 Introducing Palaeoecology
As discussed in Chapter 2, taphonomic
processes, when studied, help us know how and
under what condition the organic remain or trace
was buried or preserved. From there we can trace
back the conditions under which and the ways
in which the concerned organism might have
lived (Factsheet 3.1).
Ecology studies these conditions and ways of
living of recent organisms. Palaeoecology is the
extension of ecology to ancient materials; it is the
study of interrelations and interactions between
ancient organisms now represented by fossils and
the environments of the geological past in which
they lived.
Certain essential terms and concepts of ecology–
palaeoecology are defined in Factsheets 3.2(a) and
3.2(b). They will be referred and discussed in proper
context with or without being defined again. Here
we proceed, however, on their basis.
Each organism (a population, a species or any
higher taxon) lives in a particular environment,
being in constant interrelation and interaction with
it. Man knew it since long from his observations,
though he was not fully aware of all details and
niceties. Thus, when Leonardo da Vinci observed
that fossils in the rocks at the top of certain Italian
32
hills were of organisms that could be found in the
recent seas and when he, on the basis of this fossil
content, interpreted those rocks to have formed
under sea, he was using this knowledge. Similarly,
we can suggest the marine origin for host rocks,
now exposed on land, from abundant fossils of
brachiopods or echinoids in them, or interpret
fluvial or lacustrine origin of a shale, which bears
dinosaurian footprint on its bedding plane. In the
latter case, from the nature of the footprints, their
number, spacings, depths, etc. we may even infer
if the dinosaur walked on all four legs (for instance,
as Stegosaurus walked) or on the stronger
hindlimbs in a kangaroo-like stance (as in
Titanosaurus), thus interpreting the locomotory
habit of the concerned organism. In some other
case, we can interpret an inequivalved bivalve fossil
as of an organism which, in all likelihood, lived
attached to the bottom, as against an equivalved
inequilateral bivalve fossil representing the normal
burrow-making mode of living of those organisms.
Obviously to reach the above conclusions we need
some premise. We must recognize our fossils as
brachiopods, echinoids or dinosaurs. Then we must
know from our observations that present-day
brachiopods or echinoids are strictly marine, and
on that basis interpret their fossils as also of marine
type. But since dinosaurs are extinct, we have to
 Chapter 3 Palaeoecology 33

FACTSHEET 3.1
Palaeoecology and Taphonomy in Relation to
Process of Fossilization
(Based on Lawrence 1968, Martin 1999)

Terrestrial organisms/Epifaunal organisms

LIFE ASSEMBLAGE of ~~~Sediment–water interface ~~
Infaunal organisms
ñ
PALAEO
Death /decay ÿÞ .............................................................................................
ECOLOGY
â á
Dead remains
â
Reworked remains
á
ò
THANATOCOENOSIS Bioturbation
Immediate burial à (biological reworking
Biostratinomic â and destruction) TAPHONOMY
processes/time Þ
averaging
Biostratinomy
TAPHOCOENOSIS Buried remains (physical reworking and
â destruction via
sedimentary processes)
Diagenesis Þ Diagenesis
â Diagenetic alteration
(addition, removal,
FOSSIL Fossil record replacement)
ASSEMBLAGE
———— Collection for study —————————— ñ
depend on other evidences to come to the units were formed on the basis of their fossil
conclusion that their host rocks are continental in content, or the other way round, from different
origin, for instance, association of their fossils with characters of the host rocks, palaeoecology infers
those of big land plants, or simply the interpretation about the habitats and, may be, habits of the
if such skeletal structure could have been borne by organisms whose fossils are enclosed in them.
swimmers or crawlers in aqueous environments. Palaeoecology also may conclude about the habits
and habitats of some particular individual
organism; recall the case of inequivalved bivalve
3.2 Essential Terms and Concepts or two-leg locomotion of the dinosaur, in the
examples above (palaeoautecology). In other
3.2.1 Palaeoautecology cases like a coral limestone with a varied fossil
and palaeosynecology assemblage including coral fossils occurring
attached to the bedding, i.e., preserved in situ at
Thus, to begin with, palaeoecology interprets the their living position, palaeoecology may have to
environment or condition in which certain rock deal with a fossil community of reef environment,
34 Part One: Principles

FACTSHEET 3.2(a)
Essential Concepts and Terms

Palaeoecology: Study of ancient organisms in relation to their environment.

Palaeoautecology: Branch of palaeoecology that is concerned with studying life habits of organisms and
relation of individuals to their environments. It concentrates on growth and shapes of organisms and the
correspondence of morphology to both life strategies and habitats (Brenchley and Harper 1998).
Palaeosynecology: Branch of palaeoecology that is concerned with studying ecology of fossil communities,
including environment and relationships among the members of the community.
Ecosystem: A chosen portion, large or small, of the physical environment plus all the organisms contained in
it. It is the largest unit of study in ecology, though by definition it includes from the biosphere itself to a tiny
puddle of rainwater with two or three species of one-celled organism.
Habitat: The environment in which an organism lives. There may be more than one habitat in an ecosystem.
Thus, a rocky shore or a beach, a grassland or a forest and the intestine of a larger animal which may house a few
parasites living on it, are all some kinds of habitat. Habitats of the marine ecosystem include littoral flats to
abyssal plains, from shallow photic zone to lightless deep water, or substrates varying in different aspects, such
as soft, loose sediment covered to hard, rocky, or quiet to turbulent, etc.
Ecological Niche: Variously defined, the niche may be the environment or its features that permits or permit
the organism to live most successfully or niche may be the organism’s position in the habitat, including its way
of life and the role it plays in the ecosystem. Most habitats are occupied by several species, each with its own
ecological niche.
Species, Individual, Population and Community: Usually, a species, living in a habitat, is represented by
two or more individuals that constitute a population. A community is an array of populations of two or more
species of animals and plants (plus unicellular organisms) that inhabit a common region of a habitat and live in
close interrelation and interaction maintaining trophic and different other relationships (such as symbiosis,
parasitism, commensalism, epibiosis, etc.) among themselves.
Habit: Also referred as mode of living, habit is the way in which an organism lives. In a water body, e.g., a
sea, an organism may live attached to the bottom or substrate, it is benthic and fixosessile in habit, or it may be
water-dwelling (pelagic), either swimming in habit (nektonic) or passively floating (planktonic).

FACTSHEET 3.2(b)
Essential Concepts and Terms

Biological Environment: In addition to the physico-chemical controls of environment, organisms are also
affected by their biological environment. Two fundamental relationships of biological environment are as
follows:
Trophic Chain (food chain/food web): A closed chain of fundamental interrelationship among organisms
of a community that controls transfer of materials and energy to maintain the metabolic processes of the
organisms. It includes:
Primary producers that photosynthesize organic compounds with the help of solar energy, i.e., they
produce their own food material (autotrophs: e.g. plants). Heterotrophs that acquire food material
from other organism either by ingestion or taking in as food (animals) or by absorption (fungi). Among
them, consumers feed on others: herbivores, the first level consumers, consume plants; carnivores
feed on herbivores, or the higher level consumers on smaller carnivores of lower level consumers.
Parasites feed on living organism, producers or consumers. Scavengers feed on dead remains of either
of these or all.
(Cont...)
 Chapter 3 Palaeoecology 35

FACTSHEET 3.2(b) (Cont...)

Essential Concepts and Terms

Decomposers (mainly bacteria) break down the left-over unassimilated organic materials. Transformers
(also bacteria) chemically change the decomposed products to be used by the producers again as
nutrients.
Ideally, producers–consumers (first, second, third levels, etc.), by number, form a pyramid with the producers
at the base, the highest level consumers at the apex.
Species Relationship
Symbiosis: Two different species living in close association with each other; at least one of them is benefited
or harmed by the other. It includes:
Commensalism: None of the two is of harm to the other, though one may be benefitted. Fungi on
tree trunks (+, 0).
Mutualism: Both of the two are benefitted and are without disadvantage. This is often referred as
symbiosis itself. Coral-Zooxanthellae (+.+).
Parasitism: One of the two benefits at the expense of the other. Worms in intestines of other animals.
(+, –).
Epibiosis: Not necessarily involving two living species; one (the epibiont) attaches itself to the firm
substrate of the hard part of the other, live or dead. Worm or barnacle on mollusc shells.
Antagonism: One of the two species is definitely harmed.
Antibiosis: One is harmed, the other is of no particular advantage. (–, 0).
Exploitation: One harms the other to its own advantage (predation). (–,+).
Concurrence: Both are harmed in the process (–, –).
Tolerance: Two related species having no particular effect on either (0,0).
NB: (+) Beneficial, (0) Neutral, (–) Detrimental

its composition, relationship among its members,
as also the physico-chemical environment in which
the community lived and was preserved
(palaeosynecology).
With the organic world constantly evolving
and environments also being in a continuous state
of flux, organism–environment interrelation and
interaction too change with time. Palaeoecology
makes use of the whole geological time on the basis
of the fossil record. So, based on the case studies
of individuals as well as communities represented
as fossil assemblages, palaeoecology also
reconstructs the past ecosystems.
3.2.2 Habits and habitats
On the surface of the earth, there are two basic
types of environment available for organisms to
live in, viz. terrestrial (environment on land) and
aquatic (environment in water bodies, mainly
marine also freshwater or brackish). Organisms
inhabiting these environments show different
habits and habitats too (see Factsheet 3.2,
Figure 3.1). The aquatic environment is obviously
dominated by oceanic or marine environments.
Modern oceanic environments fall into the
following categories, littoral/tidal being part of the
oceanic environment between low tide and high
tide levels on the shore, neritic/sublittoral, the part
of the sea between low tide and 200 m depth; it
coincides with the photic zone in normal clear sea
water on the continental shelf bathyal, the part
between 200 metres and 4000 metres depth of the
ocean on the continental slope, abyssal between
4000 metres and 5000 metres on abyssal plains of
the ocean floor and hadal being deeper parts of
the oceanic trenches. Besides these divisions in
seas, aquatic environment also comprises two
36 Part One: Principles

1 2 3 4 5
6
HTL
LTL

B
A

9 10
STL
6 HTL
LTL
7 11
200 m
8
12

C
13

2000 m
14
17 18 16 15

> 6000 m

Fig. 3.1 Basic environments of land and sea.

(A) From land to sea, mountain to oceanic trench, (B) Major continental environments, (C) Basic
marine environments; Not to scale.
Index: 1. Mountain, 2. Alluvial, 3. Fluvial, 4. Lacustrine, 5. Paludal, 6. Supratidal, 7. Intertidal,
8. Subtidal, 9. Neritic, 10. Oceanic, 11. Photic, 12. Aphotic, 13. Bathyal, 14. Abyssal, 15. Abyssal
plain, 16. Hadal/oceanic trench, 17. Continental shelf, 18. Continental slope, STL (Storm tide level),
HTL (High tide level), LTL (Low tide level).

types, one on or in the floor or substrate of the
basin or the water body, and the other in the water
mass itself. Accordingly, organisms are said to be
benthic/benthonic and pelagic in habit
respectively, on the basis of which of the two
environments they live in.
Organisms benthic in habit further subdivide
into epibenthic and endobenthic, the former
including those living on the substrate, and the
latter in the substrate, within sediments.,
Epibenthics include sessile, i.e. immobile or
attached, being cemented or otherwise, fixosessile
(e.g. Terebratula, a brachiopod attached by a rod-
like peduncle or pedicle; Ostrea, a cemented
bivalve), or attached by long root-like processes,
rhizosessile (e.g., Cryptopora, a brachiopod or
Mytilus, a bivalve attached by thread-like byssus)
or simply free-lying on the bottom, liberosessile
(e.g., Atrypa, a brachiopod genus or secondarily
free-lying bivalves like ‘recumbent’ rudists or
Gryphea). Epibenthics may be otherwise vagrant/
vagile, i.e. moving, generally sluggish (e.g. regular
echinoids or asteroids, i.e. star fishes). Endo-
benthics, in turn, include burrowing (digging in
soft sediments); boring (through harder materials
like shell hard parts, rocks or wood) or nestling
(living in holes or crevasses) types of habits.
Pelagic organisms include nektons or
swimmers in the water mass (e.g. fishes or
cephalopods) and planktic/planktonic, floaters on
the water surface (e.g. planktic foraminifers among
many other planktons). They are further grouped
into neritopelagic and oceanopelagic, accordingly
as it refers to living in neritic water or oceanic
water. More detailed divisions are made on the
depth of water and parallel criterion of light
condition. Thus, these may be epipelagic/euphotic
where water depth is <200 metres and sunlight
penetrates well; mesopelagic/disphotic with water
depth between 200 metres and 1000 metres and
sun-light penetration poor; bathypelagic/aphotic
for bathyal water between 1000 metres and 4000
metres with no sunlight penetrating and
abyssopelagic in abyssal water at depths more than
4000 metres. Organisms living in the respective
Chapter 3 Palaeoecology 37
zones are termed by the zone terms themselves,
viz. epipelagic/euphotic, mesopelagic/disphotic,
bathypelagic/aphotic and abyssopelagic. Planktic
organisms may be phytoplanktons (plant affinity) or
zooplanktons (animal affinity). They are also
recognized as holoplanktic, meroplanktic or
pseudoplanktic accordingly as they are floaters
throughout life (e.g. planktic foraminifers), planktic
only in the larval stage, the later benthic (e.g.
brachiopods, anthozoans, i.e. corals) or are relatively
larger but attache themselves to a floating or
swimming body, e.g. another organism (as
reconstructed for graptolites).
Feeding habit of organisms is another
important category on the question of habit.
Basically, organisms are autotrophs and
heterotrophs, that is, those which feed by photo-
synthesis, generating food from available
nutrients with the help of sunlight (plants) and
others acquiring food either by ingestion of other
organic material (animals) or by absorption
(fungi). Heterotroph animals of aqueous
environments (particularly the primary
consumers) may be filterers/suspension feeders
that suck in water to extract nutrients from that;
swallowers/deposit feeders that gulp in sediment
rich in organic material; collectors/detritus feeders
which sweep up detritus, i.e. non-living
particulate matter derived from bodies of
organisms; scavengers that feed on dead organic
remains; grazers or selective collectors on hard
substrate; predators/hunters that prey upon living
organisms. Terrestrial organisms fall into
herbivores, including browsers/folivore or leaf-
eaters, frugivore or fruit-eaters, granivore or
grain-eaters and grazers or grass-eaters on one
hand and carnivores or flesh-eaters on the other.
3.2.3 Limiting factors
Thus, every organism has a typical living and
feeding habit and prefers a particular habitat, an
environment. As mentioned, there are two basic
types of environment. These are namely aqueous
and terrestrial. Of the former, there are again two
38 Part One: Principles

different kinds: marine and continental. In water
bodies, life is controlled by factors such as depth,
temperature, solute content or salinity, oxygen or
other gas content, the amount of suspended
particulate matter, etc. Life on land is determined
again by temperature, precipitation or rainfall and,
thus, humidity and aridity.
Potential niche of a species is defined by many
parameters. But in a particular environment some
factors have a more profound effect in determining
the range of a species than others. These are called
limiting factors. For instance, in normal sea water
salinity and oxygen levels remain broadly constant
whereas temperature may vary considerably as also
the depth of the substrate. The latter in turn affects
penetration of sunlight and, thus, the abundance
of phytoplanktons, the primary producers,
Temperature is also a major factor that controls
the nature of the biota. Thus, temperature and depth
become more vital in determining the presence or
the absence of different species than salinity and
oxygen. The major limiting factors that control
biotic distribution are listed in Factsheet 3.3.
Controlled by them, organisms live fitted to some
environment or other.
3.2.4 Adaptation
It is the theory of evolution by natural selection that
explains two different aspects of the living world,
diversity and fitness. About 2 million species are
now living and at least 99.9 per cent of the organic
kingdom of the geological past are now extinct. This
diversity is explained mainly by evolution. One
major question that the theory of evolution has to
answer is ‘where did they all come from?’
Secondly, organisms fit remarkably well into
the external world in which they live. They have
morphologies, physiologies and behaviour that
appear to have been carefully and artfully designed
to enable each organism to appropriate or make
use of the world around it for its own life. That is,
in other words, they adapt. (Factsheet 3.4).

FACTSHEET 3.3
Limiting Factors: Distribution and Abundance of Organisms

General Mainly for aqueous environment Mainly for land

Physical
Temperature Depth of water Precipitation
Latitude–Longitude Hydrostatic pressure Atmospheric condition
Light conditions Distance from shore Soil type and thickness
Land/seafloor surface Water movement Groundwater condition
Viscosity and diffusion Shape and openness of the water body
of air/water Geography of land around
Sedimentology
of water and substrate areas
Chemical
Gaseous content Solute content and salinity
Carbon dioxide, Oxygen, Colloid concentration
Hydrogen sulphide Nitrogen Calcium
Eh-PH potential Eh–PH potential

Biological
Trophic relation
ü
ý Pertain to both
Species relation
þ
Mobility
 Chapter 3 Palaeoecology 39

FACTSHEET 3.4
Adaptation

Environment around organisms to live in, sets problems for them to cope up with.
Evolution, by means of natural selection, is the mechanism for creating solution.
Adaptation is the process of evolutionary change by which the organism provides a better and better solution to
the ‘problem’ that the environment sets.
Thus, adaptation brings about ‘fitness’.
That is, every organism is adapted or ‘fitted’ to a specific mode of life and is constrained by environmental
limitations.
Adaptation helps organisms to (i) cope with changing environments, (ii) invade new environments, (iii) diversify
and (iv) function more efficiently.
Evolution produces diversity, i.e. different kinds of organisms. Adaptation may conserve, i.e. independently
develop similar morphologies that are related to similar functions.
Homeomorphy is such similarity in morphology in unrelated organisms.
Problems Evolution } Adapted
Organism « Environment ®

Solution Adaptation } Organism

Morphology is basically adaptive. Functionally neutral features rarely form an important skeletal or anatomical
part. All biologically feasible morphologies do not occur in nature. That is, they are not utilized by organisms
that actually evolve. It means adaptation is not random. It is directed towards functional efficiency. Areas of
poor adaptation , i.e., parts of the total range of feasible morphologies not occupied by or rather not realized by
organisms, are adaptive valleys or biological impossibilities.
The case is manifested by coiled shells. Not all the possible types of coiled shells are found in organisms; only
those that help the concerned organisms perform their necessary functions, occur in reality.
Example of adaptation: With the evolution of birds from reptiles, there was a successive alteration of bones,
muscles and skin of the forelimb to give rise to wing. Increase in the size of the breastbone provided anchor
for the wing muscle; restructuring of bones made them light and strong; development of feathers provided both
aerodynamic elements and lightweight insulation. This is the process of major adaptation by which birds solved
the ‘problems’ of flight. Yet there is no end to it. Some birds reversed the process, e.g. penguins adapted to
marine life by changing their wings into flippers and feathers into waterproof cover; ostriches became ground
dwellers with strong and long legs and relatively smaller wings.
A few more terms:
Adaptations are fits of organisms shaped by natural selection. Exaptations are functionally useful structures
that apparently were not shaped by natural selection. Aptations are any structures fitted to a particular function,
thus including both adaptations and exaptations. Preadaptation involves structures or groups of structures
already functional but available and suited by chance for a more innovative function. Postadaptation occurs
when the newly developed function of a preadapted structure becomes the main function of the structure in
future generations. Adaptation and preadaptation are the more used terms.

Patterns of distribution and abundance of environment fails or changes, organisms either shift
organisms are controlled, both in time and in space, (migrate) or die, or they evolve and in course of
by the interrelation–interaction of organisms and evolution they adapt. Very simply told, organisms
their environment. At any particular time, if cope or adjust with their changing environment,
40 Part One: Principles
invade newer ones and function more efficiently
in a given environment. That is, they adapt to
environment and to functionally efficient habit to
live smoothly and successfully.
3.2.5 Echinoid adaptation
Factsheet 3.4 gives examples of adaptation. In
addition, we may consider the specific instance of
echinoids, a group of ‘invertebrate’ organisms. It
is recommended, however, that relevant sections
of Chapter 13 should also be seen. The group
represented by their Regularia genera and species
in Palaeozoic, faced a major change in
environment in mid-Mesozoic. With widespread
and profound changes taking place in the land–sea
configuration in result to the break-up of the
Pangaea and with modern oceanic regimes coming
into being thereupon, there was also a change in
the substrate condition on ocean floors. With new
sedimentational regimes having been set up, the
floors were laden with sediments. Echinoids that
were adapted to rocky bottoms and lived
epibenthic life, passed through adaptation to a
endobenthic life, adapting to a burrowing or other
infaunal mode of life. Irregular echinoids took the
scene, with regular forms dwindling in the process.
Regular echinoids were broadly characterized by
tests with radial symmetry of equal-sized simple,
ambulacra containing circular pores for tubular
tubefeet, anal and oral openings placed at the
centre of the aboral and oral surfaces respectively,
diametrically opposite at the ends of the aboral–
oral axis. This fitted with epibenthic mode of
living, which met with similar conditions all
around. Adaptation to endobenthic mode led the
organism to face different environments in front
(the dead end of the burrow) and at back (sediment
water interface and the water mass to derive
oxygen and nutrients from, as well as free space to
discharge wastes), but similar environments on the
sides. Burrows also provided limited oxygen
supply, demanding special device to acquire
adequate oxygen. All this led to a bilaterally
symmetrical test with ambulacra differentiated into
unpaired anterior and two pairs of laterals behind,
development of slit-like pores for appressed
respiratory tubefeet in ambulacra on their aboral
sides, shifting of anal opening towards posterior
and mouth towards anterior, thereby defining and
lying on the symmetry plane of the test of irregular
echinoids.
3.2.6 Adaptation and functional
morphology
Morphology of organisms develops through
interplay of three factors, viz. adaptation
(controlled by ecology/palaeoecology), phylogeny
(controlled by evolution) and growth (controlled
by ontogeny). Every organism of a species or
population possesses morphological charac-
teristics that are largely determined by its genetic
attributes it inherits from ancestors in course of
phylogeny. During its ontogeny (from larval to
senile stages) too, there are morphological changes
that are also controlled by its genetic characters.
At the same time, on this phylogenetically and
ontogenetically controlled material, natural
selection acts upon to preserve through generations
only those characteristics as stable and functional
which help organisms, i.e., members of the species,
to adapt to its environment. As a result, occurrence
of adequate and abundant individuals of a species
or population in any fossil assemblage, would
suggest that the species was successfully adapted
to the environment. In that case its morphological
features must have performed some effective
functional role in its living. It further means that
morphological features are generally adaptive.
Functionally neutral features rarely form an
important skeletal or anatomical part.
All biologically feasible morphologies do not occur
in nature.That is, they are not utilized by organisms
that actually evolve. In other words, adaptation is
not random. It is directed towards functional
efficiency. Coiled shells present a case. All the
possible types of coiled shells are not found in
organisms (see Figure 3.2); only those that help
 Chapter 3 Palaeoecology 41

d1 Q
r1
t1
r2 d2 t2 R
c2 (ii)
c3

is
Expansion rate (W)
d3 r3 l
ira

ax
1
(i) isp

m
n

fro
10 a
Pl rms

e
Fo

rv
102

cu
at 0
P Axis of Coiling (iii)

er 1.
g
in
Helicoid Forms

en 0.8
4
10 (iv) 6
Q Initial generating curve 0.

(D 0.4
fg
R Generating curve after 1 coil

)o
2
6
10

0.
4 3 2 1 0
Translation (T)

e
nc
ta
is
D
(a) (b)

Fig. 3.2 Modelling morphology of coiled shells

(a) Schematic representation of a gastropod shell to mark parameters used in part (b) W- c3r3 : c2r2;
c2r2: c1r1; ... D-d1r1, d2r2, d3r3 ... T- t1t2, t1t3 ...
(b) Domains of coiled shells found in nature marked out in computer-simulation model
(after Raup 1967). (i) Gastropoda, (ii) Cephalopoda, (iii) Bivalvia, (iv) Brachiopoda.

concerned organisms perform their necessary
functions, occur in reality. This attests to their
functional efficiency or usefulness in terms of life-
activities. Varied combinations among genetic
relationship, adaptation and morphology are shown
in Factsheet 3.5.
In result, morphological features of fossils may
be analyzed to bring out their functional
importance. From there, we may infer the mode or
habit of living of ancient organisms as well as their
environment. This gives rise to a very useful and
popular method in palaeoecology, namely,
functional morphology.
Specific instances will be discussed more in
context with discussions on morphology of
different invertebrate groups. Here, as a general
discussion, the following examples may elucidate
what functional morphology is.
For epibenthic organisms (related terms:
epifauna; epibiota) living on the substrate, either
attached or moving, rocky/shelly substance
provides a firm ground for fixation or mobility.
Adequate presence of such fossils, thus, speaks of
a sediment-free substrate of the basin. Each sessile
epibenthic organism faces different environments
below and above; hence its body or hard parts
become asymmetric vertically; likewise, shells
become inequivalved in sessile bivalves or
brachiopods. The lower attached valve serves as
the abode and is larger; the upper valve serving
more as a protective lid and, thus, requiring vertical
movement to open or close the shell, can afford to
be just fitting to the other one, even smaller to
reduce the weight. Any benthic organism living in
a large water body also has to bear the hydrostatic
pressure of the huge water column above. Hence,
42 Part One: Principles

FACTSHEET 3.5
Morphology, Genetic Relation and Adaptation: Variation in Relationship

Morphology Example Relationship

Equivalved shell
with equal and relatively
thin valves
Inequivalved shell
with unequal and relatively
thick valves
Inequivalved shell
with unequal and relatively
thick valves
Similar external morphology
for smooth movement
in water
Development of
hollow bones
and transformation of
forelimbs into wings
ü
Most bivalves ï
of vagrant habit Genetically related,
ï Morphologically different
ý Adaptation to
Sessile bivalves, many
üï different environments
of which form shell-banks
ïï and different habits
ïï
in shallow seas
ïþ
ï Genetically unrelated,
ý Morphologically similar
Few brachiopods like ï Adaptation to the
Lower Carboniferous ï same environment
genus Gigantoproductus ï
which also probably built ï
shell-banks in shallow seas þ
Selachian fishes ÞÑ
Jurassic Ichtyosaurian reptile ß Genetically unrelated,
Mammals like dolphins Ñà Morphologically similar
Adaptation to the
Mesozoic flying reptiles ÞÑ same environment
Birds ß
Chiropteran mammals Ñà
(e.g. bats)

its body and the skeleton is either horizontally
spread, as in larger benthic foraminifera with disc-
shaped tests, or dome-shaped, as in echinoids, or
vertically grown as tubes or cones as in corals or
gastropods.
Endobenthic organisms (endofauna/
endobiota) may bore into harder materials like shell
hard parts, rocks or wood or may live in holes or
crevasses. But more commonly they dig burrows
in soft sediments and while doing so, find different
environments in front and at back, but similar
environments on the sides. Hence, as we noted with
echinoids in section 3.2.4, we also find in
burrowing bivalves, shells becoming antero-
posteriorly inequilateral, yet equivalved. (See also
Chapters 8 and 10.)
Nektics or swimmers develop bilaterally
symmetrical streamlined body for swift and smooth
movement in a fluid, as found in fishes or
cephalopods; this meets their hydrodynamic
requirement. On the other hand, to maintain their
vertical movement in search for food or safety from
predators, they need devices like air-bladder in
fishes or siphuncle in cephalopods. Also to
withstand the variation in hydrostatic pressure
acting on their body or shell at different depths,
they develop reinforcing structures in their
skeletons; septa in cephalopods provide such shell-
strengthening device, in the way partition walls
hold a large ceiling back from collapsing.
Planktics are fitted to their mode by dint of
their light, hollow body or hard part with increased
 Chapter 3 Palaeoecology 43

surface area or floating device such as hollow
spines, hair-like extensions, etc. as found in
planktic foraminifers and other groups. Pseudop-
lanktons require a device to cling to a floating or
swimming body; nema in graptolites may be an
example. Small, ever-moving larvae of many
sessile benthic animals like brachiopods or
anthozoans (corals) stand for examples of
meroplanktons. They explain the occurrence of a
number of vertically short-ranging yet widely
distributed stratigraphically important species in
those benthic groups.
These examples of morphological features
serving functions are a few among many others.
Functional morphology, a modern upcoming
branch of palaeontology takes care of them.
3.2.7 Ancient community to fossil
assemblage
Environment to which organisms are adapted, has
two basic components, a set of physico-chemical
conditions to delimit it and a set of organisms
specifically adapted to it. In fact, organisms tend
to live in communities, each a natural association
of living species in a given area. The member
species are interrelated with some others of the
community, in addition to their being interrelated
to physico-chemical or abiotic environment. The
most fundamental of these interrelationships is
the trophic relationship; other relationships are
defined as symbiosis, antibiosis, etc. [See
Factsheet 3.2 (a).]
A fossil assemblage which is the sum total of
fossils occurring in a particular horizon of any
particular place or time, is, in fact, the fossil record
of a community of that time (fossil community),
which lived in that area and was fossilized and
preserved in part or whole and with or without
additional members (Factsheet 3.6). A study of the
composition of an assemblage may lead to clues
about which of the constituents were linked to each
other and how or in what relationships. The fossil
community structure may then be envisaged.

FACTSHEET 3.6
A Living Community may Leave a Complete Record, or a Part of it, or None at All.

This is how:
LIVING COMMUNITY in one area at one moment may change with
(A) (B)
Migration All migrated All remained to die
Timing of death All died normally All killed simultaneously
at different times
Physical removal All removed None removed
Scavenger/ saprophyte action All destroyed None destroyed
Removal by solution, etc. All removed None removed
Preservation adequate None so All so
No addition of Earlier inhabitants Later inhabitants,
Derived fossils
Discovery and collection None found All found
Hence Record lost Record complete
NB: A and B represent two extreme cases; in A the ancient community leaves no record, in B a complete record is left.
In still other cases, there may be any position in between.
44 Part One: Principles
This community-assemblage pathway is the
domain where taphonomy merges with
palaeoecology.
At one stage, understanding of taphonomy
hinged on the following tenets. It was held that
organisms are more likely to be preserved if they
have hard parts. Their preservation is, however,
greatly enhanced by rapid burial, especially in fine-
grained sediment (low turbulence) or in the absence
of decay and scavenging.
Subsequent addition of data and improvement
of understanding have led to following
appreciations: a community maintaining trophic
and different other relationships (such as
symbiosis, parasitism, commensalisms, epibiosis,
etc.) may, and generally do, contain organisms
with lesser preservation potentialities either due
to their not having hard parts or their niche (for
instance, water for pelagic forms) not coinciding
with the places in which deposition is taking place
(in the above case, the sea floor). Hence, an
ancient community is generally not represented
in full in the corresponding fossil assemblage
(Factsheet 3.1).
A fossil assemblage is then a community of
ancient time (fossil community), in part or whole
and with or without additional members that are
found together in the fossil record of any particular
place or time. It is also called a taphocoenosis,
equivalent to most fossil communities (Ziegler
1983), created by various taphonomic processes
that act on the remains of organisms.
It may contain (i) an assemblage of dead
organisms that lived together and was preserved
in situ (Ziegler 1983) after death and decay
(Brenchley and Harper 1998) {Biocoenosis/
Autochthonous thanatocoenosis} (obviously this
will include only benthics of the area); (ii) an
assemblage of dead organisms derived from
without the area of deposition or fossilization. It,
thus, includes organisms of different habitats
(Thanatocoenosis/ allochthonous taphocoenosis)
(for example, remains of benthics of different
habitats accumulated or dead bodies of pelagic
forms showered down, on the floor of basin and
fossilized with remains of in situ benthics there);
and (iii) an assemblage of fossils of organisms,
which are allochthonous or foreign remains
derived from reworking of some other fossil
assemblage of a different time{Reworked/
derived}, generally from older deposits; in rare
cases, younger fossils may be reworked into an
older deposit (reports of microfossils leached
from Tertiary rocks and carried and deposited in
highly porous Cambrian Saline Series of Salt
Range of Pakistan). Authors (Ziegler 1983;
Brenchley and Harper 1998) differ in their
definition of different terms in relation to the
components of a fossil assemblage. Hence, the
terms are left here in parentheses and shown in
Factsheet 3.7.
During this course (ancient community to
fossil assemblage) processes like disarticulation
and chemical alteration resulting from decay,
abrasion, transportation, predation, scavenging, or
dissolution, etc. may cause loss of information
(or addition of allochthonous components, by
transportation and otherwise) about species
abundances and community diversity and structure.
These need to be taken note of and sorted out while
studying a community.
3.2.8 Palaeoecology redefines its course
Palaeoecology deals with all these aspects,
palaeoautecological or palaeosynecological,
about ancient organisms of a particular time or
of different times. In the last few decades it has,
however, witnessed a major change, from a
habitat approach to an ecosystem approach. The
goal has shifted too. It was set at delineating or
reconstructing habitats or mainly the abiotic
controls of environments of ancient organisms in
question. From there, it now tries to develop an
understanding of the various interactions and
interrelations, both biotic and abiotic, with which
ancient organisms may have lived, singularly or
in community. As it is appreciated more and more
 Chapter 3 Palaeoecology 45

FACTSHEET 3.7
Community to Fossil Assemblage or Taphocoenosis.
Living Place versus Burial Time and Place Relationship

Living and burial takes place at

the same place different places
Same time Biocoenosis Indigenous Thanatocoenosis
Ichnocoenosis (slight movement) Vertical settling of planktics-nektics
Horizontal shifting of benthics
Different times
From older Reworked {Remanie} Reworked
From younger Leaked Leaked

that the biosphere has changed closely linked with
the changes in the planet earth itself,
palaeoecology has also broadened its scope to
include the study of how ecosystems have evolved
through time. Relationship between the organic
world and the physical world has also been
reinterpreted as two way interactions: not only
environment influences organisms, organisms
influence environment too. Classically field and
laboratory observations and studies were the
methods. Palaeoecology today takes help of
additonal methods of mathematical–statistical
analysis, theoretical–geometrical modelling,
stable isotope studies and such other sophisticated
techniques to reach its conclusions.
Moreover, as a composite science palaeo-
ecological studies involve a few other major
aspects. In the organic world there are about 2
million species now living and at least 99 per cent
of the organic kingdom of the geological past are
now extinct. Organic evolution explains wherefrom
did these species come, why were they extinct and
how did they change. Besides, different organisms
inhabited the earth at different times and had
different habitats and habits. So, with a view to
reaching right answers to palaeoecological
questions, one must have clear knowledge and
understanding of the systematics and stratigraphy
of the concerned organisms too.
3.3 Methodology of Palaeoecology
As mentioned in the last section, with the scope of
palaeoecological studies broadened and changed,
the subject has adopted more modern methods.
Details of the methodology cannot be included in
one single discussion. Only a few major ones will
be mentioned here.
3.3.1 Palaeoecology works on simple
premise of principle of
uniformitarianism
However, complex and diversified the subject may
have become, it works largely on a simple basic
principle of actualism or uniformitarianism. It
holds that the relationships and interactions of
present-day organisms and their environments may
be used to arrive at the conclusions on such
relationships and interactions for similar and
related organisms of the past. For example, extant
scleractinian reef-corals are confined to shallow
tropical seas. From this, it is interpreted that
Palaeozoic bioherms with extinct tabulates and
tetracorals (rugoses), too, were formed in the then
shallow tropical sea waters.
Such uniformitarian conclusion has two parts:
(i) Palaeozoic or mesozoic bioherms (or the then
coral reefs) formed in an environment same as that
in which the present-day coral reefs form and
46 Part One: Principles
(ii) extinct groups of tabulates and tetracorals lived
in environment same as that in which present-day
scleractinian corals live.
Here the approach has its limitations. It
presupposes that processes and conditions
remained broadly similar through time, which may
not be so because of the following reasons:
1. There may have been major changes in abiotic
environment during the geological past. For
instance, till there was free oxygen in the
atmosphere, in the earlier parts of
Precambrian, modern life forms were not
likely to appear and survive; conclusions in
regard to oxygen-breathing organisms may
not pertain to those of the oxygen-less earth.
Oxygen content of the atmosphere changed
probably markedly during the Devonian also,
with the advent and evolution of land plants.
The type and amount of elements contributed
to the oceans changed with repeated uplift and
denudation of continental land masses. The
amount of carbon dioxide available to the
ocean may vary according to the extent of
tropical forests where it is generated and the
volcanic activity, particularly at the end of
Cretaceous. Continental drift or plate
tectonics has altered through geological time
the amount of continental shelf present, thus
affecting the depth parameter; and the
latitudinal situation of shelves and seas
(crossing through temperature zones) and,
thus, the number and types of ecologic niches
present. The present day landsea distribution
was initiated in Mesozoic with the break up
of Pangaea. Modern environments to which
extant forms are adapted, not being avaliable
earlier, the then organisms must have had
different kinds of adaptation, as we have
discussed in the case of echinoids. These and
other changing features of the abiotic
environment on the earth demonstrates that
we should relate the evolution and adaptation
of marine organisms to the constantly
changing face of a dynamic earth.
2. There may have been major changes in the
biotic environment during the past, on which
basis interrelationship of organisms has also
changed. Appearance and expansion of
photosynthetic cyanophytes that gave off free
oxygen to the atmosphere, might have trigered
changes in atmosphere from oxygen-less to
oxygen-bearing. Modes and demands of life
of shelled organisms were different from those
of earlier organisms without any mineralized
parts in the body. Appearance and spread of
flowering ground-plants or grasses in Oligo-
Miocene helped evolution of grazers among
land vertebrates. Morphological expressions
of grazing habit are, thus, not expected in
earlier forms.
3. Major changes in adaptation in many biologic
groups may demand critical application of the
principle. Endobenthic habit was not there in
Palaeozoic echinoids; planktic habit appeared
in calcareous foraminifers only in Upper
Cretaceous. So, earlier echinoids or
foraminifers cannot be judged on the example
of endobenthic echinoids or planktic
foraminifers.
4. Some unique events like the origin of life,
origin of different animal phyla are non-
repetitive; the conditions in which they took
place were also unique and non-repetitive.
Their interpretation must take note of the fact
and cannot be explained on the basis of any
existing examples.
5. Preservation potential of evidences demand
consideration. Modern reef-corals have
symbiotic relationship with zooxanthellae, a
photosynthetic brown-algae that restricts the
former within the photic zone. But they are
not preservable in fossils and so one cannot
be sure of any such symbiotic relationship
between reef-corals of Palaeozoic and any
photosynthetic organism, and from that,
whether they were restricted to the then photic
zone. Siphuncle of cephalopods, including
extinct ammonoids, is an important part of

their body. It helps the animals in their vertical
movement through water. But the structure is
fragile and are often not preserved.
6. Conclusions on extinct groups on the basis
of their extant equivalents may not be always
precise and correct. Thus, in the earlier
example, tabulates and tetracorals of
Palaeozoic might not have lived in the same
environment under similar controls as their
extant counterparts, scleractinians do.
Trilobites, another Palaeozoic extinct group
of organisms close to or belonging to
arthropods have their carapace, the hard
skeleton made of calcite, whereas arthropod
hard parts are made of a combination of chitin
and calcium phosphate. Any functional
significance of these two different kinds of
hard parts may not be the same.
7. Positive or negative taphonomic changes
during the entire pathway from the burial of
a balanced organic community to the
collection of a fossil assemblage in the rock
record, may affect the application of the
principle of actualism. Benthics, particularly
sessile benthics and trace fossils, belonging
to biocoenosis and ichnocoenosis are more
suitable for the purpose of palaeoecology,
though nektics and planktics, too, often stand
important in palaeoecology. Terrestrial
organisms are not likely to be preserved until
their remains are transported from land to a
basin and are covered. Transportation is likely
to cause mechanical damage and
sedimentation on land or basins there is not
generally prolific.
However, there is a basic unity betweeen
adaptation on one hand, and mode of living, habit
and environment of organisms on the other. In
result, achievable adaptative patterns are never
random and infinite, rather they are quite a few.
And so, in spite of all the difficulties and
limitations, the principle of actualism remains a
fairly effective tool in palaeoecological studies.
Chapter 3 Palaeoecology 47
3.3.2 Some major methods
Palaeoecological studies on fossils may help
cast light on the following aspects of mode of living
and activities of the concerned organisms:
(a) How did the organisms live?
(b) Were the organisms mobile or sessile?
(c) Did the organisms live attached to the
ground, or were they free-lying?
(d) How were they attached, if so?
(e) How did they feed?
(f) How did they reproduce?
Answers to any or all these questions may be
addressed in a number of ways.
3.3.2.1 Directly from preservation: Coral
(anthozoan) fossils preserved unbroken and
vertically upwards at right angles, or nearly so, to
the bedding plane are likely to have been preserved
in situ, in living condition attached to the sea-floor.
So these also suggest fixosessile habit of the
organisms. A few rarer instances may provide
significant information. In one such complete
mature shell of a brachiopod genus, Composita is
found inside the large pedicle valve of another
brachiopod, Hercosestria. The latter has an
aberrant morphology with a large subcylindrical
pedicle (ventral) valve and a small lid-like brachial
(dorsal) valve. The pedicle valve has a meshwork
at the top even above the brachial valve. The
Composita shell rests on the brachial valve.
Obviously, the animal Composita passed through
the meshwork of the pedicle valve of Hercosestria
during its larval stage and grew up inside the shell
of the latter genus. It could not have survived and
grown had the animal Hercosestria carried out its
feeding with the help of its lophophore thrown out
of the shell for food catching and creating water
currents. Such a method of food catching would
mean repeated opening and closing of the shell
with repeated fanning of the smaller valve. That
would have definitely killed the animal Composita
lying above that valve. The instance, thus,
suggested that brachiopods did not and do not feed
48 Part One: Principles
with the help of lophophore thrown out the shell,
as was thought earlier. They were really suspension
feeders which depended on the current of water
created by hair-like cilia associated with the
lophophore. It did not require any major movement
of the valves or the lophophore, under which
condition both the host and the included
brachiopod individuals could live peacefully
without interference [see Raup and Stanley (1971)
for figure and more details].
3.3.2.2 On the evidence of homologous
structures: Similar morphological features of
two different organisms having the same origin are
homologous structures. Function of such a structure
in an extinct organism may thus be inferred from
its homologous counterpart in an extant organism,
which can be directly observed and studied.
Ammonoids (an extinct cephalopod group) have
phragmocone chambered by septa and siphuncle
running across the septa from apical to apertural, or
the last, chamber. Phragmocone, septa and siphuncle
in Nautilus, a living cephalopod, are homologous
structures. So, it is inferred that the same structures
in ammonoids performed similar functions that
they perform in Nautilus, and, thus, helped
ammonoids live a nektic, predatory life in the sea.
3.3.2.3 Functional morphology: As mentioned
earlier, stable and permanent morphological
features are adaptive. Hence, the same
morphological characteristics in two different
organisms suggest similar adaptations and from
that their similar functions or utility. This is the
basis of functional morphological interpretation,
a much used method in palaeoecology. It attempts
at analysis of morphological features of fossils with
a view to interpreting their role in life activities,
how were they used and for what purpose and
thereby, how did they help in adaptation of the
organisms. From here conclusions may be drawn
about the mode of living, locomotory, feeding,
protective, reproductive and other habits and
behaviours and relationship with environment,
particularly the biological components.
Similar morphological features arise from
homology having the same origin (already
discussed above) or by homoplasy. In the latter
case, similar morphology develops in two different
unrelated organisms because of their performing
similar functions, but originating through different
physiological and ontogenetic processes. Thus,
wings of birds, bats and insects perform the same
function and to that extent are similar in
morphology, though they originate in different
ways. These are analogous structures.
So, both homologous and analogous structures
are useful in functional morphological analysis. To
interpret functions of such structures, it may be
necessary to take principles of mechanics,
hydrostatics and hydrodynamics to frame models
or paradigms. Of them, the model or paradigm
which provides the simplest and best explanation
and matches best with the physiology, anatomy or
genetics of the concerned organism, is accepted
for decision.
Septa and suture of cephalopods provide an
example. A paradigm chooses a particular function
or activity for a structure or morphological feature
and determines, theoretically, or with the help of
mechanical models or even computer simulation,
what should be the morphology to perform the
function most efficiently. Cephalopods require
smooth movement through the fluid medium of sea
water, horizontal or vertical. Streamlined,
bilaterally symmetrical body and shell of these
animals help in such movements. But particularly
during their vertical movement through water in
search of food, they meet with significant differences
in hydrostatic pressure. Such variation in pressure
may cause the shell to implode as they move to
depths. A paradigm which assumes regularly spaced
septa to provide a mechanical support to the shell
of a minimum yet optimum thickness, appears the
most plausible for interpreting functions of septa.
Sutures are the traces or the lines of contact of largely
concave septa on the inner surface of the shell. They
vary from smooth and straight in orthoceratoids to
wavy in nautiloids and a few earlier ammonoid
 Chapter 3 Palaeoecology 49

groups (goniatitines) to finally complex, frilled
pattern in ceratitines, ammonitines and a few other
later groups of ammonoids. Obviously for a
particular size of the shell, a frilled line of contact
or suture is greater in length than a straight line of
contact. In result, it provides greater strength to
the wall than a straight line of contact or suture. It
means that the evolution of septa in cephalopods
(particularly in the extinct group of ammonoids)
from simple to complex may have adaptative
bearing towards more efficient strengthening of
shells in the face of variation in hydrostatic pressure
at different depths, pointing to better adaptation to
swimming habit.
Functional morphological analysis has turned
out to be one of the most useful modern method
for palaeoecological, rather palaeontological
studies. It must be understood at the same time
that unique solution may not be arrived at in many
cases. One of the main reasons for that is the
limitation in applying the principle of actualism,
as discussed in Section 3.3.1. All traces of softer
fleshy parts of the body, particularly of extinct
organisms, may not be available on the skeletal
harder parts. Hence, it may not be possible to bring
out all the details of how the hard parts were related
to the soft parts, where and how they were linked
to the latter, and so on. Naturally that would tell
upon drawing conclusion about the full functional
singificance of the concerned morphological
features. Nevertheless, even within the present
limit of knowledge, it is generally possible to
understand to a considerable extent, the adaptive
significance of morphological features of even
extinct organisms with the help of analogous
structures. Analysis on homologous structures, in
turn, may help in augmenting our knowledge about
the phylogenetic relationship between the
concerned groups as also in determining the course
of evolutionary changes followed in their
phylogeny (for instance between extinct
ammonoids and extant nautiloids in evolution of
cephalopods). Factsheet 3.8 provides an example
of functional morphological studies.
3.3.2.4 Palichnological evidences: Ichno-
fossils or trace fossils may also provide important
information about the behaviour, habit and
environment of living of the organisms that made
them. A number of benthic animals make burrows
in soft sediments often for different purposes. Not
only the burrows made by different organisms are
different in shape, size, orientation, etc., those
made by the same organism for different purposes
are also different. It is often difficult to make out
which organism made which burrow-type; in that
case different types of burrows are recognized as
‘form-genera’ on their morphology only.
Observations on present day examples reveal that
burrow types vary at different depths of the basin
under varying conditions of waves and currents.
Their presence in rock-records may thus indicate
such respective conditions under which they
might have formed.

FACTSHEET 3.8
Functional Morphology of Hallucigenia: A Case Study

Hallucigenia (Conway 1977) is a Cambrian fossil from the Burgess Shale of British Columbia.
It was a soft bodied animal, whose like is not found in the present-day. The body flattened during fossilization
had a blob at one end, interpreted as head, and a long tentacle at the other, presumably a tail. On one side there
was a double row of long spike-like appendages and on the other a single row of slightly curved tentacles.
The spikes were paired and were thus considered as limbs for locomotion; single row of tentacles was thought to
represent some protective or food-catching device.
Similar fossils later found from China (Bengston 1991) showed that even the tentacles were paired. So, these
were interpreted as limbs and the thorny spikes protective device and the animal was, thereby, turned rather
upside down.
50 Part One: Principles
Since the morphology and anatomy of
vertebrates are better comprehended from their
recent counterparts or kins, functional
morphological analysis of these animals are easier
and more precise. Thus, position and depth of
different parts of footprints of dinosaurs, for
instance the genus Iguanodon of Cretaceous period
may tell us about whether they were made by the
organism in a running, walking or resting
condition. Traces of its big tail associated with the
footprints will also attest to their stance as the
present-day kangaroos that use the two hind legs
and the tail to rest upon while standing.
Of many methods now used in palaeoecology,
the above mentioned ones are more common.
Different other methods may be mentioned in
course of other discussions along with more
examples.
3.4 Appendix: Marine and
Terrestrial Environments
Contrasted
As an appendix to this discussion on palaeoecology,
the following section brings out a few vital aspects
of marine and terrestrial environments contrasted
(also see Factsheet 3.9). These two being the most
basic types of environments for the organic world,
their attributes throw much light on the pattern of
biotic distribution. Life abounds in shallow seas.
In fact, marine environment is characterized by
(i) approximately 300 times more space for life than
that on land and freshwater combined;
(ii) larger volume of biomass; (iii) an ambience with
readily available water, a fundamental constituent
of living organisms and a universal solvent with
the ability to dissolve many substances; and (iv)
less drastic variation of temperature than in air.
These may be the clues to explain greater
abundance of life in the shallow seas.
However, life is more varied on land, as
environmental variation is greater with larger
variety of habitats on land than in seas. This
variation on land is largely horizontal, caused
primarily by precipitation (controlling availability
of water), temperature (an interaction of latitudinal
climatic belt, relief and other factors), etc.
Environmental variation in seas is, on the other
hand, largely vertical, caused by (i) lesser amount
of nutrients (e.g., nitrates and phosphates from
decay of dead organisms) than in soil, as nutrients
sink to depths, out of use for plants in the photic
zone. These are brought upwards only by physical
movement of water (swelling); (ii) plants, the
primary producers of food, are limited by light
penetrating to a few metres in turbid water and a
few hundred metres in clear water, as only
50 per cent of the solar radiation penetrates the
sea surface and disappears rapidly with depth;
(iii) conditions becoming broadly constant and
uniform with depth with temperature coming down
to between 2°C and 4°C and hydrostatic pressure
increasing. Surface variations in seas are caused
by exchange of gases (with atmosphere), variations
in temperature and salinity and turbulence from
winds, these horizontal variation being set by
physical and chemical differences in sea water and
the atmosphere.
In addtion to these present-day differences
between environments on land and in seas, a few
other facts demand attention. There are more phyla
of animals now living in oceans than in freshwater
or on land. But majority of described animal
species are non-marine (Lalli and Parsons 1997).
It attests to the fact that the differences between
oceanic and terrestrial environments were broadly
the same in the geological past too.
Secondly, life originated in seas, with all
known phyla, extinct and extant, also making their
appearances there. Freshwater and terrestrial
environments were invaded by a few of the phyla
only in later times. On account of its larger solute
content, seawater has higher density than air. Thus,
it exerts greater thrust upwards than air does. This,
in turn, requires lesser energy to be spent in
floating/swimming in sea water than required for
walking/flying through air. So, marine organisms
 Chapter 3 Palaeoecology 51

do not need to store large amounts of energy in
skeleton to overcome effects of gravity and thus
have smaller size or less massive skeletons with
hollow bones. Terrestrial ones, on the other hand,
have larger and stronger skeletons to stand or to
move against gravity. It is probable that first forms
of life and all phyla may have preferred
environment where buoyancy permitted greater
energy conservation and lesser need for
development of skeletal parts.

FACTSHEET 3.9
Marine Versus Terrestrial: An Introduction

Life abounds in shallow seas

Marine environment is characterized by:
(i) approximately 300 times more space for life than that for those on land and freshwater combined;
(ii) larger volume of biomass;
(iii) an ambience with readily available water, a fundamental constituent of living organisms and a universal
solvent with the ability to dissolve many substances;
(iv) less drastic variation of temperature than in air.
These may be the clues to explain greater abundance of life in the shallow seas.
Life originated in seas
First life and all known phyla (extinct and extant) originated in sea; some of the latter migrated later into
freshwater and terrestrial environments.
Seawater has higher density than air.
Seawater exerts greater thrust upwards than air do.
Lesser energy spent in floating/swimming in seawater than in walking/flying through air.
Marine organisms do not need to store large amounts of energy in skeleton to overcome effects of gravity.
Marine organisms have smaller size or less massive skeletons with hollow bones.
Terrestrial ones have larger and stronger skeletons to stand or to move against gravity.
First forms of life and all phyla may have preferred environment where buoyancy permitted greater
energy conservation and lesser need of development of skeletal parts.
Life varies more on land
Environmental variation greater with larger variety of habitats on land than in seas,
Variation on land largely horizontal, caused by precipitation (controlling availability of water), temperature
(an interaction of latitudinal climatic belt and relief), etc.
Environmental variation in seas, largely vertical, caused by
Lesser nutrients (nitrates and phosphates from decay of dead organisms) than in soil, as nutrients sink out of
use of plants in the photic zone to be brought upwards only by physical movement of water (swelling/
upwelling);
Plants, the primary producers of food, being limited by light penetrating to a few metres in turbid water and
a few hundred metres in clear water, as only 50% of the solar radiation penetrates the sea surface and
disappears rapidly with depth;
Conditions becoming broadly constant and uniform with depth; temperature down to between 2 and 4°C;
hydrostatic pressure increasing
Surface variations are caused by exchange of gases (with atmosphere), variations in temperature and salinity
and turbulence from winds; horizontal variation being set by physical and chemical differences in sea water.
Thus, more phyla of animals now in oceans than in freshwater or on land.
But majority of described animal species are non-marine.
 4
Systematics and
Biostratigraphy
4.1 Systematics: Introduction
The world is so full of such different varieties
of things that it can be extremely, even
hopelessly, confusing. If each of the many things
in the world were taken as distinct and unique,
a thing in itself unrelated to any other, the
perception of the world would have disintegrated
into complete meaninglessness. If each tree as
considered as a wholly separate entity, there
would not be any tree at all, for ‘tree’ is a
collective concept not applicable exclusively to
a single object, considered without any
relationship to others. Similarly, it would be
equally confusing if each thing is designated by
a separate word, a separate name, without
indicating its realtionship to others.
In man’s endeavour to know about the organic
world, present and past, netither it is nor it was,
the objective to know each individual organism
separately, discreetly and independently. Rather,
what is attempted is to know about the kinds of
organisms that occur in nature, independent of
man’s perception and knowledge about them.
These kinds may be at different levels, species,
52
family or phylum; but at any level, a kind includes
organisms that are naturally related by some or
other kind of relationships among themselves and
the different kinds, in their turn, are unrelated and
differ from each other in their relationships.
Thus, when man attempts consciously and
scientifically to comprehend the huge organic
world, he wants to find out these kinds, what are
they, how many of them are there, how do they
differ among themselves and how the members of
a kind relate themselves to each other, etc.
At the outset he takes to aggregating organisms
on the basis of generalized characteristics and
relationships, those which define the groups. This
is the process of classifiying things, without which
no amount of in-depth study in whichever aspect
it may be, is possible.
Systematics is this total scientific study of the
kinds and diversity of organisms, including any
and all the relationships that define them. As
mentioned, it is the unavoidable initial step of any
scientific study of organisms. Yet, systematics
eventually gathers together, utilizes, summarizes
and implements all that is known about organisms,
with a view to arriving at the truthful conclusion
 Chapter 4
about the groups or kinds of organisms as they
occur or occurred in nature.
Classification, included in systematics, is an
operation along with its results which arranges
organisms into orderly groups, approximating
those of nature. Once erected, it provides a means
of reference to the groups so framed.
Taxonomy lays out the bases, principles,
procedures and rules of classifications themselves.
Some authors use systematics and taxonomy as
alternatives for the science of classification
(Clarkson 1998).
Nomenclature, an essential component of this
whole exercise, is the naming of different groups,
to help and facilitate all future references.
Thus, a kind of organism with spines on skins
and an internal rigid skeleton (and other features)
may be found to have two smaller kinds at the next
lower level of classification. One of them has five
equal and radially symmetrical regions (let us term
it ambs) and other characters, while the other has
ambs and associated features differentiated to
define a bilateral symmetry. For any reference to
an organism with spines on skins and an internal
rigid skeleton and radially disposed five ambs, we
need a name which really symbolizes the set of
characters that define the group. We call the
organism with spines on skins and an internal rigid
skeleton ‘echinoid’ and its two subdivisions, radial
and bilateral, ‘Regularia’ and ‘Irregularia’,
respectively. ‘Echinoid’ grouped into ‘Regularia’
and ‘Irregularia’ is the scheme of classification.
Choosing ambs, symmetry and such characters for
defining Regularia and Irregularia comes under the
purview of taxonomy. The names make it
convenient for us to recognize whether a new find
is a regular or an irregular echinoid.
Natural relationships among organisms is the
ultimate basis for their classification. But since
that is to be judged, the process of classifying
organisms starts with observing and working on
different kinds of tangible relationships. Thus,
there may be different bases for classification and
for that matter many different classifications.
Systematics and Biostratigraphy 53
In this regard, a few considerations are essential.
These are as follows:
1. A classification proposes subdivisions,
classes, groups or sets at successively lower
levels about which we can make
generalizations.
2. The subdivisions are constructed in
connection with some particular purpose; the
kinds of generalizations considered pertinent
depends on this purpose. When the purpose
is to study ecology, generalizations required
about the organisms are on habit and habitats;
subdivisions arrived at may be benthic, nektic
or planktic.
3. There should be one form of classification for
all organisms for the purpose of nomenclature.
Scientists agreed to adopt Linnean hierarchy
as this form.
4. The most widely meaningful basis of that
classification would be natural relationships
among the organisms themselves. Hence, the
classification is ultimately to be based on
evolutionary relationships.
5. To begin with, the most intuitive approach
relies on grouping on similarity of
morphology, at the first hand.
6. When it is established that the morphological
attributes, the phenotypes, represent genetic
characteristics, the genotypes, their
similarities are considered as measures of
evolutionary closeness. This involves an all-
embracing study of organisms in relation to
space and time, its environment, both physico-
chemical and biological.
With increasing knowledge and difference in
judgement and opinions, the biological contents of
a taxon, i.e., knowledge and understanding about
its biological characteristics (for fossils, mainly
morphological at the beginning) may frequently and
inevitably change. So, the problem that may crop
up is whether a taxon, a subdivision at some level
of classification, is really the same as one to which
a name was originally given. Herein comes the
54 Part One: Principles

concept of ‘type’. The type of a species is an
individual specimen or a few such; for a genus, the
type is a species. This ‘type’ is regarded as the
typical representative of the taxon. Since
morphology is the basis, the classification thus
arrived at (having been based on type and mor-
phology) is know as ‘typomorphic’ (also called
typological).
4.2 Types
Types or type-specimens are the actual specimens
that the author of a species used while founding
his species. It serves as the reference for all future
studies of that species by the same author or others.
Types are preserved in museums, to make
available for future work.
Different kinds of types are defined in
Factsheet 4.1.
4.3 Naming Species
Before we enter into the discussion of this section,
a few terms may need some introduction. In
biology as well as in palaeontology, Hierarchy is
a much used term. It refers to a systematic
framework for classification with a sequence of
groupings (or sets) at successive levels, in which
each group except the lowest one includes one or
more subordinate groups. Linnean hierarchy is the
one in vogue (Factsheet 4.2). A taxon is a set of
real organisms recognized as a formal unit at any
level of such a hierarchic classification. Thus,
Gastropoda is a taxon, a set of organisms that are
recognized as belonging to a class of the phylum
Mollusca. It is a taxonomic category at the level
of the class.The rank of this category signifies its
position in the hierarchy. Thus, it is included in
the phylum Mollusca at its next higher rank, is
equivalent (in rank) to Bivalvia, another class of
the same phylum and includes Prosogastropoda,
Pulmonata, etc. which are taxa of the next lower
rank.
The scientific system of naming kinds of plants
and animals revolves around the species level. It
is governed by a set of rules of nomenclature
agreed upon by the international community of
scientists. The rules entail as follows :
1. When an author is reporting a genus or species
from his collection, he provides proper
description and illustrations (photographs
and/or drawings).
2. The author then compares his description with
those from earlier monographs or other pub-

FACTSHEET 4.1
‘Types’ of Species

Holotype: Single specimen described by the original author in the original description
Paratype: One or a few specimens described by the original author as support to holotype in the original
description
Syntype: A few specimens described by the original author as original type specimens, in case there is no
holotype
Lectotype: Single specimen described by the original/later author; equivalent to holotype, but erected later
Neotype: One or a few specimens described by the original/later author as replacement of original types
Plesiotype: One or a few specimens described by the original/later author for founding new species
Topotype: Any number of specimens from the same locality and zone as the holotype or syntypes
From syntypes, on revision, there may be
= => One single specimen as Lectotype of original species.
= = = = > Single specimen as Holotype of later species; there may be several such.
Genotype is the type species of a genus; there may be geno-holotype, geno-syntypes, genolectotypes.
 Chapter 4 Systematics and Biostratigraphy 55

3. The new name must be in accordance with

FACTSHEET 4.2
the Linnean Binomial or Binominal System
Linnean Hierarchy of Nomenclature. For example, Spiroclypeus
ranjanae Tewari 1956 is a species name
Kingdom (a larger benthic foraminifera), where the first
Phylum part Spiroclypeus is the name of the genus
Class and ranjanae is the trivial name. The name
(Cohort) was proposed for that fossil by Tewari in 1956.
Order In names like Homo sapiens sapiens, the third
Family part represents the sapiens variety of the
(Tribe) species Homo sapiens. In all references to any
Genus such name of a genus or a species in a text, it
Species should be printed in distinctly different type-
Variety fonts, normally italics. Genus name begins
with a capital letter.
NB: Prefix ‘super’- is used for a category above some
4. Rule (Law) of Priority is a basic rule of
other, but below the next higher category: superfamily
is above family, but below class. Likewise, subfamily nomenclature. By this, other things being equal,
is below family, but above genus. Less used categories the earliest name given to any genus or species
are put within parentheses. Population and then is retained; synomyms, later cropped up, are
Individual come below variety rejected. Homonyms are normally supressed in
favour of the earliest name (see Factsheet 4.3).
lications; if his species or genus proves to be 5. The new species must be assigned to an
identical with an earlier one, it is identified existing genus, or none found suitable, to a
as the latter. If not, it is a new report. This genus to be created along with. The new
entire material must be published in an species or genus will then be marked as
acknowledged and well-circulated scientific sp.nov. or gen.nov, respectively to be written
journal. after the name suggested for it.

FACTSHEET 4.3
Synonym-Homonym
Synonym Homonym
Two names of the same thing Same name for two things
Absolute synonyms: Generic:
Two authors founded two genera Avalonia Walcott 1889 (a trilobite)
on the same genotype Avalonia Seeley 1898 (a reptile)*
Subjective synonyms: Specific:
Two genera on different genotypes Spirifer pinguis Sowerby 1820 (Carboniferous)
judged to be a single genus S. pinguis Zeiten 1838 (Jurassic)*
Absolute synonyms are suppressed Homonym is generally
in favour of the earliest name suppressed (marked * as above)
Subjective synonyms may remain dormant
till the generic name is redefined
NB: Examples acquired from Morley Davies (1949)
56 Part One: Principles

With more elaborate knowledge, a species may
be transferred from its original genus to another.
The following is an example:
Stage I: Two species are there, viz. Cardita
deltoidea Sowerby 1818 and Venericardia
deltoidea Sowerby 1820.
Stage II: Species are revised–C.deltoidea to
Pholadomya deltoidea (J. Sowerby) (species to a
new genus Pholadomya by Sowerby) and V.
deltoidea to C. deltoidea (J. Sowerby) (as Wood
transfers the species to Cardita). Both the second
names are retained.
When an original genus is divided into several
genera, the original name must be retained for one
of them, preferably for the species the original
author considers as the type species of the genus.
For this, the original author should name the type
species or genotype in his original description.
Stage I: Dalmon raises genus Orthis.
Stage II: Hall and Clarke raise subgenera
one Orthis s.s. (including Dalmon’s genotype) and
all other with new names.
Stage III: Subsequently the subgenera may
be raised to genera, viz. O. (Dalmanella)
elegantula Dalmon to Dalmanella elegantula
(Dalmon); O. (Schizophoria) resupinata (Martin)
to Schizophoria resupinata (Martin); O. (Orthis)
callactis Dalmon to Orthis callactis Dalmon.
A later author may unite several genera into
one; the earliest of the names of the genera being
united should be retained.
A few more examples will illustrate some more
details.
1. In 1776 O. F. Muller described the genus,
Terebratula.
2. In 1820 E.F. von Schlotheim described a new
species from Devonian of North Germany
naming it Terebratula sarcinulatus.
3. In 1830, with more work and knowledge on
brachiopods, G. Fischer de Waldheim revised
the species as type species of Chonetes
sarcinulatus (Schlotheim); that the species
was originally described under a different
genus is known from Schlotheim’s, i.e. the
original author’s name in parentheses.
4. In 1917, F.R. Cowper Reed recognized a new
species of the same genus, different enough

FACTSHEET 4.4
Systematic Position of Dinosaurs, Modern Man, Horses and Elephants
(Following Phylogenetic Systematics Scheme for Vertebrates, Benton 2005)

All included in phylum, Chordata, subphylum Vertebrata

Series Amniota
Class Sauropsida Mammalia
Subclass Diapsida Cohort Placentalia (Eutheria)
Infraclass Archosauromorpha
Division Archosauria
Subdivision Avemetatarsalia
Superorder Dinosauria
Order Saurischia Ornithischia Primates Perissodactyla Proboscidea
Suborder Anthropoidea ... Elephantiformes
Infraorder Catarrhini ...
Superfamily Hominoidea Hippomorpha ...
Family Hominidae Equidae Elephantidae
Genus Homo: Species sapiens: Variety sapiens (modern man)
Genus Equus (horse, ass, zebra, etc.)
Genus Elephas (Asiatic elephant) Loxodonta (African elephant)
 Chapter 4 Systematics and Biostratigraphy 57

to be placed in a new subgenus, Siju Limestone of Assam (along with the

Chonetes(Eochonetes) advena Reed 1917.
5. Later Eochonetes was raised to the rank of a
genus; two subspecies of Eochonetes advena
were written as Eochonetes advena advena
Reed 1917 (recognizing the original material
of Reed) and as E. advena Reed 1917 gracilis
sp.nov. by Harper in 1989.
Subsequently the second species will be
referred as E. advena gracilis Harper 1989.
In the case species identification is not certain,
the following processes are taken to as required or
suitable.
1. Monograptus cf. vomerinus means species
that may be compared with Monograptus
vomerinus. Use of aff. in place of cf. means
‘related to’ instead of ‘comparable to’.
Nummulites sp. would signify a species of
Nummulites, which can neither be matched
with any earlier species nor can be designated
as a new one for want of proper material. Use
of ‘?’ would also indicate doubt about
identification and naming.
2. For many species (ones that vary in size and/
or colour or other characters over geography),
subspecies or varietal names are used for
distinctive geographic forms. Assilina
regularia sijuensis is a variety of the species
Assilina regularia, described from Eocene
species itself).
The examples are acquired from Morley
Davies (1949) and Clarkson (1998).
4.4 Species Concept and Species
Problem
4.4.1 Dual problem
As discussed above, systematics involves
classifying organisms in a hierarchical sequence
of taxonomic categories on this or that basis, and
naming the taxa, particularly species, governed by
often intricate rules of nomenclature. However,
systematics is not just a drab branch of
palaeontology or biology, involving the above-
mentioned routine exercises. At its core, it rests
on philosophy, an outlook of how we look at the
objects under study. Since science has developed
through ages with more and more observations and
their rational explanations, the outlook with which
palaeontologists (or, for that matter, scientists) have
tried to classify the organic world has also
undergone changes. In honest pursuit of truth,
scientists have tried to reach at the most rational,
objective scheme or method. Limitations of
knowledge prevailing in the real objective situation
of the society, scientific or broader, have ultimately

FACTSHEET 4.5
Species: Dual Usage (Following Mayr 1996)

1. The species taxon: The word taxon refers to a concrete zoological or botanical object consisting of a
classifiable population (or group of populations) of organisms. The house sparrow (Passer domesticus), the
potato (Solanum tubersum) or the modern man (Homo sapiens sapiens) are species taxa. Species taxa being
concrete and particulars, can be described, compared with and delimited against other species taxa.
2. The species category: The word ‘species’ also indicates the rank in the Linnean hierarchy. The species
category is an array, i.e. the group that contains all taxa of species rank. It articulates the concept of the
biological species and is defined by the species definition. The principal use of the species definition is to
help take decision on the ranking of species level populations. “Is it a full species or a subspecies”? The
answer to this question has to be based on inference (Mayr and Ashlock 1991, 100-105). A complication is
produced by the fact that in the Linnaean hierarchy asexual ‘species’ are also ranked in the species category,
even though they do not represent the Biological Species Concept.
58 Part One: Principles
shaped the product. It is not possible to trace out
the whole history in all details in the purview of a
brief chapter. Only some essential features will be
highlighted to bring home some concepts useful
for students.
Species concept and species problem have
intrigued scientists for long. In essence, both hinge
upon how we look at species. In fact, there cannot
be an universally applicable definition of species
for all known organisms. They may be relatively
distinct and stable, as in birds and mammals, or
relatively indistinct and unstable, as in the hybrid
complexes common in micro-organisms, plants,
some crustaceans, amphibians and fish. Long since,
scientists recognized that species is the working
unit of any scientific study of organisms or their
fossils; the latter cannot be named by individuals.
As discussed in the beginning of the chapter, the
whole process becomes chaotic and confusing
then. So to name organisms, we named species.
The meaning of species is, however, beset with
controversy as to how it should be understood. This
gives rise to the species problem (see Mayr 1957
for a history). In essence, there are two different
sets of species problems, one being the problem of
how to define the species (what species concept to
adopt), and the other being how to apply this
concept in demarcation of species taxa.
What a biologist actually encounters in nature
are, however, populations of organisms. The
organisms vary considerably in size, ranging from
local demes (the community of potentially
interbreeding individuals at a locality) to the
species taxon. The biologist assigns these
populations to species. It requires two operations
that correspond to the two facets of the species
problem stated above.
4.4.2 Typological species concept
To delve further into the issue, we need a
digression. Ancient Greeks looked at a fossil as
‘half-life’, which bore the appearance of a living
form, but lacked the vital force of life. It was not
bestowed with that vitality. It was, so they
thought, on account of an omission on the part of
the creator. So it remained fixed, unchanging in
its characters, just as other objects in the
inanimate world around man remained. To know
about it, to identify and recognize it, we must look
at the essential characters of its form and
appearance that distinguishes it from other fossils.
There was no organic evolution known at that
time, there was no earthly process known as to
how organisms, at levels higher than individuals,
could have arisen.
A long period in the history of science, from
Plato and Aristotle until Carl Gustav Linnaeus (he
wrote his Systema naturae in 1778) or even later,
was reigned by this kind of ideas about organisms.
Of course, there were advancements. Among
others, the hierarchy Linnaeus proposed still serves
as the base for our studies (with additions of many
levels, though); the system of Nomenclature he
propounded is still the system scientists adhere to.
But neither the taxonomists nor the philosophers
looked at species in a way different from that in
which they would have considered inanimate
objects. They defined species, may be, differently.
Nevertheless, they all meant at heart that each thing
bore some essential properties. We must bring them
out with the help of ‘intellectual intuition’ and
must define the things on the basis of those
‘essence’. Thus, all members of a species, or for
that matter any taxon, bear and reflect the same
essential nature. In other words, they have a typical
commonality, a type of ‘essence’. Thus, came the
philosophy of ‘essentialists’ and the idea or the
concept of species or classification based on it,
known as typological concept.
The word ‘species’ conveyed the idea of a
class or array of objects, members of which shared
certain defining properties. This definition
distinguished a species from all others. Such a
group is constant, with fixed defining characters;
it does not change in time; all deviations from the
definition of the array, all variations are trivial,
 Chapter 4
irrelevant and merely ‘accidents’, that is, imperfect
manifestations of the essence, the characteristics
of the species.
Upto the 19th century this was the traditional
and the most practical species concept in biology
or palaeontology. According to this concept, a
species comprises a set of populations consisting
of individuals that are typified by common
morphological characteristics different from those
of other species. These morphological
characteristics may be best represented in totality
or in part in a single specimen; it then stands out
as the ‘holotype’ of the species; otherwise, the same
requirement is fulfilled by a number of specimens
(paratypes, syntypes, etc.), all accepted as typical
or types. The species is identified and named on
the strength of these types. Subsequently any
specimen or individual is recognized as the same
species only on the basis of whether it matches
with the types or not. This is definitely a convenient
exercise, particularly with fossils. In the latter,
morphology is the immediate and most evident
criterion to work upon, whether in cataloguing of
species taxa and their arrangement in keys and in
collections. The concept was usually referred to
as the typomorphic or typological species
concept, the Linnaean concept of species, much
prevalent in palaeontology.
But the concept has serious flaws. The most
important is that morphological characters are not
infallible criteria for recognition of species. Firstly,
individuals belonging to a particular species are
never morphologically exactly alike. There are
often numerous different morphological types
within a species as they occur in nature as distinct
biological entities. It may be either due to
individual genetic variation or due to different life
history categories (males, females, immatures)
which are morphologically far more different from
each other than are the corresponding
morphological types in different species. Besides,
differences as manifested in widely different
human races of the same species of the modern
man are more cases in point. On the other hand,
Systematics and Biostratigraphy 59
there are species that differ morphologically only
very slightly or even none at all (sibling species).
Defining a species by a fixed type ignores these
variations, however small they may be.
Secondly, morphology of a species also varies
with time. This is because the species evolves; it
is never alienated from the process of evolution,
which links organisms genetically. So any view on
species must be cast in genetical terms, if it is to
be useful in understanding the process of
evolution. Furthermore, similar morphology may
result from convergence or parallel evolution on
account of similar adaptation, the former in case
of unrelated organisms, latter with related forms
(see Factsheet 4.6). Grouping individuals into a
species solely on morphology may lead to
artificially lumping them together ignoring the
natural genetical differences among them. Thus,
simple morphological difference between two
species fails to shed any light on the true
biological significance of species.
Essentialists’ approach hinged upon
‘intuition’, a subjective element of the person
concerned. With that only, he decides what are the
essence of the species he is studying. To stand
against this subjectivism, there arose a second
group of scientists, who defined species as an array
of individuals so nearly alike (in morphology) that
they may conveniently be denoted by the same
name. Nominalists as they are, with them species
do not exist as natural groupings. They are the
products of human mind. Man attaches a name to
a group of individuals he considers to be similar.
At the same time, it is he, who decides the degree
of similarities, giving equal weight to all the
characters for the sake of objectivity. But, as
mentioned morphological similarities may arise
from convergence or parallel evolution or other
evolutionary, genetic or developmental phenomena
and, thus, cannot be of equal importance or
significance. Besides, a species for instance, the
human species is not an entity made by the
taxonomist, who gave the name. It is a natural
entity, a product of evolution.
60 Part One: Principles

FACTSHEET 4.6
Morphological Similarities (Due to Inheritance from a Common Ancestry)

Yes, due to inheritance No, not due to inheritance

HOMOLOGY HOMOPLASY
Y From common ancestry Y N N N N
Y Acquired due to evolution along similar trends Y N N N N
Y/N Acquired due to adaptive pressure N Y Y N N
Y/N Within same community N N Y N N
N Acquired due to functional similarity N N N Y N
N Acquired due to independent causes N N N N Y
Y Within same lineage N N N N N/Y
1 2 3 4 5

1. Parallelism: Development of similar characters separately in two or more lineages of common ancestry
and on the basis of, or channelled by, characteristics of that ancestry.
2. Convergence: Development of similar characters separately in two or more lineages without a common
ancestry, involving adaptation to similar ecological status.
3. Mimicry: Similarity adaptive as such and not related to community of descent.
4. Analogy: Functional similarity not related to community of ancestry.
5. Chance: Resemblance in characteristics developed in separate taxa by independent causes and without
causal relationship involving the similarity as such. Y: yes N: no.

4.4.3 Biological species concept
In fact, after organic evolution was known to man,
it was also found that species was the principal
unit of evolution. It is impossible to comprehend
evolution, and indeed almost any aspect of biology,
without having a sound understanding of species.
Increasing knowledge of evolution also brought
out clarity on the significance of morphology and
its variation in time and space. This further helped
understand the deficiencies of the typological
species concept. It led to the emergence of the
Biological Species Concept (BSC) developed in
the second half of the 19th century particularly
among zoologists.
The concept holds:
Species are groups of actually or potentially
interbreeding natural populations that are
reproductively isolated from other such groups.
In regard to speciation it states:
A new species develops if a population which
has become geographically isolated from its
parental species acquires during this period of
isolation, characters which promote or guarantee
reproductive isolation when the external barriers
break down. (Mayr 1942).
Buffon was perhaps the first among the
scientists who foreshadowed a change. Darwin, too,
had unquestionably adopted a biological species
concept in the 1830s in his transmutation notebook,
even though later he largely gave it up (Mayr 1992).
Besides, many of the biologists who accepted or
accept the morphological species concept did or
do base their decisions ultimately on the
reproductive community principle of the BSC. They
combine drastically different phenotypes into a
single species because they have observed that they
were produced by the same gene pool. On the other
hand, the biological concept uses the degree of
morphological difference as an indication of the
underlying degree of reproductive isolation.
There are other details too. Generally, species
have a geographical (space) and a temporal (time)
extension. In that case, there are local or temporally
 Chapter 4 Systematics and Biostratigraphy 61

circumscribed (i.e. of limited time range)
populations which may differ slightly from each
other. Such populations, when they are considered
to be conspecific, are combined into a polytypic
species. The major problem is to decide which local
populations to combine into a polytypic species.
The decision is based on inference, so it is always
somewhat uncertain. When the typomorphic
species concept was dominant, almost any isolated
population that differed by a morphological
character was called a different species. Since the
rise of the biological species concept, the question
is always asked whether or not such a population
would interbreed with other populations differing
in space or time, if they would meet in nature.
Conspecific populations that differ from each
other morphologically are called subspecies. If
such subspecies are part of a series of contiguous
populations, they are a purely taxonomic device.
However, they are incipient species if such
subspecies are geographically isolated. They may
in due time acquire the needed isolating
mechanisms to function as well-separated species.
It is apparent that the definition of the
biological species must be based on its biological
significance, which is the maintenance of the
integrity of well-balanced, harmonious gene pools.
In actuality, species taxa are demarcated using
morphological, geographical, ecological,
behavioural and molecular information, and not
always the actually or potentially interbreeding
character to infer the rank of isolated populations.
Furthermore, particularly for palaeontologists the
concept is rather useless for all practical purposes,
because fossils do not bear any mark of potentiality
of interbreeding as such.
Again, the biologist, rather the neontologist,
does not deal with evolutionary lineages (except
very short ones). Biological species tend to be
distinct because most belong to lineages that have
been reproductively isolated from other lineages
for a considerable time. For a palaeontologist, the
time dimension makes it impossible to apply the
commonly accepted biological species definition,
as changes within a species with its evolution
through time blurs its discreteness. These are the
aspects of species problem a palaeontologist has
to face in particular.
4.4.4 Evolutionary species concept
Solution to these uncertainties or problems with
the BSC was sought in other kinds of species
concept, using other criteria to define species.
One such was the Ecological Species Concept
(ESC), based on the niche occupation of a species.
It is not workable because of the fact that in almost
all the widespread species, there are local
populations which differ in their niche occupation.
An ecological species definition would require that
these populations be called different species even
though, on the basis of all other criteria, it is
obvious that they are not.

FACTSHEET 4.7
Cladogram of Cladistics

In the diagram, A and B, two taxa share common ancestor; they are then
A B C D sister groups. They are separated from C, on the strength of a
synapomorphy, i.e. a new character developed in course of evolution.
Taxon C is then a sister group of A and B combined. Likewise, D is the
sister group of A, B and C combined. The taxonomist assumes that a
lineage splits dichotomously; he compiles a character data matrix without
adding any weightage to the characters. Larger number of characters
increase the data base, which can be tackled by well-known computer
programmes, such as PAUP, Hennig 86, NONA, Mac Clade, etc.
62 Part One: Principles
From their observations, several authors chose
‘evolutionary potential’ as another criterion, for
they found that species were not constant but the
product of evolution and were still potentially
continuing to evolve. So, they defined species as,
for example, “a species is an evolved or evolving
genetically distinctive, reproductively isolated,
natural population” (Emerson 1945). But though
it was necessary, it was not a sufficient criterion.
Everything else in the living nature also has the
capacity to evolve. Every population, every
structure and organ is the product of evolution and
continues to evolve, genera and higher taxa evolve,
and so do faunas and floras. Most of all, the
capacity for evolving is not the crucial biological
criterion of a species, it is the protection of its gene
pool.The biological species concept omits
‘evolving’ from the species definition for this
reason. Simpson also attempted to make evolution
the basis of a species concept: “An evolutionary
species is a lineage (an ancestral-descendant
sequence of populations) evolving separately from
others and with its own unitary evolutionary role
and tendencies” (1961, 153). He replaced
reproductive isolation of the biological species
concept, a criterion not useable for palaeon-
tologists, with such terms as ‘maintains its identity
and ‘evolutionary tendencies’. But, as criticized
by the biologists like Mayr (1996), maintaining
identity was a rather vague process. It was also
not clear if it includes geographical barriers. It was
also not well-defined about what evolutionary
tendencies were and how could they be determined.
The ESC encounters three additional major
difficulties: (1) it is applicable only to monotypic
species and every geographical isolate would, by
implication, have to be treated as a different
species; (2) there are no empirical criteria by which
either evolutionary tendency or ‘historical fate’ can
be observed in a given fossil sample (Simpson
1961, 154-l60) and (3) the definition does not help
in the lower or upper demarcation of
chronospecies, even though the main reason why
the evolutionary species concept was apparently
introduced, was in order to deal with the time
dimension, not considered in the non-dimensional
biological species concept. Indeed, Simpson’s
definition is essentially an operational recipe for
the demarcation of fossil species.
Evolutionary taxonomy based on this
evolutionary species concept appears to serve a lot
for palaeontologists. They start with morphological
similarities (phenetic similarities). Yet considering
that such similarities reflect phylogenetic affinity,
they end in inferring phylogenetic relationships.
Thus, taxa erected on morphology (or such other
tangible similarities) earns the status of naturally
related life forms. Obviously, this is judged in the
background of the order of stratigraphic succession
and geographical distribution.
However, as indicated above, uncertainties and
subjectivity in judging evolutionary, i.e., genetic
which is, in other words, natural relationships led
some taxonomists to adopt a numerical taxonomic
method. Proponents of this method quantified
phenetic resemblance and held that considering a
large number of characters would minimize
subjectivity and uncertainty. But here too, the
selection of suitable mathematical–statistical
analytical method depends on the author.
Moreover, considering all characters with equal
‘weightage’ may over or underestimate taxonomic
importance of one character or the other. On the
contrary, placing weightage would bring in
subjective judgement on which character is to be
weighed high or not.
4.4.5 Phylogenetic species concept
Another species concept, rather a methodology
based on bringing out phylogenetic relationships
of species, is variously termed Phylogenetic
Species Concept (PSC) and cladistics or
phylogenetic systematics, respectively. The father
of this school is Willi Henni, a German
entomologist; ideas have, however, since changed
from what Hennig thought.
The post-Hennigian PSC, is enunciated as: A
species is a diagnosable cluster of individuals
 Chapter 4 Systematics and Biostratigraphy 63

FACTSHEET 4.8
Phylogenetic Relationships

There may be three kinds of phylogenetic relationship:

Monophyletic: Groups (also called clades) whose members have arisen from a common ancestor and includes
all descendants of that ancestor, e.g. Phylum Chordata, subphylum Vertebrata, family Equidae,
genus Hipparion, etc.
Paraphyletic: Groups which include only the most primitive descendants of a common ancestor, but exclude
some advanced descendants. Reptilia is paraphyletic, as it arises from a common ancestor, but
excludes some descendants, the birds and the mammals, which have also arisen from the
same ancestor.
Polyphyletic: A group whose members have arisen from different ancestors and whose latest common
ancestor is not included in the group; they show convergence, that is similarity of characters
on account of similar adaptation. Arthropoda, with members with segmented body and
articulated appendages, is now considered polyphyletic. A group of nektic vertebrates including
fishes, seals and whales would be polyphyletic.

within which there is a parental pattern of ancestry
and descent, beyond which there is not, and which
exhibits a pattern of phylogenetic ancestry and
descent among units of like kind. Individuals
of a cluster are linked by ‘synapomorphous
resemblances’ (i.e. shared derived characters).
According to this school, in any group of
organisms, characters (i.e. observable attributes of
organisms) or combination of characters may be
primitive (symplesiomorphic) or derived. Recency
of common origin of the groups of organisms
(species or taxa at any level) under study, that is
their closeness in phylogenetic relationship, is
revealed through shared derived characters
(synapomorphies), meaning through possessing
new characters that have appeared in course of
evolution.
One common example, easy to comprehend,
comes from vertebrates. All vertebrates have
backbone, a primitive character of them. But it is
a derived character shared by all vertebrates, when
the group is judged against invertebrates or plants.
This method known as outgroup comparison
is the key to distinguishing synapomorphies from
primitive characters. The outgroup consists of
everything that lies outside the group under study,
i.e. the ‘ingroup’. In the above example, inverte-
brates or plants make the outgroup, vertebrates the
ingroup. Homologies (see Factsheet 4.6) need to
be identified and separated from analogies. The
former are considered as ‘derived characters’.
However, it requires time and judgement to find
out and analyze primitive or derived status of
characters. The results are depicted
diagrammatically in cladograms, which reflect
phylogenetic relationships albeit without any
reference to geological time or succession of their
occurrence (Factsheet 4.7).
Phylogenetic species concept has drawn both
attention and criticism. Biologists often consider
it nothing more than the revival of a purely
morphological species concept (Mayr 1996).
Moreover, as held by the proponents of this
concept, speciation occurs when “gaps develop in
the fabric of tokogenetic relationships” (i.e. genetic
relationships among animals which arise through
the phenomenon of reproduction). In that sense,
the concept is not much different from the
evolutionary species concept of Simpson. It has
also been questioned if parthenogens and other
non-sexually reproducing species are phylogenetic
species or not.
In the post-Hennigian days PSC was
enunciated by various authors. We have already
64 Part One: Principles
cited a definition by Eldredge and Cracraft. In
reality, however, there exist vast differences not
only among different phylogenetic species
concepts, but also within individual investigators’
temporal concepts. The differences may be
summarized (following Baum and Donoghue
1995) as relating to how to define species: some
PSCs do so based on characters, whereas others
do so based on historical relationships or
ancestry. However, relatedness is determined in
all the phylogenetic schools based on examination
of characters and character states. But in the
analysis, convergence poses a problem, much
greater than generally thought of. Thus, deciding
between homologies and analogies may not just
be academic.
Further, it is argued that a cladogram is not an
evolutionary tree; it is an analysis of taxonomic
relationships and does not take into account the
succession in the rock record. A solution is sought
in combining information from cladograms with
biostratigraphic data. There is generally a good
correlation between the cladogram and the rock
FACTSHEET 4. 9
Allopatric Speciation
Allopatric speciation takes place when a new species
develops with a population which has become
geographically isolated from its parental species
acquiring during this period of isolation, characters
which promote or guarantee reproductive isolation
when the external barriers break down.
Allopatric populations need not be isolated by large
distances or formidable geographic barriers.
Particularly in organisms which do not have means
to spread over large distances, or in those organisms
which breed in limited or fixed territories (temporal
or physical), the inhabitants of adjacent biotopes
may be geographically or temporally disjunct, and
hence become allopatric. Parasites are allopatrically
distributed on their host organism, even if the
individuals constituting the host species are
sympatric.
record; in cases, newer ‘unexpected pattern’
(Clarkson 1998) may emerge.
In this connection, we may refer to Callomon
(1985), where he concluded the following:
1. In the analytical stage of taxonomy,
classification is based on single or a few
selected morphological characters and is
purely descriptive.
2. Differentiating groups on morphological
differences, instead of similarities, still makes
a classification primarily morphological.
Stratigraphical data are rarely used for the
purpose. Species and genera are thus
morphospecies and morphogenera.
3. Differences among some groups are vertically
less than those among others horizontally, i.e.
of the same age. Such vertically similar
species that is similar species of different ages
are often combined into a single genus. This
is also morphological and the taxa are
morphospecies or morphogenera.
4. In horizontal classification, all members of a
contemporaneous but morphologically diverse
FACTSHEET 4.10
Sympatric Speciation
Sympatric speciation is speciation without complete
geographic isolation. The processes leading to
divergence and eventual reproductive isolation are
diverse. It is most commonly reported in insects,
presumably due to insects’ reduced vagility,
generally reduced mobility, and highly specific
niche requirements. In general, however, sympatric
speciation has not been received with great
enthusiasm. Stasipatric model is a sympatric
chromosomal mode of speciation.
Sympatric speciation may take place in highly
different heterogeneous environment where
conditions remain constant or stable and evolutionary
processes work on a broad morphologically
(or otherwise), i.e., phenotypically and hence
genetically, variable, yet small-sized ancestral
population in which genetic drift takes place relatively
easily accelerating the processes of speciation.
 Chapter 4
assemblage are regarded as merely variants of
a single variable unit, the biospecies. It
becomes almost a rule that given large sample
size, the inferred species exhibits equally great
intraspecific variability, and morphological
similarities between such successive specific
assemblages indicates ‘probable’ ‘linear,
evolutionary relationship’. At this point,
classification enters into synthetic stage and
stratigraphical data are prerequisite. Kayal and
Bardhan (2006) apply this concept of
phylogenetic systematics to middle Jurassic
Reineckeiid ammonites of Kachchh, India, a
case study from India.
In conclusion, it may be added that BSC is
applied to the living species, where natural
interbreeding and viability or fertility to produce
offsprings may be tested. In practice, however,
morphological species concept is much more used,
in which species are recognized on the strength of
unique characters. As mentioned, most biologists
consider phylogenetic species concept as very near
to, or even revival of, morphological species
concept, where species is defined as a clade of
‘diagnosable geographic forms of the same basic
kind’ (Benton 2005). Palaeontologists, particularly
the vertebrate palaeontologists, seem to be taking
up both morphological and phylogenetic species
concepts for their work (Benton 2005).
4.5 Speciation or Origin of Species
4.5.1 Phyletic gradualism
For many years since the publication of Darwin’s
On the Origin of Species, evolution was perceived
as having proceeded in a stately fashion at a steady
rate. Phyletic gradualism is this traditional thought
with respect to rates of speciation. Phyletic
gradualism is visualized as follows (Eldredge and
Gould 1972):
New species arise by even and slow
transformations, involving large numbers, usually
Systematics and Biostratigraphy 65
the entire ancestral population into its modified
descendants, occurring over all or a large part of
the ancestral species’ geographic range.
Under such circumstances the fossil record of
a species should consist of a long graded series of
continuously intermediate forms and
morphological ‘gaps’ in a phyletic series are due
to imperfections in the fossil record.
The idea of phyletic gradualism has its
inconsistencies. Guided by this idea,
palaeontologists search for graded series, or
intermediates or ‘missing link’ among various
‘forms’ or species. Modern evolutionary theory
however answers in negative. Besides, if species
can arise solely by the slow change of preceding
forms, then a new species can only arise if its
predecessor changes to the extent that it becomes
unrecognizable as itself. In other words, a new
species can only arise when the ancestor is
extinct.
4.5.2 Punctuated equilibria
A paper titled Punctuated equilibria: an alternative
to phyletic gradualism (Eldredge and Gould, 1972)
ushered in a new idea. It held that the
“imperfections” of the fossil record were
reflections of the reality of evolution, and not gaps,
which, as Darwin envisaged, would demonstrate
continuity and steady rate of evolution, once they
were filled. Evolution, rather speciation, takes
place in not the entire, but only a small part of
populations, only large enough to permit changes
in gene frequencies due to random drift. It is
allopatric in character. These parts are presumably
peripheral isolates. As a result, their examples are
much rarer to find in the fossil record; the large,
main population would be preserved in much
higher proportions than the small isolate. As to the
rates of evolution, rather speciation, they are quite
rapid (in an estimate it is held as 100,000 years for
the origin of a species with a subsequent life span
of 10 million years), but the actual speciation
events are quite difficult to find in the geological
66 Part One: Principles

record because they are restricted to very thin layers

FACTSHEET 4.11
of strata. Many breaks in the fossil record are, thus,
real. The history of evolution is not one of stately Categories and Unit-terms in Stratigraphy
unfolding, but a story of homeostatic equilibria or
Stratigraphic Principal stratigraphic
stasis, disturbed or punctuated only ‘rarely’ categories unit-terms
(i.e. rather often in the fullness of time) by rapid
and episodic events of speciation. It is, thus, not Lithostratigraphic Group
any phyletic gradualism, but a ‘punctuated Formation
equilibria’ that characterizes the scenario. Members
After the introduction of the idea of punctuated Bed(s)
equilibria, whose proponents even considered at Biostratigraphic Biozones:
one stage that phyletic gradualism simply does not Assemblage-zones
Range-zones
occur, fossil evidences for punctuated equilibrium
(various kinds)
became almost overwhelming. Soon palae-
Acme-zones
ontologists sought for unbroken chains of Interval-zones, etc.
intermediate forms, now considered a rarity, an
Chronostratigraphic and equivalent
exception. The PSC developed on this idea totally Geochronologic units
negating phyletic gradualism. Eonothem Eon
However, scientists now agree that it is not a Erathem Era
matter of whether phyletic gradualism or System Period
punctuated equilibrium is the one and only Series Epoch
mechanism for evolutionary change, nor it is a Stage Age
matter between palaeontologists and non- Chronozone Chron
palaeontologists, the former standing for phyletic Other stratigraphic Zone (with appropriate
gradualism and the latter for punctuated categories (mineralogic, prefix)
equilibrium model. The two may really represent environmental, seismic,
extreme views on the complex process of organic magnetic, etc.)
evolution. NB: If additional ranks are needed, prefixes ‘sub’ and
‘super’ may be used with unit-terms for additional
ranks, though the International Stratigraphic
4.6 Biostratigraphy Commisssion recommends prudent and restrained
use to avoid complicating the nomenclature
unnecessarily.
4.6.1 Background material
Some background material may help introducing 4.6.2 Introducing biostratigraphy
biostratigraphy. However, for brevity’s sake, that
material is represented here in factsheets. Factsheet Stratigraphy is the branch of geology which
4.11 presents different categories units used in undertakes studies on rock successions, including
stratigraphy. Factsheet 4.12 includes some observation and description of the members of
definitions related to biostratigraphy, discussed those successions and correlation between them.
subsequently. Factsheet 4. 13 summarizes the Thus, stratigraphy, with its principles and
etymology and type locality of different periods of procedures, produces systematic and thorough
Phanerozoic Eon, and authors or authorities who study of composition, geometry, sequence, history
helped in erecting these periods. Criteria for and genesis of rocks, on which there may develop
definition are also shown. a systematic ordering of the rock record, in regard
 Chapter 4 Systematics and Biostratigraphy 67

FACTSHEET 4.12
Some Definitions (Hedberg 1976)

1. Biostratigraphy is the element of stratigraphy that deals with the remains or evidences of former life in
strata and with the organization of strata into units based on their fossil content.
2. Biostratigraphic classification is the organization of strata into units based on their fossils content.
3. Biostratigraphic unit is a body of rock strata unified by its fossil content or palaeontological character and
thus differentiated from adjacent strata. A biostratigraphic unit is present only within the limits of observed
occurrence of the particular biostratigraphic feature on which it is based.
4. Biostratigraphic zone (Biozone) is a general term for any kind of biostratigraphic unit. Biozone is a short
alternative term for biostratigraphic zone. Bio should be used in front of the term ‘zone’ to distinguish
biostratigraphic zones from other kinds of zones whenever there is any danger of confusion. This is particularly
important in preventing confusion between biozones and chronozones; both may be named from a fossil or
fossils, but they are quite different from each other in concept.

FACTSHEET 4.13
Phanerozoic Periods of Stratigraphical Column: Summary Details Including Criteria of Definition

System Name Type Locality Named or Proposed by Date Proposed Remarks: Definition Based on

Cambrian W. Wales Adam Sedgwick 1835 Defined mainly on lithology

Ordovician W. Wales Charles Lapworth 1879 Set up as an intermediate unit between the
Cambrian and Silurian to resolve boundary
defined by fossils
Silurian W. Wales Roderick I. Murchison 1835 Defined on lithology and fossils
Devonian Devonshire, Roderick I. Murchison 1840 Boundaries based mainly on
S. England and Adam Sedgwick fossils
Carboniferous C. England William Conybeare 1822 On lithologically distinctive coal-bearing
and William Philips strata, but recognizable by distinctive fossils
Mississippian Mississippi Alexander Winchell 1870 Þ The Mississippian and Pennsylvanian
Valley Ñ are subdivisions of the
ß Carboniferous; not used outside
Pennsylvanian Pennsylvania Henry S.Williams 1891 Ñ the United States
à
Permian Perm Province, Roderick I. Murchison 1841 Identified by distinctive fossils
Russia
Triassic S. Germany Frederick von Humboldt 1843 Defined lithologically on the basis of a
distinctive three-fold division of strata; also
defined on fossils
Jurassic Jura Mtns, Alexander von Humboldt 1795 Defined originally on lithology
N. Switzerland
Cretaceous Paris Basin Omalius d’Halloy 1882 Defined on the basis of strata of distinctive
chalk beds
Tertiary Italy Giovanni Arduino 1760 Originally defined on lithology; redefined
with type section in France on the basis of
distinctive fossils
Quaternary France Jules Desnoyers 1829 Defined on lithology, including some
unconsolidated sediment
68 Part One: Principles
to how they are arranged and related in time and
space framework. To the extent it leads to
interpretation and understanding of rock
successions in fuller details, as ingredients, as well
as sequences of events in the geological history of
the earth, stratigraphy was regarded as one of the
final tools for a geologist.
However, stratigraphy, as understood now,
provides methods of analysis and interpretation,
central to many fields of geological investigation,
viz. analysis of basin dynamics and evolution, etc.
There are three major classical approaches to
stratigraphy : lithostratigraphy, biostratigraphy
and chronostratigraphy. Other methods, e.g.
magnetostratigraphy, geophysical log
stratigraphy, seismic stratigraphy, developed
with subsurface studies, augment stratigraphical
studies. Holostratigraphy is the combined use of
all possible methods.
Whatever be the ramifications, strati-
graphical studies of an area or succession, starts
with observation and description of the local
succession of units, erected objectively in terms
of tangible mappable or traceable criteria.
Lithology (or such other tangible characters such
as magnetic, electrical, seismic or other
signatures in subsurface stratigraphy) serves for
the purpose. With this, the local succession is
correlated with regional stratigraphical units and
successions, and ultimately with the standard
stratigraphical column (Factsheets 1.2 and 4.13),
the global reference frame. With most
Phanerozoic rocks, this correlation is generally
done on biostratigraphy.
Biostratigraphy is the study of the relative
arrangement of strata based on their fossil content
(Matthews 1984). In other words, it is the
characterization, classification and correlation of
rock units, in one word their systematization, on
the basis of their fossil content.
In addition to the physical characteristics of
sedimentary rocks, lithology, or other physical
properties (e.g. magnetic, seismic, electrical, etc.)
that can be remotely sensed and measured, fossils
provide highly useful means of subdividing sedi-
mentary rocks or their sequences into identifiable
stratigraphic units. They further make possible
ordering and relative age-fixing of strata and their
correlation on local, regional, continental or even
global scale.
In fact, on account of their objectivity, though
rock units are the basic starting points in
stratigraphic studies of an area, their use is limited
to that small confine. Verifiable physical
correlation of rock units is limited by outcrop
patterns and availability of subsurface data, and is
thus restricted to small areas. Beyond that, rock
units are generally diachronous. Thus, a beach
sandstone prograding southward across a shallow
marine environment, will have an older northern
part and will be youngish towards south, though
with the same lithology. Besides, rock units do not
carry with them any connotation of time. The same
beach sandstone with virtually similar lithology
may be repeated in a sequence whenever the
required environment is available.
Biostratigraphy is a study, also based on
objective, verifiable materials, the fossils that help
overcome these difficulties.
4.6.3 Essence of biostratigraphy: index
or guide fossils
The concept of biostratigraphy is based on
observation that organisms have undergone
successive changes through geological time. The
variations are for two main reasons: evolutionary
changes and those due to ecological differences
(Factsheet 4.14, also see Figure 4.1). Bio-
stratigraphy is based largely on evolutionary
changes, though it is always difficult to distinguish
these from changes due to ecological controls,
without critical examination.
Fossil record suggests that evolutionary
changes which produce new species or variations
in characters of species are directional and
irreversible. Once a species is extinct, it does not
reappear in the fossil record; once its morphology
 Chapter 4 Systematics and Biostratigraphy 69

FACTSHEET 4.14
Types of Biostratigraphic Zone

Types Relative Importance of Causes Major Usage (scale/domain)

Assemblage zone env > evn local

Range zone evn > env regional/ global
Concurrent range zone evn > env local
Acme zone env > evn local
Interval zone env > evn local
or evn > env regional/global
env: environment evn: evolution >: dominant over
Causes Effects
Biostratigraphically more useful Biostratigraphically less useful
Evolutionary Rapid evolution Slow evolution
Stratigraphic range short Stratigraphic range long
Morphological distinction greater Morphological distinction lesser
Part Evolutionary Isochronous Diachronous
Part Ecologic Horizontal spread From migration
Ecologic Facies-independence Facies-controlled
Taphonomic Preservation potential high low Preservation potential low
Relatively durable structure Delicate structure and
and morphological details morphology

has changed in a certain trend, it does not trace
back the same course of changes, in all details or
in totality. The so-called degeneration is now
refuted (see Chapters 11 and 12 for examples). As
members of a new species increase in numbers,
they may eventually become abundant and
widespread enough to show up in the geological
record as the first appearance of the species (FAD:
First Appearance Datum). It then reaches an acme
or maximum abundance. When it is no longer able
to adjust to shifting or unfavourable environmental
conditions, its members decrease in numbers and
eventually disappear marking the extinction or last
appearance of the species (LAD: Last Appearance
Datum). Some species exist for only a short span
of geological time. Others may persist longer.
Whatever it be, the span each of them lives through
on the earth and is represented by its fossil record,
marks a particular segment of geological time and
corresponding part of the stratigraphical column.
Thus, each species and its fossils, represent a
particular age and a non-repetitive, irreversible
occurrence. Some of them have greater
stratigraphical values, in the sense that they are
more useful in finding out the relative age of their
host rock units and in correlating them with other
units of the same relative age. Such fossils or
species are called index fossil/ guide fossils or, in
some cases, zone fossils. The characteristics that
make them useful that way are as follows:
1. They should have preservable hard parts to
ensure easy fossilization.
2. They must be morphologically distinct,
favouring their easy and clear-cut
identification.
3. They must be abundant, increasing thereby the
chances of collecting their fossils.
4. They must have had a lifestyle or life cycle
allowing rapid dispersal around the world or
70 Part One: Principles

C C
C
B B
B

A A
A

(a) (b)

upper limit

lower limit
A B
(c) (d) (e)

Biohorizon

Biohorizon

(f)

Fig. 4.1 Different types of biostratigraphic zones.

(a) Concurrent range zone, (b) Lineage/phylozone, (c) Acme zone (width of the bar gives relative
abundance), (d) Taxon range limits, (e) Oppel zone, (f) Interval zone.

at least over very large areas; planktons or
smaller nektons or benthos with meroplanktic
stage are best suitable in aqueous
environment; airborne spore-pollens also
fulfil the requirement.
5. With the above-mentioned mode of living, they
may be distributed widely across geographic
as also ecologic boundaries; it means they may
be found in rocks of different environments;
however, it is now appreciated that even
planktons which are passive drifters in currents,
may be ecologically limited by temperature
gradients, those of warmer water not finding it
hospitable in colder parts of the oceans.
6. The most significant of all, they must be
relatively short ranging with a rather abrupt
beginning in the rock record and a rather abrupt
demise not too separated in time or vertically;
their presence on the earth is, thus, indicative
of a short duration of time.
 Chapter 4
4.6.4 Relationship with other
stratigraphic units and correlation
Relationship of biostratigraphic units with their
lithostratigraphic or time-stratigraphic counter-
parts is far from simple and uniform. Boundaries
of the former may or may not coincide with those
of lithostratigraphic units. Commonly, even
formations, the mappable lithostratigraphic units,
can be subdivided into smaller biostratigraphic
units on distinctive fossil assemblages. These help
date and correlate parts of the sequences in more
details, and over wider geographic extent, thus
allowing reconstruction of geological history for
more precise segments of geological time.
Reverse may also be true, where a biostratigraphic
unit may encompass more than one member or
even formations, indicating a rapid change in
sedimentological conditions within a relatively
short time.
Though fossil record of each species represents
a particular geological age, records of the same
species from two different sequences may not be
time equivalent. Organisms are often restricted to
biogeographic provinces and this provinciality may
create problems. A species may exist in one
province for longer periods of time before
broaching a particular barrier and spreading into a
nearby province. Thereafter, it may die out in any
of the two provinces and continue in the other. The
local vertical range of the species in any province
will be different from that in the other. Besides,
either of these ranges will be shorter than the total
vertical range of the species. Such instances explain
why two different kinds of units, viz.
biostratigraphic and chronostratigraphic are taken
help of. The former serves more for classification
of local or regional sequences (often called
zonation), while the latter is essential in correlations
over longer distances, regional, intercontinental
or global.
The above-mentioned simple constraints or
limitations, however, call for critical judgement.
A single fossil occurrence or a suite or succession
Systematics and Biostratigraphy 71
of fossils does not necessarily reveal simple or
accurate biostratigraphic information (Matthew,
1984). Of different types of organisms, nektons
and planktons are more suitable for biostratigraphy,
simply because they are more likely to be widely
distributed. However, facies-controlled benthos
may prove useful if they have a meroplanktic stage
of life cycle. Conversely, planktons or nektons too
may be facies governed and thus less useful in
correlation.
Thus, benthonic species of anthozoa, e.g.
Calceola sandalina or of brachiopoda, such as
Pentamerus oblongus, are used in biostratigraphy,
the former marking Middle Devonian and the latter
Lower-Middle Silurian age. Graptolites, widely
used in correlation of Siluro-Devonian successions
on account of their distribution as
pseudoplanktonic forms, were however, too fragile
to survive in agitated, shallow water environments
and are, thus, excluded from rocks of such
environments. Similarly, species of planktonic
foraminifers otherwise widely acclaimed as index
forms, differ in cold and warm waters.
Secondly, the assemblage in question, must
represent the same time period as the host stratum.
As mentioned in Chapter 2, bioturbation and
physical reworking also cause time-averaging
(temporal mixing, i.e. mixing of materials of
different time) of different communities and may
lead to increased diversity and variation in
morphological features of fossil lineages. False
LADs are most serious because bioturbation and
reworking preferentially mix sediments upwards,
fossils of older age, thus, occurring in rocks of
younger age. Death assemblages or
thanatocoenosis, too, may give a ‘wrong’ signal,
particularly in areas of slow sedimentation; in such
cases, as sedimentation is prolonged over a longer
time, chances of elements of different assemblages
getting mixed up increases.
Thirdly, the same assemblages or sequences
found in different localities generally prove
contemporaneity. But such occurrences may also
contain forms (species or genera) that are
72 Part One: Principles

diachronous and facies-governed. They should be for instance, or when different apparently useful
recognized and omitted from biostratigraphic work. biostratigraphic indicators may yield contradictory
The absence of a typical assemblage of an age correlations, such as is found in marine rocks near
does not necessarily mean the absence of rocks of continental margin, where marine planktonic forms
the corresponding age. The same age may have may be associated with some palynofossils
been represented by a different facies or an reworked from land from rocks of older age and
assemblage of a different faunal province, the latter brought down to the sea with the clastic input from
in the cases of major palaeogeographic changes the land.
having taken place before deposition of the All such cases need particular critical attention
concerned successions. for a meaningful biostratigraphic conclusion.
Problems may also crop up when a non-marine Figure 4.2 presents an idealized diagram on
vertebrate or land plant assemblage is matched with how biostratigraphic classification and correlation
a marine assemblage of ammonoids or foraminifer, are done.

Locality 1 Locality 2

F 51 Z61 ACF Assemblage ACF

41 51 ACE Taxon range E Assemblage ABR ABR Z52 F72

31 41 ABCD Taxon range D Taxon range D ACDQ 42 62

Concurrent
range BC
Concurrent range CP ACPQ 32 52
31 ABC Assemblage ABC
21 Assemblage AP AP 22 42
21 AB Assemblage AB 32
22
F11 Z11 A Assemblage A Assemblage A A Z12 F12
Lithostratigraphic formation

Lithostratigraphic formation
Biostratigraphic zone

Biostratigraphic zone
Zone type

Correlation

Zone type

Fossils
Fossils

Fig. 4.2 Correlation of two local successions (diagrammatic).

 5
5.1 Introduction: Evolution
Organic evolution provides the single frame which
binds together the vast organic world of the
present and the past, from the simplest bacteria or
any other monera to the huge mammals, the
intelligent animal, man or the gigantic trees of rain
forests. It explains how life appeared and evolved
from its simplest type to the most complex through
time and all over the globe. Palaeontology, in turn,
studies the organic world and its environment on
the surface of the earth with a view to using them
in reconstruction of the history of the earth itself.
Obviously, the subject cannot stand on its right
premise without taking organic evolution into
consideration. Knowledge of organic evolution, on
the other hand, cannot be complete without
knowing how the organic world changed through
the geological past, a subject matter of
palaeontology.
It is unwise to try to present an all embracing
account of evolution, even a meaningful summary
of it, including its genetic and palaeontological
aspects, in the span of a chapter of some few pages.
73
Evolution of
Organisms
In fact, that seems unnecessary too. There are many
relevant materials at hand; for instance, Clarkson
(1998, Chapter 2) provides a beautiful summary;
Colbert, Morales and Minkoff (2002) provide a
useful brief, yet an updated narration of vertebrate
evolution and Benton (2005) another rich current
account. Here only a few general aspects of
evolution that have come out of palaeontological
studies will be briefly discussed, to be followed
by four case histories of evolution, of general
palaeontological importance and relevant to Indian
students as well. Discussions will concentrate on
the following issues:
1. Some basic concepts and place of
palaeontology in the study of organic
evolution.
2. Phylogeny, phylogenetic tree, phenotypic
trends and evolution as interaction of
organism and environment as exemplified
in a few case histories like that of Equidae,
Proboscidea and Hominidae among verte-
brates and ammonoid among invertebrates
(of these, the ammonoid evolution is
discussed in Chapter 12).
74 Part One: Principles
5.2 Organic Evolution
5.2.1 Two modes of evolution
Organic evolution takes place at all levels of the
organic world, from populations to phyla through
a continuing interaction of organisms and their
environments. Thus, phylum Mollusca evolved
from its primtive Monoplacophoran organisms to
the three major classes, Gastropoda, Bivalvia and
Cephalopoda, in the face of exploring and
exploiting three different modes of life, epibenthic,
endobenthic and nektonic, respectively
(see Chapter 8). On the other hand, commonly
perceived evolution of organisms takes place at
the level of species, more particularly populations.
This vast span is divisible into two parts, defined
on two different modes of evolution micro-
evolution and macroevolution.
The conception and definition of micro-
evolution varies slightly with the evolutionary
biologists, essentially neontologists including
geneticists on the one hand, and palaeontologists
on the other. Thus, as conceived by evolutionary
biologists, microevolution occurs within
populations, as also within species, over relatively
short periods of time; for population geneticists it
involves genetic changes, viz. mutations, then
changes in gene frequencies within populations
driven by selection, migration and drift.
Geneticists view microevolution as not the
whole story and consider speciation as transition
between micro- and macroevolution. Speciation,
i.e., origin of new species (also see Chapter 4) is
splitting of populations in evolutionarily
independent units. To that extent, it involves
changes different from, yet interconnected with
those of microevolutionary changes within
populations and macroevolutionary changes at
higher levels (Stearns and Hoekstra 2001).
Palaeontologists, in their turn, consider
microevolution as pertaining to small-scale
changes within species and especially concerning
transformation of one species into a new one, i.e.,
speciation (Clarkson 1998). Phyletic gradualism
and punctuated equilibria model, treated in Chapter
4, are the two different processes visualized for
explaining speciation. But the occurrence of
gradualistic and punctuational changes in the same
succession suggests that both the modes may be
real, not contesting and rather complementary.
Macroevolution involves larger scale changes
at the levels between species and involving larger
groups, such as families, oders and phyla. It, thus,
covers changes such as adaptive radiations, mass
extinctions, etc. Palaeontology particularly
contributes to revealing and understanding
macroevolution.
5.2.2 Rates of evolution
Organic evolution not only differs in modes, it takes
place at different rates too. Even within a mode
itself evolution is never uniform in its rate.
Thus, changes in species include stasis and
punctuations, the former slower changes
interrupted by rapid changes at punctuation at
which new species are born.
A different case of varying rates of evolution
in microevolution is obtained at, what is known
as, heterochrony. Different organs of the body of
an organism appear and grow in a definite sequence
during ontogeny. The timing is genetically
controlled. What size and shape an organism will
attain depends on the timing of rate of
development. But, ‘changes through time in the
appearance or rate of development of ancestral
characters’ (Gould 1977, quoted from Clarkson
1998), the heterochrony, may occur when time-
coordinated development is decoupled. For
instance, a brachiopod will grow to the normal size
and shape, when there is a coordinated
development of size, shape and timing. But, if there
is a mutation that causes genetic timing mechanism
that stops the growth at an early stage than when it
was really supposed to act, the organism will
develop with a size less than its normal one. That
is, there will be a dwarf, as compared to the

ancestor. On the contrary, there may be a giant, if
the mechanism acts late, that is growth stops later
than its usual schedule.
There may be changes in shape or morphology,
if different structures develop at different rates. In
one case, the adult of the descendant may retain
some or whole of the juvenile characters of the
ancestor. It means the characters did not change as
fast as they should have. This is known as
paedomorphosis.
In the second type, the descendant adult may
be morphologically advanced than the ancestor,
meaning faster than usual rate of change; it is called
peramorphosis.
It is now considered that many micro-
evolutionary changes in fossil record may have
taken place through heterochrony. It may be
important for macroevolutionary changes too. A
good example comes from trilobites. Most
trilobites have holochroal eyes, in which many
small lenses are closely packed together. This is a
primitive type too, appearing in early Cambrian.
In early Ordovician, there appeared a new type of
eyes characterizing the phacopid group. In these
schizochroal eyes, lenses are larger, fewer and
separated by cuticular material. Closer scrutiny
reveals that schizochroal eyes are very similar to
what an adult trilobite with typical holochroal eyes
had in its juvenile stage of ontogeny. It is, thus, a
case of paedomorphosis that retained more
primitive holochroal juvenile eyes in more
advanced or descendant adult schizochroal-eyed
trilobites (Clarkson 1998; Figure 5.1 (a)).
Three paedomorphic processes are
differentiated (McNamara 1990): (i) progenesis
occurring with early onset of sexual maturation,
whereby the organism with all its structures
affected develops into a small and juvenile edition
of its ancestor; (ii) neoteny, in which
morphological development takes place at a slower
rate, may affect all or some characters and the
descendant is of nearly the same size as that of the
ancestor (Figure 5.1a); and (iii) post-
displacement, in which some single character
Chapter 5 Evolution of Organisms 75
starts to grow later than others do. Peramorphosis,
too, may be of three types: delayed sexual maturity,
rapid morphological development or early start of
growth. Morphology of a descendant may be a
complex of normal, paedomorphic and
peramorphic characters.
5.2.3 Rates of evolution, appearance
and extermination in evolution
Evolution was not uniform in its rates; neither was
it uniform in producing the number of organisms.
This is another significant aspect of evolution,
namely the appearance or origin of new forms, as
well as disappearance, extermination or extinction
of existing forms. The number of species living
on the earth at any given time is determined and
maintained as a sort of equilibrium condition, by
the positive process of origin of new species, as
well as emergence of newer higher level taxa and
the negative process of extinction. There are
periods of adaptive radiation or explosive
evolution that are episodes of sudden appearance
and rapid proliferation of new forms and there are
mass extinctions which are episodes of rapid
large-scale demise of taxa of related or unrelated
groups. However, these episodes or periods of
rapid changes, addition or deletion, are interspersed
with periods of relatively slower evolution. This
makes it necessary to discuss rates of evolution in
relation to appearance and extinction.
In the continuous process of organic evolution
through constant and continuing interaction
between organisms and their environment, there
may appear a successful new organic group,
generally with new body plan or a new type of
adaptation. Normally, this is coupled or interlinked
with a major change in environment, often of global
span that creates pressure on development of that
new body plan or demands a new type of
adaptation. This creates an increased geographic
range and a variety of niches to the evolving group,
which then utilizes the situation with rapid origin
of newer taxa, at different levels, species to higher.
76 Part One: Principles

(i) (ii)
(A)

(i) (ii)
(B)

(i) (ii)
(C)

Fig. 5.1 (a) Paedomorphosis including neoteny.

(A) (i) and (ii) Chimpanzee skull at foetus and adult stages, (B) (i) and (ii) Human skull at foetus
and adult stages, A(i) and B(i) are very similar; but slower rate in development of human skull
makes it much different from the chimpanzee skull at the adult stage, as found from A(ii) and
B(ii), (C) Trilobite eyes: (i) holochroal and (ii) schizochroal.
In holochroal eyes polygonal lenses are in contact and covered by a single cornea; in schizochroal
eyes each lense has its own corneal cover and the adjacent lenses are separated by a different
interstitial material. Trilobites with holochroal eyes seem to have schizochroal type at the larval
stage; hence, adult schizochroal eyes are interpreted as formed by paedomorphosis, i.e. retention
of juvenile character of ancestor in the adult of descendent.

100
80
60
40
20
0
Cm O S D
600
200
Fig. 5.1(b) Appearance, extinction abundance of animals through time.
(A) Appearance (dashed line) and extinction (solid line) of animal families through geological time
(B) Overall increase in number of families during the same time span (a and b modified from Babin
1980).
They find new adaptation to the available niches
and evolve quickly, setting up the adaptive
radiation. Thus, at this stage of evolution,
appearance or origin of new taxa is overwhelmingly
greater than disappearance or extinction of existing
taxa, producing what is called tachytelic evolution.
Once the group in question has found new
adaptations stabilized, with many different taxa
fitted to small differences in niches to their
respective advantage, they start evolving relatively
slowly adjusting to their changing environment.
However, initial quick adaptation to a narrow limit
of environmental parameters set significant
limitations to their evolution too. Many of the new
taxa become extinct with slight variations in
environment, often replaced by new ones in the
same stock or group and with similar adaptations.
There is, thus, more or less equal number of taxa
becoming extinct and appearing anew. Such an
evolution is called horotelic evolution.
Chapter 5 Evolution of Organisms 77
LC UC P Tr Jr K Cz
(A)
(B)
More commonly, after the adaptive radiation
or its initial burst, there is a rapid decline in number,
with only a few taxa or stock persisting with very
little changes. They seem to maintain the narrow
range of adaptation to a specialized or particular
set of environment, as a sort of a conservative
stock, and may produce what are known as living
fossils. Such extremely slow evolution, in which
evolutionary changes are apparently absent, is
known as bradytelic evolution. Lingula, an
‘inarticulate’ brachiopod adapted to a rather
uncommon environment of mud-flat and shoals
exposed at low tide is an example; coelacanth fish
is another.
In organic evolution, mass extinctions are
periods or stages in which extinction overrides
appearance, meaning a sudden extinction of a large
number of organisms and their groups. Massive or
not, extinction is an integral part of evolution. It is
discussed below.
78 Part One: Principles

5.2.4 Extinction with many newer species trying to adopt to the

available new niches. In the marine Phanerozoic
Organisms may eventually become extinct. Just as
fossil record, there are thus 12 ‘ecological-
new taxa arise through time, they decline and
evolutionary’ units, recognized by some authors that
disappear. There is, thus, a continuous changeover
are punctuated by extinction events of a greater or
of fauna, with innumerable minor extinction and
lesser magnitude. Within each of these units, both
replacement episodes occurring between any two
populations and communities were relatively stable
major adaptive radiations. Periods of mass
and their respective structure ecologically
extinction are really pulses that are interspersed in
controlled. When major extinction events disrupted
this continuous process when many taxa become
the structure of ecosystems, not only were new
extinct almost simultaneously.
structural and physiological innovations established
Natural selection, the primary process of
during the subsequent adaptive radiations, but also
microevolutionary change, generally works on
community types were completely reconstituted in
enough genetic variability to produce individuals
new and different patterns.
that are adapted to the new conditions. These are
There are several mass extinction periods
selected to survive. In case it fails, that is, where
documented in the fossil record. The most severe
there is no successful adaptive shift possible,
was the late Permian crisis wiping out marine
extinction ensues. Competition, predation, changes
invertebrate families by 57 per cent (~ 95 per cent
in the physical environment and chance changes
species disappeared). In the late Ordovician (end-
in population are external factors that bring about
Ashgillian) 22 per cent of all families died out, in
extinction. Competition leads to extinction when
the late Devonian (Frasnian-Famennian) 21 per cent,
one or more species invade the range of an inferior
in the late Triassic (Norian) 20 per cent and in the
competitor. The latter then becomes extinct.
late Cretaceous (Maestrichtian) 15 per cent. The last
Predation works to extinction when the predatory
of these both and marine faunas became extinct.
species feeds on a variety of prey species. Physical
Potential causes of the deterioration of global
conditions are varied; climate on land and
ecosystems, in turn causing mass extinctions, may
temperature, salinity and dissolved oxygen in
be listed as in Factsheet 5.1.
marine realm are of great importance.
Large-scale extinction episodes terminate each FACTSHEET 5.1
of the geological periods and the systems
themselves are defined on the basis of the faunas Potential Causes of Extinction
they contain. At the end of the Cambrian, the
Earthbound mechanisms:
typical trilobite faunas of the period changed Extinction of key species (plant or animal) of the food
globally. Most of the families became extinct and web, may disrupt the entire network of trophic
were replaced by entirely new stocks. relationships
Once new ecosystems are set up, they tend to Global temperature decline
become rather rigid in an evolutionary sense. Not Marine regression
much innovative changes seem possible to take Volcanism
place in organisms living in such an ecosystem. Oceanographic effects, viz. circulation, changes in up-
Organisms tend to evolve slowly. But once it again welling systems, either adding to overturning of stag-
breaks down, the organisms face the rapid doom nant bottom water or diminishing it and thus depleting
planktonic nutrients, etc.
and the potential for new replacements, seemingly
Extraterrestrial mechanisms:
opportunistic, become available. At such times, the
Supernova radiations
older fauna (or flora or both) become extinct and Bolide (asteroid) impact, etc.
new and promising developments tend to prosper

5.2.5 Divergence and convergence
One more aspect of evolution seems to draw
general attention. It is the question of divergence
and convergence, understood in palaeontology in
context of morphology. Thus, divergence means
appearance of many morphologically different or
distinct organisms from relatively fewer ancestors.
Obviously periods of adaptive radiations are also
the periods of maximum divergence being
produced. Availability of many different niches sets
the scope for different adaptations and, thus,
divergence.
In contrast, convergence produces
morphological similarities in descendants of
different ancestors. Here again, environment has
its role to play; living in a similar kind of
environment may necessitate similar kind of
adaptation and thus convergence. Parallel
evolution is a phenomenon whose effects are
comparable to adaptive or evolutionary
convergence. Both parallel evolution and
convergence lead to homeomorphy that is
development of close morphological similarity in
two or more unrelated groups (Factsheet 4.6).
In parallel evolution, two closely related stocks
with minor morphological differences undergo a
series of similar evolutionary changes through
time. In it, the ancestors are nearly similar, whereas
in convergence they are quite distinct. Both can
occur in either phyletic (within a lineage) or
phylogenetic (within a group) trends. Convergence
may be isochronous that involves species or groups
of the same time, or it may be heterochronous
involving forms of different ages.
5.3 Palaeontology in the Study
of Organic Evolution
Organic evolution embraces the whole organic
kingdom. Much of this kingdom would have re-
mained unknown to man, but for the fossils. It is
the study of fossils that has shown that different
kinds of organisms appeared on the surface of the
Chapter 5 Evolution of Organisms 79
earth at different times and lived through some range
of time (known as stratigraphical or geological range
of organisms or their fossils). Again, it is the study
of fossils that has revealed the presence of vast
number of organisms which do not live any more
today; they are extinct, leaving behind their fossils
only to mark their one-time existence. Organic
evolution owes it to palaeontology that to obtain its
fuller picture, it could find the answers to questions
such as what were the extinct groups, how were
they linked with the extant groups, what brought
about their extinction, and such other questions.
Spectacular example comes from the well-known
dinosaurs; no less known to palaeontology students
are the examples of trilobites or ammonoids.
Recent finds of rich records of soft-bodied
organisms in Precambrian, without any trace of
them afterwards are additional vital cases in point
(see Appendix 1).
As discussed in section 5.2, fossils have not
just recorded extinctions; they have also provided
evidence of appearance of newer and newer
groups, often to fill in vacuum, as it appeared,
created by the extinction of some group or others.
Any theory of evolution must take these
appearances into full account. For example, if
scleractinian corals that appeared in Triassic after
rugose corals had become extinct in Permian, were
they related to the latter or not is definitely a
question evolutionary studies need address.
Well-preserved fossil records have proved that
evolution of a group after its modest appearance
is not a simple story of gradual increase in number,
variety and complexity of its members. Rather,
almost invariably, there are periods of adaptive
radiation, when a large number of genera and
species appear suddenly in the record, live for a
short time and become extinct shortly afterwards.
These are the junctures in the evolution of a group
when it evolves or evolved fast. Likewise, there
are periods when the group evolved or evolves slow
without much addition to its number or varieties
or to changes in the character of its members; the
fossil records are there to prove it. One cannot
80 Part One: Principles
expect such documentation of rates of evolution
from the portion of the organic world, living at a
small segment of the present time only.
It is the fossil record that has also brought
the fact to man’s knowledge that all the major
phyla, representing major types of body
organization, appeared very early in the history
of life. Smaller groups and subgroups of these
major groups appeared and evolved through
interaction with their environments, physico-
chemical as well as biological, local as well as
global. In course of this evolution, there emerged
groups above species level, the macroevolution.
At the same time there were changes within
species and origin of new species, the
microevolution. Palaeontology contributed to
elaboration of understanding of both.
Even the far-from-complete stratigraphical
successions prove convincingly that each organism
has a particular range of its life on the earth and
that once extinct it never reappeared. The
occurrence was thus irreversible, a fact that not
only made a fossil useful for marking
stratigraphical age, but also for building up the
sequence of organisms as it appeared successively
in course of evolution. From these, palaeontologists
could draw up lines of descent at all taxonomic
levels and the trends, patterns and courses of
morphological (phenetic) or evolutionary changes
that took place in those lines. Summing up such
data, phylogeny of the group could be
reconstructed and judged on the background of
stratigraphical and palaeoenvironmental as well as
tectonic and climatic data. It would then provide
the entire evolutionary history of the group set in
the backdrop of changing face of the earth with
plate tectonics and major climatic, sea level and
other changes through geological time.
Organic evolution and its fuller understanding
entails a lot more issues and questions, decided or
debated. Genetic mechanisms are important
considerations. Yet, as mentioned earlier, this brief
introduction is aimed at presenting only the few
vital issues that may be of concern for the beginners
of palaeontological studies. With this introduction,
we may now move onto looking at some case
histories that will provide many instances of the
questions and phenomenona discussed above.
5.4 Introduction: Proboscidea,
Equidae and Hominidae
Proboscidea (elephants and their ancient), Equidae
(horses, etc.) and Hominidae (human and related
forms) are the three well-known groups of land
vertebrates. The first is an order itself; Equidae
belongs to the Order Perissodactyla, Hominidae
to Primates, all under the phylum Chordata
(see Factsheet 4.4). Traditional views on the
systematic position of these animals have been
largely challenged by phylogenetic systematics
based on cladistic methods. That demands a bit of
elaboration on this point to be included in a
separate section in this chapter.
Both Proboscidea and Equidae range from
Eocene to Recent, whereas Hominidae ranges from
Pliocene to Recent, though it might have appeared
in Miocene. Proboscideans are and have been
forest-dwellers, equids prefer and preferred
grassland, though the earlier members of the family
were forest-living. Hominids might have started
in a mixed chequered environment of grassland
and forest, though its present-day representatives,
the modern men have made virtually the whole
world habitable for them.
Fossil records of the three groups are not
equally rich. Of the three, equids present a very
well-preserved record in North America that has
been fairly well-studied too. Records of hominids
are rather scanty. But intrinsic interest in the
evolution of our own kins, have led scientists to
undertake significant studies on the basis of
available materials. Unabated interest has also
helped continue the search for more and more
fossils, yielding encouraging results. Probo-
scideans did capture man’s imagination and
attention since long. Distinct, large fossil

specimens were also suitable for studies. But the
fossil record of this group is not quite rich and the
evolutionary picture is also rather complex.
In spite of these differences, evolutionary
pictures of these three groups stand representative
to demonstrate very many interesting features and
aspects. So, they have been included together in
this chapter. Several invertebrate groups such as
trilobites, graptolites, echinoids and of, course,
ammonoids among cephalopods have good
documentation of their evolution in the fossil
record. Among them, only the evolution of
ammonoids will be taken up as a representative
case, in the chapter on Cephalopoda (Chapter 12).
It is fairly well-established that evolution of
Equidae hinged on the basic adaptive change
from browsers to grazers and accompanying
problems of locomotion and feeding. No such
clear-cut conclusion can be made with the
evolution of Proboscidea. They were forest
dwellers with big bodies. So the major adaptive
changes might have hovered around feeding and
concomittant changes in skull, jaws, etc. For
humans, many conclusions can be made about
adaptive changes in course of their evolution.
Interpretations of functions of different parts of
body, their significance in the habit of living can
be made quite easily. It is readily verifiable. In
summary, evolution of these three groups have
immensely helped augment our knowledge about
the organic evolution in general.
5.5 Revised Views on Vertebrates
5.5.1 Phylogenetic systematics and
molecular phylogeny
As mentioned in Chapter 4, cladistic method has
emerged as a powerful alternative means for
classification of organisms, now claiming
increasingly wider acceptance among scientists.
For vertebrates, two principal analytical techniques
are adopted to establish their phylogenetic
relationships: cladistic analysis of morphological
Chapter 5 Evolution of Organisms 81
data and molecular phylogeny reconstruction. The
former has already been introduced. Here we
present a brief introduction on molecular
phylogeny.
Many kinds of organic molecules record
evolution and relationships among organisms may
be brought out by comparing them from different
organisms. Sequences of nucleic acids, particularly
different RNA molecules have been widely used.
Polymerase chain reaction (PCR) technique
invented for cloning small samples of nucleic acids
to analyzable quantities and computer programmes
for dealing with large amount of data to find out
the phylogenetic trees have helped this branch of
studies develop since 1980s.
5.5.2 Some revisions
Molecular evidence has now made it clear that
relationships of early placental mammals, called
basal placental relationships, were controlled by
biogeography. There is, however, difference in
deciding if eutherian vertebrates (Eutheria or
Placentalia is a cohort in the class Mammalia, it
includes all placental mammals and is the most
dominant group of the class. Marsupial mammals
are grouped in a second cohort, Marsupialia) arose
in southern continents of Gondwanaland in Early
Cretaceous and were disjuncted by their split later
in the same period, or if first eutherians were
originated in the Laurasia (Eomaia from China
from 125 million years ago) and their descendants
spread out from there to Africa by the end of
Cretaceous and to South America by Palaeocene.
In any case, early in the history of placental
mammals there lived in Africa a group, named
Afrotheria, diverged from the basal placentals to
form an independent clade with common shared
derived characters. The clade included as different
groups as elephants (proboscideans), golden moles,
tenrecs and aardvarks (the latter three are insect-
eating mammals). Of them Proboscidea, along with
Hyracoidea and Sirenia form a clade Paenun-
gulata within Afrotheria.
82 Part One: Principles
After the divergence of Afrotheria and
Xenarthra (in South America), the stem of
placental mammals gave rise to Boreotheria which
later diverged into two sister groups, Laura-
siatheria and Euarchontoglires. The former,
Laurasiatheria includes Chiroptera (bats), Artio-
dactyla (cows, goats, giraffes, hippopotamuses,
pigs, etc), Cetacea (whales), Carnivora and
Perissodactyla (horses, rhinoceroses, tapirs) and
others. The latter, Euar-chontoglires, includes
Primates, Rodents (rats, etc.), Lagomorpha
(rabbits, etc.), etc.
The new molecular phylogeny holds that
Perissodactyla form a sister group of Carnivora and
Pholidota (pangolins: ant-eaters) together.
As distinct from artiodactyls, even-toed ungulates,
perissodactyls are odd-toed ungulates (1, 3 or
5 toed).
In Euarchontoglires, primates (along with a
few other smaller groups) belong to Archonta, a
sister group of Glires that includes rodents and
lagomorphs.
Revised relationships of primates, particularly
in relation to Homo, the human genus, is shown in
Factsheet 4.4.
5.6 Order Proboscidea
5.6.1 Introduction
Elephants, belonging to the order Proboscidea, are
familiar and fascinating creatures to man. Modern
elephants are actually the last representatives of a
dying group. Still numerous as individuals, they
are limited at the present time to two genera, each
with a single species, one Asiatic (genus Elephas)
and the other African (genus Loxodonta). Remains
of these great beasts were among the relics of
extinct animals first known to civilization, to the
ancient Greeks and Romans as well as the Tlascan
Indians of Mexico.
Fossils show that at various times during the
Cenozoic era, the proboscideans lived on all the
continents of the world except Australia and
Antarctica. These great beasts evolved along
numerous lines of adaptive radiation, some of
which continued into Pleistocene times, coexisting
with humans. It has made the phylogenetic history
of the proboscideans very complex, with several
instances of parallel evolution.
5.6.2 First proboscideans
The first proboscideans known from the fossil
record were from Africa; the oldest is from lower
Eocene of Morocco, Phosphatherium (Gheerbrant,
Sudre and Cappetta, 1996) followed by
Moeritherium, from upper Eocene-Oligocene of
Egypt. They belonged to the moeritheres. Colbert,
Morales and Minkoff (2002) however, hold that
the earliest moeritheres lived in Southern Asia
during early and middle Eocene times.
Anthracobune, first described in 1940, and the
more recently described Pilgrimella and
Lammidhania are very primitive moeritheres.
5.6.3 Phenotypic characteristics
and trends
Proboscideans, today and in the past as well, are
characterized by a number of following features:
1. Reduced jugal and orbit that opens in the
maxilla.
2. Enlarged second upper incisors to become
tusks in later forms.
3. The absence of lower canines and first
premolars.
4. Broad molar teeth with thickened cusps.
5. Limbs adapted for supporting body weight.
In addition, Moeritherium, the early
proboscidean which represent a morphological
archetype, presents the following characters:
1. Generalized plan of body and specialized
skull
2. Deep skull
3. Upper and lower second incisors enlarged as
short projecting tusks

4. Long-bodied animal (~1 m tall), size of a pig,
probably living in freshwaters, like small
hippos.
On this archetype, proboscideans show the
following dominant phenotypic trends among
varied patterns of developments:
1. Increase in size. Almost all the proboscideans
became giants.
2. Lengthening of the limb bones and the
development of short, broad feet. This has
been a common evolutionary trend among
very large mammals.
3. Growth of the skull to extraordinarily large
size, even in proportion to the rest of the body.
4. Shortening of the neck. Because the skull and
its associated structures became large and
heavy, the neck was reduced in length to
shorten the lever between the body and the
head.
5. Elongation of the lower jaw. In many of the
later proboscideans, there was a secondary
shortening of the lower jaw, but lengthening
of the jaw was an early primary trend.
6. Growth of a trunk. Elongation of the upper
lip and the nose probably went along with
elongation of the lower jaw. Subsequently the
nose was further elongated to form a very
mobile trunk or proboscis.
7. Hypertrophy of the second incisors to form
tusk, used for defence and for fighting.
8. Limitation and specialization of cheek teeth
in various ways, as adaptations for chewing
and grinding plant food.
5.6.4 Proboscidean radiation
After Eocene, barring the emergence of the
barytheres (a peculiar, little known and palaeonto-
logically less important group from Egypt), there
were two principal lines of proboscidean
development, diverging from the moerithere stage.
These were the deinotheres and the euele-
phanotoids (otherwise called elephantiforms).
Chapter 5 Evolution of Organisms 83
The deinotheres were modestly proportioned,
standing 3 m (10 ft) height at the shoulder with
long legs. They had a pair of lower tusks curved
back under the chin from the lower jaw towards
the body. There are different opinions on its use.
In one suggestion, they were used as hooks,
employed for digging into the ground and pulling
up roots or plants (Colbert, Morales and Minkoff
2002). In another, it is held that they may have
been used in scraping barks from trees (Benton
2005). The upper tusks were lost. They had well-
developed trunk.
The deinotheres evolved in the Old World
along a very narrow path of adaptations. Their
record shows some pecularities. They seem to have
evolved from the moeritheres rapidly in Eocene,
though this is not confirmed by fossils. However,
the group leaves no record in Oligocene and
became extinct in mid-Pleistocene. During this
course, between early Miocene and Pleistocene,
there was little morphological progress. There was,
thus, an example of rapid initial evolutionary
development followed by a long period of
evolutionary stability at a high level of
specialization.
The elephantiforms had two groups,
palaeomastodontids of Eocene-Oligocene of Egypt
and the elephantoids. The latter shows rapid
morphological modifications and divergence into
groups of superfamilial rank during Miocene. They
were the large, trunk and tusk-bearing giants that
spread to almost all corners of the earth during
middle and late Cenozoic times.
Thus, there were:
1. Long-jawed gomphotheres (e.g. Gompho-
therium, more commonly known as
Trilophodon; gomphotheres had four short
tusks and spread from Africa to Eurasia and
North, even South America. They also
included Miocene-Pliocene genus Serri-
dentinus, upper Pliocene Syconolophus, lower
Pleistocene Stegomastodon and during
Pleistocene Cuvieronius in South America).
84 Part One: Principles
2. The short jawed mastodonts, held as a
paraphyletic group (Benton 2005: they arose
perhaps in Central Asia, and spread rapidly
over Asia, Europe and Africa to reach America
in early Miocene: Mastodon americanus was
abundant in North America and the first
humans in America, not the first Europeans
there, might have been contemporaneous with
the mastodon at around 8000 years ago).
3. The stegodonts (including in the Old World
Stegolophodon during late Miocene and early
Pliocene and then Stegodon in late Pliocene
to Pleistocene).
4. Finally the elephantids (mammoths and
elephants) which evolved with great rapidity
and profusion during Pliocene and
Pleistocene.
5.6.5 Trends in elephantid evolution
Increase in body size, teeth reduced to few in the
jaw at any time, tusks and a trunk, were mutually
linked phenotypic trends found in all these groups,
the clades. As elephantoids grew taller, their head
heavier, the latter had to be supported by a short
neck to add mechanical advantage to the
movements of head with shortening of distance
between it and the body. This put constraints on
the tall animal to reach the ground with its mouth.
This problem was met with by the development of
a long proboscis or trunk.
Elephantids show changes in teeth characters,
adapted to feeding upon harsh vegetation of forests.
It needs teeth strong enough to break the branches
and stems, sharp enough to cut down their hard
cover or bark into pieces to get through to their
softer inside and large enough to stand high rate
of wear during lifetime. With this is added the fact
that modern elephants live long (<60 years) which
prolongs tooth-wear too.
The problem is solved in several ways. It is
found that Moeritherium had all six cheek teeth in
each jaw (as in other mammals), the modern
elephant has only one or two in each jaw at any
time. It means, of the 6 cheek teeth, the first 3 are
milk molars lost by 15 years age. Number 4 come
into use at 18–28 years of age; number 5 at 40–50
and number 6 at age 50 or more. Teeth are also
pushed forward on the jaw as they erupt and as
they replace earlier ones. This trait is
phylogenetically or evolutionarily shared with
sirenians (sea cows); besides, kangaroos have the
same trait, a convergence in nature.
A second trend in elephantid teeth is the
development of cement to fill the valleys between
the lophs or rows of cusps fused together; the latter
are made of tough enamel cover on soft dentine.
As a result, when worn out, the crown surface of
the tooth presents alternating zigzag ridges of hard
enamel and valleys of dentine and cement (in the
sequence enamel, dentine, enamel, cement,
enamel, dentine, enamel, and so on). This makes
an effective cutting surface on each tooth. (For
details of mammalian dentition, see Appendix 2).
5.6.6 Mammoths and extinction of the
order
Mammoths were elephantids that lived during the
Pleistocene Ice Age and spread from Africa over
much of Europe and Asia and later, North America.
They make a monophyletic group. DNA analysis
has related them to the African elephants. The
woolly mammoth is also famous for its fossils in
Siberia and Alaska, where, in the best case, the
entire body, the thick fat layer and the hairy surface
and even the food material in mouth and stomach
have been preserved. Mammoths lived with early
humans. They were extinct in Europe 12,000 years
ago, in North America about 10,000 years ago. A
dwarf variety may have lived in Russia upto 4000
years before present.
Extinction of these large mammals which were
abundant in Pleistocene, may have been due to the
advent of the Ice Age as well as to hunting by man,
which lived contemporaneously with these
animals. As a result of extinction, this spectacular
group of terrestrial animals dwindled to a mere
 Chapter 5 Evolution of Organisms 85

fraction, finding representation in only two genera 5.7 Family Equidae

in the recent times. It has, however, left the record
of evolution, with instances of rapid changes
followed by slow conservativism, of parallel
evolution and some unique adaptations in food
gathering and mastication, which were coupled
with changes in anatomy, particularly development
of large size of the body and skull. For a broad
phylogeny see Figure 5.2 and for Indian records
see section 18.4.5 to 18.4.8.
5.7.1 Introduction
The family Equidae (Lower Eocene-Recent with
only one genus, Equus, remaining in the present-
day; it includes horses, zebras, asses, etc.)
belongs to the order Perissodactyla of the class
Mammalia. The fossil record of the family,
particulary from North America, presents a more

Mastodonts Elephants

Gomphotheres
Stegodonts
Q
HYRACOIDS

PROBOSCIDEANS

P
?
S
LID

S
TY

IAN
OS

EN

Deinotheres ?
SM

M
SIR
DE

Moeritheres

DS
ITH OPO
E E MBR

Fig. 5.2 Diagrammatic phylogenetic tree of Proboscidea showing only the major superfamilial groups of
the order and its related orders.
Uncertain diversification points are left with question marks. E stands for Eocene, O for Oligocene,
M for Miocene, P for Pliocene and Q for Quaternary (adapted from various sources for example, Colbert,
Morales and Minkoff 2002, Benton 2005).
86 Part One: Principles
or less continuous sequence of genera and species.
Because of their tendency to live in large herds,
equids have been buried and fossilized in great
numbers, almost since the early stages of their
phylogenetic history.
All this has made it possible to reconstruct a
fairly detailed picture of the evolution of the group.
Ideas on the equid evolution have, however,
changed. Earlier it was cited as an outstanding
example of ‘straight line evolution’ or
‘orthogenesis’. It was maintained that these
animals evolved with little deviation, along a straight
course of change, with a set of progressive changes
acting upon successively younger forms, from
Eocene Hyracotherium (or Eohippus) to Equus.
However, though there were many progressive
changes or trends of evolution that could be
followed through time, the reverse was also true
and there were various branches in which some
features showed progressive changes, while others
were conservative. On the whole, the phylogeny
is never a single line of gradual transformation of
orthogenesis (the only part in the phylogeny that
could be considered straight to some extent is
between Eohippus to Miohippus during Eocene to
Oligocene) and is repeatedly splitting and
intricately radiating to produce, rather a bush, than
a tree.
5.7.2 Ancestry
Ancestry of equids is not well-documented by
fossils. It is possible that the earliest equid,
Eohippus, a prototype of the odd-toed ungulates
or perissodactyls, was derived from condylarths
of Palaeocene.
5.7.3 General observation
Tooth pattern and anatomy, particularly of limbs,
of Eocene equids point to a browsing habit of the
animals. By the end of Oligocene, the animals
attained the status of advanced browsers, of which
Mesohippus might have been of forest-dwelling
and Miohippus of plain-living types, with otherwise
more or less similar characters. They were capable
of eating leaves and soft plants, and able to run
fairly rapidly and for sustained periods over a
hard ground.
There was a branching out or adaptive
radiation in early Miocene, rather at and around
the Oligocene-Miocene boundary, probably in
response to an increase in the variety of environ-
ments available to them, especially including the
spread of early grasses and other flowering ground
plants. One line remained conservative
(Archaeohippus) in all characters; in another
(Anchitherium-Hypohippus) some characters (size)
showed progression, but others (teeth and feet)
remained conservative.
The main line (of Merychippus) produced
grazers with changes in most characters that
continued in the later part of equid evolution. To
these changes were added the effects of migration
of different forms at different stages of evolution
from the main centre of North America to South
America and/or Eurasia. The emigrants often
continued in the newly invaded continents with or
without significant evolutionary changes.
Equus, culminating most of the progressive
changes in the main line of evolution or holding
over to certain character of its ancestor, viz.
Pliohippus, migrated from North America to other
continents, but became extinct in both the American
continents by the end of Pleistocene, continuing in
the Old World till the present. Recent horses in the
new world are feral, i.e., of domestic origin.
5.7.4 Phenotypic trends
Evolution of equidae reveals a number of
phenotypic changes. These are as follows:
1. Change in size involves an overall increase,
with, however, some decrease in some
branches and from Pleistocene to Recent.
2 (a) Changes in limb ratio (which may also be
called the speed ratio), i.e. proportion of
lower to upper limb segments or
tibia:femur in hindlimbs and radius:

humerus in forelimbs. Increase in these
ratios helped the animals to bear their body
weight more efficiently and, thus,
facilitated the change in the habit of living
from browsing to grazing (in Oligocene-
Miocene). The ratio showed an increase
upto Merychippus that is after the change
from browsing to grazing types and
subsequent decrease in grazing forms.
(b) Changes in ulna and fibula included an
overall reduction, which caused a
limitation to the range of movement.
3. Changes in foot-mechanism:
(a) Changes in foot posture from plantigrade
(walking on the sole of a padded foot) in
browsers to unguligrade (walking, rather
trotting on hoofs) in grazers.
(b) Reduction or loss of lateral digits or toes
from five to one, with emphasis on the
middle toe.
(c) Perfection of the hoof.
These changes are correlated to the
change from a padded foot in browsing
forms to a springing foot in grazing forms.
Changes took place in rather three rapid
transitions :
(i) From early padded foot with 4
functional toes in front and 3 in hind
legs, there came padded foot with 3
toes with emphasis on the middle
hoof, in late Eocene.
(ii) In late Oligocene-early Miocene,
changes from Parahippus to
Merychippus produced feet with
three functional toes, reduced
lateral toes functioning only for
buffering and stopping; loss of pads
made the foot unguligrade; these
changes also affected a complex
springing mechanism.
(iii) In early Pliocene, in Pliohippus, foot
became one-toed, with springing
Chapter 5 Evolution of Organisms 87
mechanism simplified and strength-
ened and check ligaments developed
for the purpose.
4. Changes in skull and dentition:
(a) Increase in hyposodonty i.e. in the height
of the crown of molar teeth. This took
place slightly in loose correlation with
size in early forms and browsers. It was
rapid in Miocene during final transition
to grazers (Parahippus-Merychippus);
thereafter the change stopped.
(b) Lengthening of the face in front of eyes
and increase in height to accommodate
the high-crowned cheek teeth; found in
all but the early forms.
(c) Perfection of dental battery in elongation
and complexity of teeth.
(d) Premolars become molariform with all
but the first premolar, molarized
completely by the early Oligocene.
(e) Complexity of the cheek-tooth pattern:
from bunodont to lophodont to complex.
This change was practically absent in
early browsing forms and began to appear
in late Oligocene in ancestral grazers,
becoming rapid in Miocene.
(f) Widening of incisor teeth.
(g) Increase in size and complexity of brain.
(h) Development of a post-orbital bar behind
the eye region separating orbit from the
temporal region as found in Merychippus.
These trends too, are related to the change from
browsing to grazing habit. The trends in the skull
may also have an allometric relationship with
(i.e. related to the change of) increase in gross size.
Thus, all but the first premolar were progressively
molarized completely by the early Oligocene,
whereas complication of cheek-teeth pattern was
practically absent in early forms and in all the
browsing forms but began to appear in ancestral
grazing forms (late Oligocene) and progressed
rapidly during Miocene. In early forms and browsers
88 Part One: Principles

hypsodonty increased slightly in loose correlation
with their size; but the trend gained momentum in
Parahippus-Merychippus lineage in Miocene.
The above phenotypic trends are interpreted
from the fossil record of the family. They, in turn,
help bring out the phylogeny of the group.
The true picture is far more complex than what is
represented in Figure 5.3. The figure, however,
gives a broad idea of the life history of the group.
Figures 5.4 and 5.5 show a few trends.

Q Eqs
Hdn Eqs
P
Hdn. gr.
Hpn

M Plp
Ptp Hpn. gr.
Myp
Hyp
Arp

Prp
O Anc
Mop

Pth

Epp
E

Orp

Eop/Hrc

Fig. 5.3 Diagrammatic phylogenetic tree of Equidae showing the two main feeding habits, viz. browsers
and grazers, a few important genera and a few generic groups.
Abbreviations used:
E (Eocene) O (Oligocene) M (Miocene) P (Pliocene)
Q (Quaternary) (Grazers) (Browsers) Eop/Hrc (Eohippus/Hyracotherium)
Orp (Orohippus) Epp (Epihippus) Mop (Miohippus) Anc (Anchitherium)
Prp (Parahippus) Arp (Archaeohippus) Hyp (Hypohippus) Myp (Merychippus)
Ptp (Protohippus) Plp (Pliohippus) Hpn (Hipparion) Hdn (Hippidion)
Eqs (Equus) Pth gr. (Palaeothere group) Hpn gr. (Hipparion group) Hdn gr. (Hippidion group)
 Chapter 5 Evolution of Organisms 89

Equus
Q
One-toed

P Pliohippus
Gr.

Merychippus

M
Parahippus

three-toed
Mesohippus
O Br.

Hyracotherium
E

Fig. 5.4 Equid evolution.

The animal, front leg and toe-characters against change in habit from browsing to grazing. Change in
body size is broadly representative; three-toed front leg characterize the browsers and early grazers;
one-toed front leg found in grazers. Approximate geological ages are given in the same abbreviations
as in Figure 5.3.

(i)

(ii)

(i) (i)
(ii)
(ii)
(iii) (iii) (iii)
(a) (b) (c)

Fig. 5.5 Equid evolution.

(a) Skull, (b) Crown surface and (c) Hypsodonty of teeth through three stages, viz.
(i) Eohippus, (ii) Parahippus and (iii) Equus, the first two browsers and the third grazer.
The vertical lines in (a) indicate lengthening of skull in front of the orbit and deepening of the skull to
accommodate high-crowned teeth.
Crown surface (b) is shown with the teeth of left upper jaw; in Eohippus and Equus the last premolar
and the first molar are shown, in Parahippus only the molar. Molarization of premolars and complexity
of crown surface are to be noted.
In (c), brachydont habit of teeth of browsers and hypsodont habit of grazers are highlighted.
For more details about teeth see Appendix 2.
90 Part One: Principles
5.7.5 Migration and evolution of
equidae
The phylogeny as shown is best represented in the
fossil record of North America. The Old World
(Europe and Asia) fossil record includes sharply
different genera without intermediate forms. It
appears that the evolution proceeded in a series of
abrupt steps. In Indo-Pak subcontinent too, the
Siwalik deposits contain Hipparion in Dhok Pathan
Formation and then Equus in Tatrot-Pinjor, the two
genera being widely separated in equid phylogeny.
The problem is explained from the North
American record of true phylogeny which indicates
that the saltations are peculiarities of fossil record
and not of evolution. Different genera migrated
from North America to the Old World as also to
South America at different times in the history of
the group and the emigrants made their appearance
in those continents without leaving any transitional
forms in the sequence there.
Equids, like many other mammals originated
simultaneously in both the Old and the New
Worlds. Hyracotherium is found in Europe,
Eohippus in North America, both from Lower
Eocene rocks. In Europe Hyracotherium became
extinct; but in North America the equine evolution
proceeded successfully giving rise to a very
complete sequence of forms.
The region in which the most continuous
record of fossil equids has been discovered in
North America lies in the western part of that
continent in the Rocky Mountains and the high
Great Plains immediately to the east. Marked
changes in environmental conditions, such as in
topography, climate and vegetation, took place in
this region during Cenozoic.
Orogeny producing the Rockies started in late
Cretaceous-early Cenozoic. Broad valleys in
between the newly formed mountain chains housed
equids of the area as well as the sedimentation
basins, in which the equid remains were entombed,
mostly in fluvial deposits. In Oligocene, deposition
may have ceased there temporarily, taking place
to the east in what is now the High Plains. The
fauna, including the equids, too, shifted eastwards
to the latter region.
A climatic change was imminent there with a
decrease in temperature and rainfall. The cause of
temperature change is not known, but the drop in
rainfall may be correlated with the rising Rockies
producing a rain-shadow area to the east, in the
then low-lying ‘High Plains’. This brought about
changes in vegetation too, which was, however,
partly due also to the appearance of grasses, or
flowering ground plants at that time.
Eocene horses lived in subtropical forests as
it is evident from their different characters. The
crown pattern of their molars are not very different
from that in the primate molars. Thus, they
probably lived on a diet of succulent parts of
plants.
Towards the end of Miocene the grazing habit
evolved along with gradual changes in cheek tooth
crown pattern; functionally the changes were
directed to increasing the number of ridges on the
crown, spacing them more closely and aligning
them transverse to the length of the jaw so that
they could be most effectively used by a lateral
grinding motion of the lower jaw. Hyposodonty
and the amount of cement deposited between the
crown ridges, increased together at an accelerated
rate during this time. Digits (toes) in foot changed
into one, with springing mechanism.
In Pliocene, woodlands probably became
restricted along river banks and as ‘discontinuous
islands’ of forest in a region of a prairie grassland.
Grazers perfected themselves with selection;
browsers became extinct. Temperature dropped
further, culminating in a sequence of four advances
and retreats of subarctic conditions in the region,
along the southern margins of Pleistocene ice-
sheets sited further north. Large size has
thermodynamic advantages, and probably the small
Pliocene horses (Nannipus) lived in more southerly
latitudes. Certainly, only the large horses,
Hipparion and Equus, crossed the northern Bering
Ridge into Eurasia in Pliocene and Pleistocene.

5.8 Family Hominidae
5.8.1 Classical view on humans in the
organic world
Hominidae is a family of primates which includes
humans and their related organisms, the modern
man being Homo sapiens sapiens, that is the
sapiens variety of the species H. sapiens.
Classically, the family included two genera
Australopithecus (meaning southern ape man) and
Homo (meaning man). A third genus Ramapithecus
was thought to be the earliest hominid, ancestral
to Australopithecus, the latter, in turn, being
ancestral to and appearing earlier than Homo.
In so far as morphology, anatomy, behaviour
mode of living, etc. are concerned, apes are the
closest organisms to these hominids. It was on this
closeness that the ape-ancestry of man was
suggested by Darwin. So, to understand evolution
of humans, apes would naturally figure out in the
discussion. Apes include the lesser apes and the
great apes.
As an organism, modern man and its related
organisms belong to the Order Primates, in which
the Suborder Anthropoidea (‘human’-like)
includes ‘higher’ primates, monkeys and apes. It
has two groups (Infraorders/Cohorts) that evolved
independently; they are Platyrrhini or New World
primates (of mainly South America) and
Catarrhini or Old World primates (of Africa, Asia,
Europe). Hominoidea, a superfamily under
Anthropoidea and then Catarrhini, includes the
apes and humans, modern and fossil. Hominoidea
was divided into Hylobatidae (gibbons, the lesser
ape), Pongidae (the great apes, including gorilla,
chimpanzee, orang-utans, etc. and their fossil
equivalents) and Hominidae. The last family is
subdivided into two subfamilies, viz. Australo-
pithecinae and Homininae. As mentioned,
hominids included two genera Australopithecus
and Homo, belonging respectively to the two
subfamilies.
Chapter 5 Evolution of Organisms 91
At one stage of study of human evolution, it
was believed that the great apes and hominids
diverged from a common ancestor at around
12–18 million years before present. Ramapithecus
was the representative of the hominids at that stage.
Australopithecus appeared then; it was divided into
a gracile (species afarensis and africanus) and a
robust type (boisei and robustus). Homo was
derived from the gracile type, H. habilis being the
first species of the genus.
This idea and knowledge about the hominids
may be best described as in a constant state of flux.
With more and more discoveries of fossils and
newer and newer techniques, particularly those of
molecular biology, as well as emergence of newer
concepts like cladistics and phylogenetic
systematics, these classical ideas have been
drastically changed. In such a situation, it is rather
undesirable to repeat the prevailing earlier views
uncritically. In the present discussion, the author
intends to include as much updated information as
possible within the limit of this chapter. Sometimes,
it may appear a little confusing with the two
conflicting views, earlier and changed. But for the
sake of presenting a truer picture, as at present
conceived and which may very well change in the
future, we should be acquainted with how the views
changed or from what to what.
5.8.2 Tertiary apes
As mentioned, tracing of human ancestry would
require knowledge about the apes. Origin of the
broad group Anthropoidea, to which all fossil and
modern apes and humans belong, is still debated.
The traditional view favoured origin ‘from Africa’;
new finds from China and Thailand put it to Asia,
though the latter reports have been questioned
(Benton 2005, Box 11.2, p. 369). The early
anthropoid fossils come from Middle Eocene to
Oligocene rocks.
Anthropoids were diverse in Miocene of
Africa. There were two families of ape-like
92 Part One: Principles
organisms, viz. Dryopithecidae and Rama-
pithecidae. The former appeared earlier. The genera
included the following:
1. Proconsul of Lower Miocene was a quadruped
animal, yet possessing synapomorphies viz.
the absence of tail and relatively large brain
size, sharing them with true apes.
2. Kenyapithecus, also of Lower Miocene is
found from East Africa, Turkey and central
Europe. It was thought to be among the earliest
Hominids, but now considered related to
orang-utans.
3. Sivapithecus (= Ramapithecus) is reported
from Northern India, Turkey, Pakistan and
China. This genus was also like orang-utans,
being adapted to feeding on tough vegetation
and quadruped in locomotory habit. It might
have also been climbers on trees.
Lufengpithesus, Griphopithecus, etc. were
similar forms described from China and
Turkey of Middle to Upper Miocene. At one
stage of studies, Ramapithecus was considered
the ancestral or first hominid; but that was
based on scanty fossil material of the genus,
and that, too, of some non-diagnostic teeth,
etc. Subsequent discovery of adequate and
more diagnostic fossils of skull, etc.
particularly from China, prompted complete
revision of the idea and group the genera as
close to apes of the orang-utan type.
4. Dryopithecus reported from Spain to Hungary
in Europe might have been a contemporary or
slightly older genus.
5. Gigantopithecus was a massive animal
reported from late Miocene of India (species
being G. giganteus); and late Pleistocene of
China (G. blackii). It is thought to have lived
in wider stretches of South East. Incidentally
this giant ape might have triggered the
imaginative stories of Yeti or King Kong.
Fossil record of apes is almost barren or poorly
studied, between latest Miocene to earliest
Pliocene. In Plio- Pleistocene, ape fossils continue
to be scarce, though hominid fossils (as defined
previously) are well represented.
5.8.3 Changed view on phylogeny
Phylogenetic systematics or cladistics and
molecular phylogeny has significantly altered these
ideas on systematics and phylogenetic relationships
of different hominoid groups, even genera, as also
the ideas on ancestry of humans (see Factsheet 5.2;
see also Lewin 1984; Jones Martin and Pilbeam
1992; Benton 2005; also see Figures 5.7 and 5.8).
Hominidae is now considered as one of the
three families of Hominoidea; the other two are
Proconsulidae and Hylobatidae. By this
definition, Hominidae includes both the great apes
and the humans, in a lineage that branched off in
early Miocene. No fossils representing this stage
has yet been discovered. However, Proconsulidae
developed subsequently to give rise to the genera
mentioned in section 5.8.2.
Living Hominidae have two subfamilies:
Ponginae (orang-utan and its fossil relatives) and
Homininae (gorilla, chimpanzee and humans) that
arose from a tree-climbing ancestor and differed
between themselves in locomotion. Thus,
Ponginae animals move by suspension from trees
with long arms (brachiation) and slow climbing
and Homininae were quadrupeds on land (gorilla,
chimpanzee) or biped (humans).
5.8.4 Different genera and species of
‘hominids’
The two former subfamilies Australopithecinae and
Homininae are now grouped in Hominini, the
genera Australopithecus and Homo being
considered as ‘sister groups’. In fact, the genus
Australopithecus is now named as different genera
by many authors, such as Sahelanthropus
tchadensis, Ardipithecus ramidus, Praeanthropus
anamensis, Praeanthropus afarensis, Australo-
pithecus africanus, Paranthropus robustus,
 Chapter 5 Evolution of Organisms 93

FACTSHEET 5.2
Changed Views on Ape-Human Relations and Human ancestry

Previous Idea Changed Idea

Present
Lesser apes Great apes Hylobatidae
Human (Homo)
Ponginae
Australopithecus, etc.
Gorilla
Chimpanzee Human
Gigantopithecus
5 Ma Giganto-
pithecus Homininae
Sivapithecus
Ancestor to Hominids
10 Ma

Ramapithecus
Dryopithecidae including Ramapithecus 15 Ma
Sivapithecus
Proconsul

20 Ma Proconsulidae

Early apes including 25 Ma

> 25 Ma Early apes
Aegyptopithecus

Paranthropus boisei and Paranthropus aethiopicus
(Factsheet 5.3). Australopithecus and the other
genus Paranthropus, as also the newly added
Praeanthrpopus (species of the two latter genera,
earlier identified as belonging to Australopithecus)
were roughly the same size as chimpanzees with
their brain slightly larger than that of the latter. Like
chimpanzees, they were climbers, yet they could
walk upright like humans.
Australopithecus perhaps coexisted with
Homo for a certain period; the former was extinct
around 2.3 million years ago, while Paranthropus
may have continued still later up to about 1
million year ago, while the genus Homo made its
appearance around 2.4 million years ago. Which
species of Australopithecus was the immediate
ancestor of Homo is not yet known, though a
gracile or light-built variety of species like
A. africanus is held as the likely candidate.
Different species of Homo are Homo
habilis, H. rudolfensis, H. erectus, H. ergaster,
H. heidelbergensis, H. neanderthalensis and
H. sapiens. A new species has been added to the
list very recently. It is Homo floresiensis from
Indonesian island, which may be an extinct human
species marooned there, while mainstream H.
sapiens sapiens colonized the mainland. The new
species is said to be 18,000 years. old and is a dwarf
species, being a 90 cm tall female. However, doubts
have been expressed if it is a dwarf variety of Homo
sapiens or even Homo at all.
The above enumeration reveals continuing
changes in the knowledge about these animals, as
more and more fossils are found and studied.
Factsheet 5.4 shows distribution of these species,
particularly the major and better-known ones. It,
however, points to a dominantly African occurrence
of earlier species, contrasted by gradual spread of
the species of Homo throughout the world.
94 Part One: Principles

4 3

(a)

2
8 6 8
9 5

10
6
7

11
12

13 (c)

(b)

Fig. 5.6 Major bones of vertebrate skeleton.

(a) Terrestrial quadruped, (b) Biped erect human, (c) Biped nonerect ape
Number index: 1 cranium/skull, 2 maxilla-premaxilla/ upper jaw, 3 mandible/ lower jaw,
4 cervical vertebrae, 5 humerus, 6 radius,
7 ulna, 8 pelvic bone,
9 carpals, metacarpals, phalanges, 10 femur,
11 fibula, 12 tibia,
13 tarsals, metatarsals, pahalanges.

5.8.5 Evolutionary stages: Are there any?
Such being the representatives of the precursors
of humans and their distribution, we may now look
at some patterns of hominid evolution. We deal
with the Hominini of the hominids, meaning
Australopithecus and Homo. Their evolution
shows some major steps, phenotypic in earlier parts
and cultural later. In fact, previous authors
demarcated three stages in this course of human
evolution. They were as follows:
Stage I: Australopithecus (and related genera
later separated from it) and the earliest species of
Homo, i.e. H. habilis: ~ 6–4 million years to ~1.6
million years (end of habilis; later views, however,
exclude habilis from this stage).
 Chapter 5 Evolution of Organisms 95

4.0 5.0 Qy 1 2 3 4 5
0 0

2 P 6 2
7
8
10 M 9 10
10 [E]
[D]
11
20 20
[A] [B] [C]
Ol
30 30

40 40

E
50 50

60 Pl 50

mybp 0 5 15 25 –1.0 +1.0 3.0 5.0 mybp

MT 0/1k

Fig. 5.7 Hominid- ape phylogeny and its environment.

(A) Broad pattern of temperature variations, (B) Oxygen isotope ratio, (C) Time scale in million years,
(D) Broad phylogeny of great apes and hominids, (E) Some major extinct apes.
Index: MT (Mean temperature in degree centigrade at middle latitudes), O/1K O18: O16 (ratio per
thousand mybp million years before present), Qy (Quaternary), P (Pliocene), M (Miocene),
O (Oligocene), E (Eocene), Pl (Palaeocene).
1 Hominid Homo and Australopithecines, 2 Chimpanzee Pan, 3 Gorilla Gorilla,
4 Orangutan Pongo, 5 Gibbon Hylobates, 6 Gigantopithecus,
7 Lufengpithecus, 8 Sivapithecus, 9 Dryopithecus,
10 Kenyapithecus, 11 Proconsul.

Stage II: Particularly H. erectus of ~1.8–0.3
million years.
Stage III: Includes H. sapiens, along with
its variety neanderthalensis (now considered a
different species), and the H. sapiens sapiens
variety of modern man since 130,000.
5.8.6 Species of Australopithecus and
Homo contrasted and compared
But before assessing these stages, it may be useful
to look at the essential features of Australopithecus
and Homo, as well as how were these ancient
organisms related to modern humans differed from
apes. Factsheet 5.5 shows that the four better
known australopithecine species, separate into
two types. Praeanthropus (Australopithecus)
afarensis and Australopithecus africanus are
lightly built and are, thus, called ‘gracile’, and
Paranthropus (Australopithecus) robustus and P.
boisei, are heavily built and are known as ‘robust
types’. Factsheet 5.6 brings out a few differences
between Australopithecus and Homo; they pertain
96 Part One: Principles

Anomaly
Epoch
K-Ar N
age R Polarity Epoch (P) - Event (V)
0 1 2 3 4 5 Normal (N) -Reverse (R)
BRUHNES (P/N)

Pleistocene
1
1 Jaramillo (V/N)
5
MATUYAMKA (P/R)
Olduvai (V/N)
2 Reunion (V/N)
10 (a) (b) 'X' (V/N)

Pliocene
2
3 Kaena (V/R) GAUSS (P/N)
Mammoth (V/R)
15

4 3 Cochiti (V/R) GILBERT (P/R)

20 4-5 2-3 1 0 4-5 2-3 1

5 12b
13 13
25 15
(c)
12a
25
(i) (ii)
Fig. 5.8 Hominid-ape phylogeny in the backdrop of climatic-palaeomagnetic events.
(a) Phylogeny, (b) 0 to 4 million years time before present enlarged to show palaeomagnetic events,
(c) Hominid-ape divergence: (i) Classical view and (ii) recent view on molecular biology
Number index: 12(a) and 12(b) probable positions of common ancestor, 13 - Ramapithecus. Other
numbers same as in Figure 5.7.

FACTSHEET 5.3
Australopithecus and Homo: Brief Introduction

I. Australopithecus:
1. Brain: small in size, 375-600 cc; brain weight relative to body weight greater than in apes; no
Broca’s area, nor Wernicke’s area.
2. Posture-gait-frame: Upright posture; bipedal standing or running gait, though both imperfect. Hands
freed from locomotion; vision made stereoscopic helping more precise reading of depth and distance
and putting more pressure on brain.
Bowl-shaped pelvic girdle; distinct s-shaped lumber curve and forward foramen magnum; leg limb
segments shorted than their counterparts in arms.
3. Cranium-dentition-face: smooth parabolic dental arch; no simian gap, premolar non-sectorial; large
teeth and jaw; non-projecting canine; prognathous face, no chin; heavy eyebrow ridges, low slanting
forehead.
4. Culture: tool-making (?), at least tool-using; family life questionable.
(Cont...)
 Chapter 5 Evolution of Organisms 97

FACTSHEET 5.3 (Cont...)

Australopithecus and Homo: Brief Introduction

II. Homo habilis: Brain 680 cc; Broca’s and Wernicke’s areas; profuse tool-making; variety of tools for
varied purposes; family, even social life, though questioned.
H. erectus: Brain 775–1225 cc; low slanting forehead; fire making, conserving and using.
H. sapiens (archaic)/H.heidelbergensis: Brain 1100-1400 cc; eyebrow ridge in some individuals;
forehead low/high slanting.
H. sapiens neanderthalensis/H. neanderthalensis: Brain 1200–1600 cc; shorter lower limb; low
forehead; eyebrow ridge present; skull and face more human like than simian; wore clothes from hides;
made shelters; could make articulated speech; performed rituals.
H. sapiens sapiens: Brain 900–2300 cc; (av. 1350 cc) bipedal standing and running; frontal lobes in brain;
skull widest above ears; small teeth and jaw; orthognathous face; chin formed; no eyebrow ridge; high
forehead; clan society formed.
III. A few recently added or revised genera and species
Revised names of some Australopithecus species
Sahelanthropus tchadensis Ardipithecus ramidus
Praeanthropus anamensis P. afarensis
Australopithecus africanus
Paranthropus robustus P.boisei
P. aethiopicus
Species of Homo
Homo habilis H.rudolfensis H.erectus H.ergaster
H.heidelbergensis (earlier ‘archaic’ sapiens) H. neanderthalensis H. sapiens

mainly to brain, but more significantly to loco-
motory, feeding and living habits.
Factsheet 5.7 presents characters of early
species of Homo. On comparison with those of
Factsheet 5.5, these bring out further the
differences between Australopithecus and Homo.
They also suggest changes in the characters of
species of Homo, as the genus evolved and its
species appeared successively in the broad series
of: Homo habilis, H. erectus, archaic or primitive
H. sapiens, H. neanderthalensis and lastly
H. sapiens sapiens.
5.8.7 Differences between apes and
man: Phenotypic distinction of
humans
It has already been mentioned that a direct
descendancy of man from the modern apes was
envisaged earlier, even by Darwin. This is now
ruled out. Rather, both the African apes and the
human appear to have a common ancestry in
Miocene. Of the two African apes, chimpanzee
seems to be closer linked to humans, as inferred
particularly from molecular biology. This is an
issue that will be taken up in a later section. Here,
it may be indicated that notwithstanding molecular
biological differences, modern apes and modern
man show important similarities as well as
differences. The former suggests their commonness
in origin and the differences help understand the
trends in which a man would have evolved.
Apes and humans resemble each other in a
bipedal gait, albeit imperfect in apes; frontally
directed vision, a result of the gait itself; broadly
similar dentition types; larger brain in relation to
the body weight as compared to that found in any
other organisms; greater and longer extent of care
for new borns; similar type of living in small
groups, etc.
 98
FACTSHEET 5.4
Principal Fossil Sites of Hominids Around the World

Sites Australopithecine Homo

Gracile / Robust habilis / erectus / ‘archaic’ sapiens/ Neanderthal/ Early sapiens/ sapiens/ Rock art
AFRICA Hadar y
Middle Awash y y
Omo River ?y y y y y
LakeTurkana/Koobi Fora ? y y y y y y
Part One: Principles

Olduvai Gorge y y y
Laetoli y y
Starkfontein y y
Swartkrans y y
Taung y
ASIA Caves (South Africa) y y
Zhoukoudian, China y y
Dali, Maba (China) y
Lantian (China) y
Longgu (China), Phillippines
Uzbekistan, Iraq, Israel y y
Sri Lanka, Niah (Indonesia) y
Trinil, Sangiran (Java) y
Narmada (India) y
AUSTRALIA Willandra Lake (Australia) y
Koonalda cave (South Australia) y
EUROPE Swanscombe (UK) y y
Steinheim (Germany) y y
Neandertal (Austria) y
Altamira (Spain) y
Lascaux, etc. (France) y
Vogelherd (Germany) y
Grotte du Prince (France) y
Pedra Furada and Pema (South America) y
<30000 yr
SOUTH AMERICA <30000/14500yr
NORTH AMERICA
 Chapter 5 Evolution of Organisms 99

FACTSHEET 5.5
AUSTRALOPITHECINE Species Compared

Australopithecus A. africanus Paranthropus P. robustus

afarenis boisei
Height (m): Weight (kg)
1-1.5:30-70 1-1.4:30-60 1.2-1.4:40-80 1-1.3: 40-80
Physique Light build; some ape Light build; probably Very heavy build; Heavy build; relatively
features (e.g. shape of relatively long arms; relatively long arms; long arms; moderate
thorax; long arms more ‘human’ features; marked sexual sexual dimorphism
relative to legs; curved probably less sexual dimorphism
fingers and toes; dimorphism
marked to moderate
sexual dimorphism)
Brain size (ml) 400-500 400-500 410-530 530
Skull form Low, flat forehead; Prominent crests top Crest on top and Skull; long broad,
Higher forehead; of shorter face; brow back of very long, flattish face; moderate
projecting face; ridges less prominent broad, flattish face; facial buttressing
prominent brow ridges skull; strong facial
buttressing
Jaws/teeth Relatively large Small incisors-like Very thick jaws; Very thick jaws; small
incisors and canines; canines; no gap small incisors and incisors and canines;
gap between upper between upper incisors canines; large molar large molar-like pre-
incisors and canines; and canines; larger like premolars; very molars; very large molars
moderate-sized molars molars large molars
Distribution Eastern Africa Southern Africa Eastern Africa Southern Africa
Known date >4 - 2.5 3.0 -< 2.5 2.6-1.2 2-1
(million of
years ago)

FACTSHEET 5.6
Some Differences Between Australopithecus and Homo
Australopithecus Homo

Brain Smaller; 450 cc Larger; ~750-800cc

Habit Better climber Better walker
Worse climber than chimpanzees
Food Fed mainly upon fruits and seeds; also Had meat as diet; hunted/scavenged carcasses; associated
had small animals as food scratched bones and tools (Oldowan variety) attest
Living Some individuals lived together in a Less dependent on trees for feeding and security
grove of trees, fed in daytime, slept at
night on the trees; when food became
scarce, they moved away to a new grove
much like present-day baboons
100 Part One: Principles

FACTSHEET 5.7
Facts and Figures on Early Human Species (See Figure 5.10)

Homo habilis (small) Homo habilis (large) Homo erectus

Height (m) 1 ~ 1.5 1.3-1.5

Physique Relatively long arms Robust but ‘human’ Robust but ‘human’
skeleton skeleton

Brain size (ml) 500-600 600-800 750-1250

Skull form Relatively small face; Larger, flatter face Flat, thick skull with large
nose developed occipital and brow ridge

Jaws/teeth Thinner jaw; smaller, Robust jaw; large narrow Robust jaw in large
narrow molars molars individuals; smaller teeth
than H. habilis

Distribution Eastern (+ southern) Eastern Africa Africa, Asia and

Africa Indonesia (+ Europe)

Known date 2-1.6million 2.4-1.6 million 1.8-0.3 million

(years ago)

“Archaic Homo Neanderthals Early modern Homo

sapiens” sapiens

Height (m) 1.5-1.7 1.6-1.85

Physique Robust but ‘human’ As ‘archaic H Modern skeleton;

skeleton sapiens’, adapted for cold adapted for warmth

Brain size (ml) 1100-1400 1200-1750 1200-1700

Skull form Higher skull; face Reduced brow ridge; Small or no brow ridge;
less producing thinner skull; larger nose; shorter, high skull
midface projection

Jaws/teeth Similar to H. erectus, Similar to ‘archaic Shorter jaws than

teeth may be smaller H. sapiens’; teeth Neanderthals;
smaller except for chin developed; teeth may
incisors; chin in some be smaller

Distribution Africa, Asia and Europe and Africa and

Europe W. Asia W. Asia

Known date 400,000-100,000 150,000-30,000 130,000-60,000

(years ago)

Factsheet 5.8 lists modern humans’ distinction
from primates. These include biological features
that determine perfect or erect bipedal gait in man,
the large human brain capable of improved brain
activities, dextral hands free from locomotion, and
capable of tool use and tool making particularly
by dint of possessing an opposable thumb in each
hand. Apart from these, there are differences in
feeding, in social and reproductive habits, and in
thinking and culture, etc. Factsheet 5.9 elaborates
important differences in the skeletons of modern
man and recent African great apes.
Synthesizing all these data of Factsheets 5.5
to 5.9, it may be summarized as follows. (Also see
Figures 5.6 and 5.9.)
Advent of Australopithecus was marked by
attaining a bipedal, upright or erect standing and
running posture; it meant development of:
1. erect posture of the body, in which the animal
could stand, and more particularly run, freely
without bending its knees. Apes can stand for
a while, but cannot run with erect posture;
2. basin shape of the pelvic girdle, which holds
the upper portion of the body in its position;
3. development of the S-shaped lumber curve
in the vertebral column that holds the heavy
skull right on its top;
FACTSHEET 5.8
Modern Humans Distinguished from Primates
Locomotion and Habitat
Bipedal walking
Open vegetation habitats
Brains, Thinking and Culture
Large brain capable of thinking
Tool use
Language as articulated symbols of thoughts
Development of cognitive skills
Information transmitted within and across
generations via thoughts, language and skills.
Chapter 5 Evolution of Organisms 101
4. shifting of foramen magnum (the opening
through which the spinal chord enters the skull)
towards the anterior, which prevents body from
stooping from the weight of the skull:
5. forelimbs (now turned hands) shortened to
become shorter than hind limbs (now turned
legs); the latter requires to be longer to hold
the body up efficiently;
6. forelimbs freed from locomotory function; on
the other hand, with development of free-
moving, opposable thumbs in each hand
(forelimb), helped perform skillful activities
such as dexterous tool-using and-making and
7. shifting of eyes from side of the skull in
quadrupeds to its front in erect, permanent
bipeds, led to development of stereoscopic
vision, which meant improved vision with more
and more perfect assessment of depth of field
in view, the distance of the object and its three-
dimensional shape and size. This, in turn,
helped in developing improved brain activities.
However, all these traits did not develop at one
stroke, even at the earlier stages of evolution of
Australopithecus.
In addition, to these developments in posture
and gait, there were changes in the dentition
pattern, as follows:
Diet and Foraging
Animal protein
High amounts of starch
Food sharing
Social and Reproductive Aspects
Infants dependant for longer span of life
Females and males strongly bonded on
economic and sexual bondings
Social life based on home
102 Part One: Principles

FACTSHEET 5.9
Differences Between Modern Man and Great African Apes

Modern man Great African apes

Erect posture of the body, in which the animal could Apes can stand erect for a while, but cannot run
stand, and more particularly run, freely without with erect posture.
bending its knees.
Basin shape of the pelvic girdle to hold the body. Pelvic girdle upwardly elongated and slanting.
S-shaped lumber curve in the vertebral column holds No lumber curve.
the heavy skull right on its top.
Foramen magnum (the opening through which Foramen magnum posterior; the heavy skull cause
the spinal chord enters the skull) shifted towards the the body to stoop forward.
anterior; the heavy skull sits on its top and the body
is prevented from stooping.
Forelimbs (upper limbs, i.e. arms) shorter; longer Arms longer as in ancestral brachiating arboreal
hind limbs (legs) help hold the body up efficiently. habit of primates.
Smoothly parabolic dental arch. U-shaped dental arch of apes.
Loss of simian gap between incisor and canine. Simian gap found in apes.
Non-sectorial first lower premolar. Sectorial first lower premolar.
Non-projecting canines. Projecting canines.
Higher ratio of brain size: body weight. Lower ratio of brain size: body weight.
Maximum width of skull shifted from the level of Maximum width of skull at the level of ears.
ears to above them.
High, vertical forehead. Low, slanting forehead.
Shorter jaw and smaller teeth. Longer jaw and larger teeth.
Concomitant orthgnathous face with a vertical front Longer jaw causes forwardly protruding prognathous
profile of face and sharp chin. face, no chin.
Thinner, delicate eyebrow ridges. Heavy eyebrow ridges.
NB: The changes did not take place at one and only one stage. Rather there were broadly different stages. Thus, posture and
dental characters changed earlier. Brain too. Finer details of skull and face changed later. Thus, there were
australopithecines that appeared intermediate between the modern man and apes (see Figures 5.6 and 5.9).

1. Smoothly parabolic dental arch, in place of
U-shaped dental arch of apes.
2. Loss of simian gap, the gap between incisor
and canine, found in apes.
3. Development of non-sectorial first lower
premolar.
4. Development of non-projecting canines.
A second trend of changes was related to brain.
Between African apes and man, the former has a
relatively smaller brain as compared to their body
weight. The ratio between brain size and body
weight was lower in the species of Australo-
pithecus. But it changed fast in the species of
Homo, meaning that successive species of the
genus had larger brains as compared to their body
weight. (Figure 5.11). Thus, there was increase in
cranium size from 450 to 600 cc in Australo-
pithecus and related genera to 775–1225 cc in

H. erectus and 1100-1450 cc in transitional forms
of H. sapiens (called Archaic sapiens, now
recognized H. heidelbergensis) further changes to
1200-1600 cc in H. neanderthalensis or 900-
2300 cc in sapiens sapiens occurred at a later stage.
In this connection, it may be added that more
significant than simple increase in size, there were
changes in the human brain that involved
development of a more complex and coordinated
organization, which could correlate and synthesize
the sensations to perceptions and conceptions and
from there to thought processes. The role of speech
in this process is now acclaimed important. In
H. habilis, it is found that the frontal lobe of the
cranium contains convolutions marking the advent
of Broca’s area and Wernicke’s area. These two
are now recognized as specific areas in human
brain that control speech and coordination-
synthesization processes, respectively. However,
in habilis, the larynx or the sound box was not
dropped down to a sufficient depth inside the
trachea, so as to be able to produce different kinds
of well-articulated sounds. Even then, the presence
of varied kinds of tools, presumably for different
types of uses, are found associated with habilis
fossils. Both the number and the varieties of these
tools point to a concerted effort that could not have
been possible without well-articulated
communication of a sort of planned programme.
However, later studies do not stand much in support
and it is suggested that speech and thinking, both
seem to have developed not in habilis but in the
next stage in H. erectus.
Cranial endocasts in fossil skulls of H. erectus
and the capacity of the species to make, conserve
and use fire (as found most distinctly and
spectacularly in the Chinese occurrence at
Zhoukoudian caves), suggest that thinking and
communication through articulated speech were
well-advanced in H. erectus.
Subsequent changes, often referred as changes
of third stage, involve specially the changes in skull
morphology (see Figure 5.10) mainly in response
to increase in cranium size, loosely in correlation
Chapter 5 Evolution of Organisms 103
with changes in brain construction and
organization.
1. Thus, maximum width of skull shifted from
the level of ears to above them.
2. Slanting forehead found in apes was no longer
there in man; it was a high, vertical forehead.
3. Jaw length was small and the size of teeth
decreased.
4. As a result, the forwardly protruding
prognathous face changed to orthgnathous
face with a vertical front profile of face.
5. There developed a sharp chin.
6. Heavy eyebrow ridges were lost to its thinner
more delicate shape.
These phenotypic distinctions of humans do
not, however, follow any persistent and distinct
trends. Rather, they developed in sets, each broadly
at certain point in phylogeny. Thus, the posture or
gait and dental characters changed first, even at
the stage of Australopithecus or early Homo.
Changes in brain took place then in the species of
Homo, particularly in erectus. Finer details of skull
and face changed later, largely at the stage of the
species sapiens.
Almost all the traits of anatomy possessed by
the modern man developed in Cro-Magnon man
that belonged to the same spcies as the modern
man. Subsequent changes were intraspecific. But
beyond anatomy, were the changes in levels of
cultural developments from the Stone Age (Pala-
eolithic, Mesolithic, Neolithic) (see Factsheet 5.10)
to the modern electronics. Present human varieties,
though apparently segregated and anatomically
different, are mere intraspecific variations; the
whole of the mankind belongs to the single species
in which populations can potentially and
successfully interbreed.
5.8.8 Human ancestry: Revisited
It has already been pointed out that the ideas on
ancestry of humans and their relations with apes
have changed with time and studies. The great apes
104 Part One: Principles

(a) (b) (c) (d)

(e) (f) (g) (h)

(i) (j) (k) (l)

(m) (n) (o) (p)

(q) (r) (s) (t)

Fig. 5.9 Ape and human contrasted.

(a) Rib cage of modern ape, (b) Rib cage of modern man, (c) Vertebral column of modern ape,
(d) Vertebral coloumn of modern man, (e) Pelvic girdle of modern ape, (f) Pelvic girdle of modern man,
(g) Hand of modern ape, (h) Hand of modern man, (i) Jaw (up.) and teeth of modern ape, (j) Jaw (up.)
and teeth of Australopithecus, (k) Jaw (up.) and teeth of modern man, (l) Jaw (lr.) of modern ape,
(m) Jaw (lr.) of Australopithecus, (n) Jaw (lr.) of modern man, (o) and (p) Skull of modern ape, (q) and
(r) Skull of Australopithecus, (s) and (t) Skull of modern man.
 Chapter 5 Evolution of Organisms 105

Australopithecus africanus

Homo habilis

Homo erectus

Homo sapiens (archaic)

Modern human
(a) (b) (c)
Fig. 5.10 Hominid species compared.
Skulls of Australopithecus africanus and different species of Homo compared in (a) front view,
(b) side view, (c) back view.

and hominids were once thought to have diverged
from a common ancestor at around 12–18 million
years ago during Early to Middle Miocene.
The great apes included both the African and the
Asian types, which were grouped together into the
family Pongidae. Later studies have led to conclude
that the Asian great apes, viz. the orang-utan
belonged to a different subfamily Ponginae which
separated from the African apes and humans
together. The latter, belonging to the subfamily
Homininae, then diverged from a common ancestor
during latest Miocene; Gorilla separated earlier and
Chimpanzee, held closer to humans than Gorilla,
separated next. Hominini, a sister group of the
Chimpanzee, included Australopithecus (including
the presently revised genera) and Homo. Figures
5.7 and 5.8 present the relevant phylogenetic trees.
Fossil finds have, no doubt, been of help in
arriving at this picture, though at the same time it
must be admitted that they have not said the final
words. Neither have the comparison between the
modern man and the modern apes provided the
106 Part One: Principles

1500

AH s

800 e (a)
bv
h
AG
500

NG NH afc ch g

0 50 100
bw
0
s
s 1
e
e
(b )
2
h h
3

A A

200 1000 0 80
bv bw

Fig. 5.11 Estimated brain volume bv (cm3) — body weight, bw (kg) of apes and hominids contrasted
mutually (a) and against time (b).
All values are approximately represented. Bars reveal markedly faster post-natal changes in human
than in gorilla, leading to socialization (compiled and approximated from different sources)
s (sapiens), e (erectus), h (habilis) (all of Homo), afc (africanus), rb (robustus) (of Australopithecus A)
ch (chimpanzee), g (gorilla). AH (adult human), NH (newborn human), AG (adult gorilla), NG (newborn
gorilla).

FACTSHEET 5.10
Stages of Human Evolution including Culture

Historical Archaeological Evolutionary Absolute dates

stage stage stage (thousand yrs.:K)

Primitive community Upper Palaeolithic, Modern man 40K–20K

Mesolithic, Neolithic
and Early Bronze
Advanced primitive horde Mid Palaeolithic Neanderthal 200K–40K
Early primitive horde Later stages of Pithecanthropus/ ~2,000K–200K
Lr. Palaeolithic Homo erectus
Primitive proto horde Early stages of Australopithecus 4,000K– ~2,000K
Lr. Palaeolithic

answers. In fact, fossils of any common ancestor
of apes and man are yet to be found. The
conclusions are now based on the study of
molecular evolution, from the study of proteins
such as fibrinopeptides, myoglobins, haemo-
globins, etc. as also from information on DNA-
DNA hybridization, which assumes that the DNA
evolutionary clock ran at about the same rate along
different lineages. It is now known that when two
species separate, the genetic material that is the
DNA in the two lineages develop changes or
mutations. The longer the separation time, the
larger will be the sum total of changes. As it is
assumed that the rate of the change remained the
same, it becomes possible to obtain a measure of
the time of separation of two species from a
common ancestor, from the measure of
accumulated biochemical differences. All these
studies on biochemical compounds, referred above,
amply demonstrate that humans and the African
apes are related closer to each other than they are
to the Asian apes and that humans are closer to the
Chimpanzee than to Gorilla. It is also inferred that
the separation of apes and humans took place at
around five million years before present, that is,
towards the end of Miocene or the beginning of
Pliocene. Absolute dating on some of the available
fossils of Australopithecus (or its related genera)
appears to reach at similar dates.
5.8.9 A probable scenario
In addition to this account of hominids and apes,
their phenotypic characteristics, distribution and
molecular biological aspects, we may visualize a
probable scenario of what happened at around 2.5
million years ago from the present that led to the
emergence of a creature with larger and improved
brain from an animal, bipedal, land-dwelling,
walking upright yet climbing trees habitually, an
animal that was named subsequently as
Australopithecus. The scenario is presented as
follows (see Stanley 1989, 1993 for more details).
Factsheet 5.11 shows the probable environments
Chapter 5 Evolution of Organisms 107
that prevailed at different sites of Africa from which
early hominid fossils have been recovered. It
demonstrates essentially the presence of a mosaic
of rivers and lakes, grasslands and forests in a
varying climatic set-up, often seasonal. It was in
such an environment in eastern and southern Africa
that the cradle of human evolution developed, in
which Australopithecines and early Homo might
have reigned. Several facts are further relevant in
this regard.
Factsheet 5.12 shows a reconstruction of living
habits of early humans. Based on studies of
behaviour and habits of aborigines or primitive
tribes, as also varied evidences of tools, etc.
associated with fossil finds, this reconstruction
reveals that answers to many questions on human
evolution may have to be sought in evidences or
studies beyond the simple, classical
palaeontological or even biological-biochemical
efforts. They may pertain to social and cultural
evolution of humans that are now considered not
only correlated with, but also stemming from and
controlled by the development and activities of
human brain. Without this understanding,
knowledge of human evolution for even a beginner
in palaeontology may remain inadequate.
It is now known that brains of all newborn
primates are quite large, about 10 per cent of their
total body weight. They also grow rapidly before
birth, in all these species. In monkeys and apes,
the rate slows down significantly after birth and
brains of adults are not much larger in result.
(Figure 5.11).
In Homo, however, the brain continues to grow
rapidly for about a year even after the birth; the
rate then drops down to a more moderate one. In
consequence of this prolonged rapid growth for
even a year, both human children and adult humans
possess a much larger brain than their counterparts
among apes and monkeys (about twice that of a
chimpanzee, at the adult stage). Larger size of the
human brain is also accompanied by a unique,
significantly different organization of human brain
that enables human to store, correlate and
108 Part One: Principles

FACTSHEET 5.11
Early Hominid Environments in Africa

Locality (age in mil yrs.) Palaeoenvironment

1. Transvaal, South Africa All were mosaic environments, with Makapansgat Member 3 and
(Makapansgat 3, Sterkfontein Member 4 being less open (more bush/woodland) than
Sterkfontein 4 and 5 Swartkrans Member 1 and Sterkfontein Member 5; this suggests a
Swartkans1, Kromdraai trend from wetter to drier conditions through time.
and Taung) (3–1.4)
2. Olduvai, Tanzania Salt lake with surrounding floodplains with seasonal streams and dry
(1.9<1) woodland savanna; the lake dries up from tectonic changes.
3. Koobi Fora, Kenya A freshwater lake with floodplains, gallery forest and dry-thorny
(3.3–1.4) savanna subsequently fluctuated from fresh to brackish.
4. Omo, Ethiopia Dry savanna flanking river banks with gallery forest and dry-horny
(3.3–1.4) savanna developed from a presumably forest environment.
5. Hadar, Ethiopia Lake and associated floodplain, with braided streams and rivers.
(3.6–2.6)
6. Laetoli, Tanzania Savanna woodland, with well-defined wet and dry seasons.
(3.7–3.2)
7. Middle Awash, Ethiopia Fluvial conditions, with extensive tectonic activity associated with the
(4.5–3.9) formation of the East African rift.
8. Tabarin, Kenya Lake margin, with locally variable savanna elements.
(5–4)
Thus, we can visualize a more or less open habitat, with grasslands and water bodies, with or without forests
around, to be the environment in eastern and southern Africa, the cradle of human evolution in which
Australopithecines and early Homo might have reigned.
Environment was a mosaic of
Water bodies Vegetation Climate Tect.
LF LB/S Rfld Sv Fs Wet Dry Ssn Ds
1. ---- - - - - y y y y6

2. --- - y y y y - y -
---- - y5 - - - - - y
3. --- y - y y3 y4 - - -

---- y y - - - - - -
4. --- - - y y? - - -
--- - - y y3 - y - -
5. --- y - y2 - - - - -
6. --- - - - y y y y y
7. --- - y - - - - - y (EAR)
8. y - - y y1 - - -
NB: LF, Freshwater lake; LB/S Brackish water or salty lake; Rfld, River with floodplain; Tect, Tectonic condition; EAR,
East African Rift; Sv, Savanna; Fs, Forest.
Ssn, Seasonal; y1, locally developed; y2, braided streams; y3, thorny savanna; y4, gallery forest; y5, drying lake; y6, dry
with time; Ds, disturbed (compiled from various sources) (y? uncertain)
 Chapter 5 Evolution of Organisms 109

FACTSHEET 5.12
Development of Living of Earliest Humans is an Unwritten Chronicle
of Initiative, Interaction and Interdependence

Division of labour

Women and children collect plant foods Men collect meat by scavenging and hunting

All carry cobbles (of volcanic and other rocks)

Activities at the living site primarily include

Stone tool-making Food processing Sharing of produce

and ideas

leads to

Increasing social and economic interaction and interdependence

exerting

Selective pressure for social, intellectual and communicative traits and skills

Increase in intelligence

leads to

Better technology Better communication More purposive social skills More complex living style

leads to

Increasing social complexity with more communication and interaction

leads to

Further increase in intelligence paving way for more developed, socially coordinated living and activities
110 Part One: Principles
synthesize its sensations and perceptions into
conceptions and ultimately to thinking. Thus, even
a human child can think and act accordingly, unlike
any other animal, including its kins, the apes.
A general slowdown of maturation process
brings about prologation of high rate of brain
growth, particularly in early life. Evidence of this
delay in also found in the fact that human baby
teeth are replaced by permanent teeth, when it is
much older than corresponding ape-babies. The
effect is that the brain attains the larger size, finds
longer time for developing its organization and for
developing the thinking faculty, so characteristic
of man. It is with the help of this faculty that man
overcomes his disadvantages, a defenceless
condition; he neither has any means for quick
movement nor any teeth, claw or horn to resist
predators.
The question is: what could have made this
change happen, as it appears, in some, may be one
lineage of Australopithecus at around 2.5 myrs
ago from now. It was around that time when
continental glaciers had started expanding, as could
be inferred from different evidences, particularly
palynofossils. This drastic change in climate told
upon forests, which shrank in extent; grasslands
spread instead. Under such circumstances,
Australopithecus, habitual climbers for food and
security, faced ecological crisis and thence,
extinction. However, from one lineage of that
genus, there arose a new organism, which was
provided with an anatomy to sustain land-dwelling
without climbing through greater span of days. For
it, the squeeze of forests were not that alarming as
it was for its ancestors. This organism standing and
walking erect on two legs, and with hands free from
locomotion or tree climbing, could use them
otherwise. There was, thus, tool-making; also
among them, mothers could carry their rather
helpless infants more on their laps, may be delaying
the maturation process of these offsprings. The
delay, however, may have proved a boon; the brain
had longer time to grow. It must be added here
that the erect posture also helped the stereoscopic
vision of humans to become more perfect, thus
helping the brain, in turn, to receive more
sensations in a more precise frame of distance and
depth. Avoiding predators, developing hunting
tools and techniques, etc. among others, added
more advantages to these organisms of the genus,
Homo that surpassed the disadvantages of their
being helpless without speed and teeth-claw-horn
or with a prolonged infant stage.
The genus Homo had several species (see
Factsheet 5.4), of which only one survives today.
Of them, H. habilis and erectus are better known
traditionally; ‘Neanderthal man’ is considered a
species by some authors, or a variety of H. sapiens
by some others. Factsheet 5.13 summarizes the
important milestone events of the hominid
evolution.
5.8.10 Appendix: On man’s uniqueness
The above account of man’s evolution suggests
that man had the following links or continuities
with his ancestors, viz. (i) biological inheritance,
(ii) inheritance of a group or horde life, and
(iii) inheritance of a communicative system. At the
same time, he had differences or breaks with his
ancestors principally by way of (i) development
of the upright posture, the freeing of hands,
formation of a developed vocal tract, enlargement
and development of the cerebral cortex, all in the
anatomy, (ii) transition from a proto horde stage
to social organization, and (iii) transition from an
animal communication system to an articulated
expression (language) which allowed for more and
more developed power of thinking.
Of these, the last ones, viz. speech, language
and thinking developed thereupon, impart man
with the power that not only distinguishes him, but
helps him earn uniqueness, to do all these
exercises, as we have been doing over these pages,
to find and disseminate a truthful picture of the
world around him.
More concretely, the whole set of factors
characterizing and controlling the pre-human
 Chapter 5 Evolution of Organisms 111

FACTSHEET 5.13
Major Milestone Events in Hominid Evolutions

Late Pleistocene hominids Decreased activity; less robust skeleton

35,000–10,000 yrs Rapid improvement in stone tool technology, hunting, foraging
Further spread in habitats
Artistic expression
Cultivation and cattle use and raising

100,000–35,000 yrs Improved stone tool technology

Rituals with dead
High activity, robust skeleton

Early Pleistocene hominids Spread of habitats

1.5–0.1 mil. yrs Sign of thinking to make different types of tools
Fire making and conserving

Plio–Pleistocene hominids Bipedalism more perfected

2.0–1.5 mil. yrs Dexterous use of hand in tool use and making to get and
process food
Protein in diet from scavenging and hunting
Increased brain function, mobility and defence
Maturation rate changed

Earliest hominids First known tool-making

3-2 mil. yrs Lived on open savannas

5-3 mil. yrs Bipedal, occasionally arboreal

Lived in a mosaic of grassland, woodland and thick bushes

Hominid ancestors Origin in African equator

8–5 mil. yrs

stage of development (‘biology’), especially the 5.9 General Comments

developing anatomical features, brain
organization, conditions of living, horde mode of
life, mode of communication, etc. all interacted
in a long course. The interaction carried into effect
the elements which qualitatively transformed the
pre-human (australopithecine) into the human
stage. The latter emerged with its own set of
characteristic and controlling factors, especially
social mode of living, thinking ability, language,
consciousness, humanness, etc. Thus, human
characteristics involve many things other than
simple anatomical ones.
The above discussion on the evolution of the three
groups of well-known large land-dwelling
vertebrate animals brings out certain interesting
aspects of evolution. As it appears, both the
Equidae and the Proboscidea made their
appearance in Eocene and evolved rapidly in an
essentially forest environment; in the same forests
evolved and originated Anthropoidea, the group
of ‘higher’ primates including apes and humans.
The warm ambience of the earth that prevailed
since Cretaceous and reached maximum in
112 Part One: Principles

Palaeocene, or was represented by warm seas in Australopithecus need be recognized as different

Middle Eocene with proliferation of carbonates and
carbonate-secreting marine organisms, witnessed
a drastic change in climate with cooling at the EOB
(Eocene-Oligocene Boundary; see Factsheet 5.14),
rather Upper Eocene. It must have had its effect
on the large land–plant vegetation, represented in
forests and was followed by the growth and spread
of flowering ground plants, i.e. the grasses. The
change is remarkably represented in the
evolutionary picture of the three groups in question.
Proboscidean and primate records dwindled;
equids were changing their adaptation from forest-
living browsing to grassland-living grazing habits.
All the three groups were represented by newer
genera and species following this phase in Mio-
Pliocene. Deinotheres, the proboscideans that
continued remained conservative, but the
elephantiforms underwent significant radiation and
divergence. Equids were experimenting
successfully on grazing habits. While primates
were giving rise to hominoids, the precursors of
humans, at one stage some lineage gave rise to
Australopithecus, the southern ape-man discovered
first in South Africa from the fossil of child, called
Taung child. However, it is now appeciated that
many of the species originally ascribed to
genera. Be that as it may, Australopithecus and
related genera developed between 5 and 2 million
years known, and presented two morphotypes,
gracile and robust. The former were lightly built,
omnivorous and may be the ancestral line to Homo.
The latter robust, heavier in built and largely
herbivorous, are now recognized under
Paranthropus, a dead end in hominid evolution.
Appearance of Homo took place sometime during
Uppermost Pliocene–Lowermost Pleistocene, and
as discussed may have a bearing with the advent
of the Ice Age. The latter may also be related to
the dwindling of both the Equidae and the
Proboscidea, which in result are now represented
by a single genus and two genera, respectively.
Primates, particularly apes, have a better
representation in the post-Ice Age forests, though
human is represented by a single species, which
has, however, proved to be the ruler of the entire
organic world, by virtue of his unique
characteristics of mental faculty. The three-letter
word ‘man’, thus, represents an all-powerful
organism that not just delves into the mystery of
his own coming into being, but it aspires also to
solve the bunch of mysteries that connect the whole
organic world.

FACTSHEET 5.14
Broad Climatic Changes Through Cenozoic
1.6 mya Ice sheets form in northern hemisphere, The last ice age begins with several interglacial and
glacial periods. The last interglacial episode begain about 10,000 years ago and the Last Glacial
Maxima (LGM) occurred at about 18,000 years ago.
6.0 mya Rapid cooling takes place.
15 mya Antarctica; more moisture and copious snow-pack. Antarctic ice cap develops.
30-25 mya Antarctica much colder but still no ice cap.
35-30 mya First glaciers on Antarctica.
38 mya Rapid cooling of surface water in the south and of deep water everywhere; Terminal Eocene
Cooling Event.
<50 mya Slight cooling takes place; no glacier or ice cap on Antarctica.
> 50 mya Uniform climate and a warm ocean with abundant carbonate.
mya: million years ago
 6
Major Events of
History of Life
The much diversified organic world has a long
history (of about 3.6 to 3.9 billion years: Stearns
and Hoekstra 2001), which is characterized by the
origin and extinction of innumerable groups of
organisms as well as by many episodes-events and
interactions-interrelations. The present chapter
attempts at providing a very brief outline of some
essential aspects of that history.
6.1 Stages in the History of Life
Among the events in the history of life, a few
brought about fundamental changes. These stages
may, thus, be considered as major episodes in the
history of life on the earth. These can be marked
as follows:
1. Origin of life: With this event, a process was
initiated on the earth, which eventually came
to be known as organic evolution and which
produced the organic world of the past and
the present.
2. Origin of organic cell: Itself a major step,
it brought in its wake two phases. The first
113
was the appearance of cells without nucleus,
i.e. prokaryotic cells, and the second was cells
with nucleus, i.e. eukaryotic. The earliest
organisms were prokaryotic, though most
organisms now are eukaryotic.
3. Origin of multicelled eukaryotes: Right
from the appearance of multicelled
eukaryotes, the organic world assumed a
diversity. So what followed was the beginning
and continuation of diversification-adaptive
radiation.
4. Origin of hard parts or biomineralization:
The whole stretch of the history of life from
its origin to earliest diversification of
mutlicellular organisms covered nearly
85 per cent of the total span of the history.
During this whole course, organisms were
soft-bodied and dominantly micro-organisms.
But the rest, less than one-fifth of the history,
was dominantly the history of organisms with
hard parts.
5. Advent of life on land: Life originated in
aqueous environment, i.e. seas. As discussed
in section 3.4 and Factsheet 3.10, it had
114 Part One: Principles
advantages: buoyancy permitting greater
energy conservation and lesser need of
development of skeletal parts. Yet the next
major step in the history of life was the advent
on land. It provided larger variety of habitats
thus, giving rise to more varied adaptation.
6. Diversification-adaptive radiation on land
and in water in response to ever changing
environment on the surface of the earth, with
land-sea distribution changing with
movement of continental plates, along with
changes in sedimentational patterns, climate
and other factors.
6.2 Some General Difficulties
A palaeontologist seeks to unfold this history of
life. The task is, however, neither smooth nor easy.
As already discussed, (i) fossils are the main tools
of a palaeontologist, and (ii) the fossil record is
biased or incomplete for taphonomic reasons.
It has also been indicated particularly in connection
with the evolution of vertebrates, including
hominidae, that evidences from molecular biology
and such other branches have helped solve many
problems of palaeontology and history or evolution
of organisms.
It may be reiterated here that the question is
specially significant on questions related to the
earliest parts of the history of life. As it has been
stated, nearly 85 per cent of the history of life
involves dominantly micro-organisms and
exclusively soft-bodied forms. Besides, this is a
history of a time period which measures in billions
of years for most of its part, from the origin of
life some 3.6 to 3.9 billion years ago to about
800-900 million years ago. There is every
possibility that the record is affected by strong
diagenesis, deformation and even metamorphism.
So, palaeontologists had to take help of other
branches of science to know the earliest history
of life.
6.3 Origin of Organic World and
Early Stages of Its Evolution
6.3.1 Two fundamental characteristics
of life
Life is characterized by two essential features
among a few others. They are, namely;
(i) metabolism of a living thing, which is a
‘coordinated system of chemical reactions’ that
contribute to and help in the maintenance of the
living thing itself, and (ii) reproduction or
‘hereditary replication’, which is ‘a system of
copying’ in which the new product resembles the
old (quotes are from Stearns and Hoekstra 2001).
Hence, origin of life must be hinged at a point,
when on the surface of the earth there appeared
some such thing that could initiate and perform
the two functions, referred above. To carry on the
chemical reactions of metabolism, the living thing
would need ingredients or materials, as well as
energy from the environment around. At the same
time, these chemical reactions themselves would
produce new materials for the sustenance of the
living thing and reproduction, as well as energy
released from the process.
6.3.2 Experimental verification of
pre-biotic happenings
In the 1920s, Oparin and Haldane acting
independently suggested that under a reducing
atmosphere, a variety of organic compounds could
be synthesized in an aqueous solution, with
energy supplied by lightning, ultraviolet radiation,
hot springs, lava outpourings, solar heat, etc. The
condition came to be known as the ‘hot dilute
soup’, which was subsequently reproduced in a
famous experiment of Miller in the 1950s.
A mixture of several gases such as methane,
ammonia, water vapour and nitrogen, and some
simple inorganic compounds were kept in a water
solution and subjected to electric discharge
 Chapter 6
simulating lightning. It produced a few
biologically important organic compounds, like
amino acids, which were the building blocks of
proteins, while the latter were important
ingredients of the living systems. Some nucleotide
bases and carbohydrates were also produced in
later experiments.
Both theoretically and experimentally, it was
then proved that life might have originated at a
certain stage of the physico-chemical actions-
reactions and evolution of the then non-living
world, referred as a ‘pre-biotic environment’,
when some such organic compounds with
potentialities for growth and replication were
formed from the available existing materials under
the influence of and controlled by natural sources
of energy, like radiations, in a reducing atmosphere
without oxygen. Those compounds combined
among themselves, guided by their structure and
characteristics. And through that they gave rise to
newer and newer compounds which could acquire
and assimilate necessary ingredients, elements and
compounds from the surrounding nature and could
build up, on that basis, substances that were
required for their sustenance, and at the same time
create energy for carrying on these chemical
actions and reactions. Also, at some stage in the
span of their existence they replicated themselves,
i.e. they could produce newer compounds
like themselves. Self-sustained growth and
development of the existing material or body and
replication became the two characteristics that
were not to be found in any other material of the
surface of the then earth. These compounds were
not organisms in the truest sense, but they were
the precursors of life no doubt.
They were fundamentally different from the
hitherto prevailing ones. It was not just one more
type of material; it was something qualitatively
different which initiated a fundamentally different
process on the earth so far unknown the process of
life and living. The emergence of these compounds,
rather the change from the already existing
Major Events of History of Life 115
compounds to these new ones, took place under a
set of unique conditions, which was not to be
repeated any further. It was because the products
of these processes once formed, themselves added
new materials to the environment, which set on
new processes of life and living, thereby creating
a separate domain within the existing world of non-
living things.
It must be added that many biologically
significant compounds like nucleic acids (RNA and
DNA), which are the chemical basis for
reproduction and heredity, have never been
obtained from any experiment. So how they could
have formed in the prebiotic environment and many
other important processes are not yet known.
Besides, it has been suggested that the initial
environment would have developed on the surfaces
of pyrite crystals or surfaces of droplets of clouds,
and not in an aqueous solution. The points remain
still unsettled, though without negating the main
premise of formation of organic molecules from
the inorganic world.
6.3.3 Cells and organic evolution
Nevertheless, the set of new compounds led to the
formation of organic cell, another different kind
of entity on the surface of the earth. It not only
continued the processes that had already begun;
but created an internal environment within its
confine that ensured continuation of the processes
like nutrition- growth and replication. It was again
not just that the organic compounds that were
interactively associated or combined within the
confines of the cell-membrane, did make another
kind of substance on the earth. They formed a
relatively secured or protected, apparently self-
sufficient entity, an organism itself.
Thus, was intiated another new process on the
surface of the earth. It was the organic evolution.
To begin with, inorganic and organic worlds
differed slightly between themselves, but the
differences were fundamental and qualitative; the
change was irreversible.
116 Part One: Principles
6.3.4 Key role of environment
The environment in which this evolution of the
qualitatively different object on the surface of the
earth took place was also not like the present. It
had some characteristics that helped entombment
or fossilization and subsequent preservation of the
organic materials of that time.
Oxygen acts on organic materials in two
contrasting ways before and after the death of
organisms. All living matter, i.e. organisms are
made of different chemical compounds. In general,
the most characteristic and ubiquitous of these
chemical constituents is the compound CH2O.
After the death of the organism, it is acted on by
the free oxygen in atmosphere to change into
carbon dioxide and water (CH2O + O2 — CO2
+ H2O). Thus, organic materials of dead bodies
are destroyed through oxidation. But during the
lifetime, oxygen interacts with other constituents
of body, viz. haemoglobin, etc. to oxidize food and
other nutrients to create energy. Energy, thus,
produced (heat, etc.) does not affect organism’s
life on account of the enzymes present in the body
that act as buffers. Instead of consuming the body
material through its reaction, oxygen, thus, acts as
one of the essential ingredients for life.
This explains life’s existence and sustenance
in an ambience of the atmosphere with free oxygen.
On the other hand, at the earliest stages of evolution
of life, however, the atmosphere did not contain
much free oxygen. Whatever little amount was
there, it was readily consumed by the huge amount
of volcanic gases, or by its reaction with the then
rocks. Contemporary organisms could not make
any use of that. Early organisms were chemo-
trophic, meaning they produced necessary
substances for their sustenance synthesizing the
materials acquired from the environment through
simple chemical reactions. At the same time, as
there was not much oxygen in the atmosphere, the
breakdown of CH2O of their body materials after
the organism’s death was also hindered. This added
to the preservation of potential of organic materials
of that time. If and when properly covered or buried
under sediments and not destroyed by subsequent
diagenesis or metamorphism, they were likely to
be preserved. This answers how fossils of bacteria,
etc. could be found in rocks as old as early
Precambrians.
6.3.5 Evidence of early life
Life appears to be as old as the oldest rocks.
Graphite in the banded iron formations of Isua,
Greenland, some of the oldest known sedimentary
rocks, attest to the presence and concentration of
carbon derived from the then organisms. Carbon
isotope ratios (13C to 12C) of this carbon are the
same as that of biological systems today.
Stromatolites serve as further and more direct
evidence (see Chapter 17 for more details on
stromatolites). The oldest stromatolite-like
structures come from the Pilbara Shield of
Australia. The rocks are 3.4 to 3.5 billion years
old. Definite stromatolites are known from
Bulawayan Group of Rhodesia (Zimbabwe),
possibly of 2.8 billion years age, and Pongola
Supergroup of 3 billion years age.
Two points about these Archaean stromatolites
are relevant. First, their rarity does not necessarily
mean the absence of the organisms that built them.
Archaean shelf deposits are rare and, thus, the rarity
of stromatolites may be due to the paucity of
conditions that could have allowed their
preservation.
Secondly, it is not yet decided for certain, if
the organisms were photosynthetic like the present-
day cyanophytes that build stromatolites or were
non-photosynthetic like certain thread-like bacteria
which are found to form stromatolitic structures
in the present-day hot springs as in Yellowstone
National Park.
Stromatolites are organosedimentary structures
that suggest the existence of organisms to produce
them. More direct evidence come from actual
fossils of cyanophytes (cyanobacteria) and other
bacteria found from the Warawoona group of West
 Chapter 6
Australia or Fig Tree Group of South Africa. The
filamentous, transversely partitioned fossils of 3.5
billion years old rocks of West Australia are similar
to modern cyanobacteria; whereas spheroidal
fossils of 3 billion years old Fig Tree rocks also
resemble cyanobacteria.
These earliest organisms were prokaryotic and
belonged to Monera. They were unicellular,
without any nucleus in the cell and their DNA was
not arranged in chromosomes. They are often
classified into Archaebacteria (that live in anoxic
conditions and include halophiles, living in brines,
methanogens or methane producing and
thermophiles, living in hot springs) and Eubacteria
(including cyanobacteria, loosely and wrongly
referred as blue-green algae and certain other
photosynthetic bacteria such as green or purple
bacteria). There are, however, some controversies
on who, chemotrophs or heterotrophs, preceded
the other.
6.3.6 Chemotrophs, autotrophs or
heterotrophs
Whether the earliest organisms were chemotrophs,
autotrophs or heterotrophs, is still a debated issue.
Modern cells use ATP or Adenosine triphosphate
as a source of energy; early cells also probably had
and used ATP. They acquired ATP from the
environment, which might have formed from
simple gases inorganically. They also obtained
amino acids, sugars and other compounds from
their environment. The process stood, in effect, like
eating; the organisms, thus, claiming the character
of heterotrophs, the animals.
There might have been also bacteria, like some
present-day ones, which had the ability of
fermenting organic compounds, that is, breaking
down complex compounds into simpler ones and
using the energy, thus, liberated in building some
other necessary compounds. Sulphate reducing
bacteria make one kind, which remove oxygen
from sulphates and release hydrogen sulphide,
Major Events of History of Life 117
obtaining energy therefrom. Methanogens release
methane through their metabolic activity. Such
bacteria are and were chemotrophs.
Which of the two, the early heterotrophs or
chemotrophs, came first is not yet definitely
known. However, the emergence of autotrophs
was definitely a later event. These were
fundamentally characterized by photosynthesis,
which might have been triggered under the
influence of frequent meteoric impacts of that
time. With the help of their chlorophylls, these
autotrophs could use solar energy, the sunlight,
and transform it to chemical energy. With its help
they could convert carbon dioxide and water into
energy-rich sugar and also use the energy released
from the breakdown of sugar in other chemical
reactions. But they had no use of oxygen produced
during these processes. Rather that could have
proved detrimental for the body, burning it up.
For early autotrophs to survive, reducing
atmosphere was, thus, more suitable.
6.3.7 Internal and external environment
fortified the organic world and
opened new vistas as well
The new material components of the earth, the
living things were, thus, in constant interaction
with their environment. They were sort of
experimenting different ways to improve their
basic or vital processes, metabolism and
reproduction. What they needed was improvement
and strengthening of their internal environment,
which identified them as organisms. Early
heterotrophs, chemotrophs or autotrophs were
different manifestations of such attempts, which
also meant different kinds of adaptations that
developed in these early organisms. All these
continue till to date; chemotrophs persist in typical
specialized anoxic conditions; autotrophs and
heterotrophs continue as plants and animals. But
right then, in those early days, the first evolving
autotrophs faced the problem; they were adopted
118 Part One: Principles
to reducing condition, yet they were producing
oxygen, risking burning up and oxygen could help
in many chemical reactions in turn. It required a
step to evolve some such means or material to
guard or prevent oxygen from acting foul on the
organism. The formation of enzymes proved to be
that step.
Enzymes might have been produced at the
next stage of evolutionary processes. Once they
formed, they created a more secured internal
environment for the organism. Photosynthesis
itself might have worked as the cause of
emergence of enzymes. At the same time, creation
of enzymes prevented burning up of organisms’
body by oxygen liberated during photosynthesis.
Autotrophs were then gradually adapted to an
oxygenating atmosphere from their adaptation to
reducing atmosphere. This made way for their
development. Besides, the more they spread in
number through successful reproduction, the
more and more free oxygen was added to the
atmosphere to make it oxygenating. At the same
time, with abundant organic material on the
surface of the earth, condition was created for the
appearance of unicellular heterotrophs, to
accompany unicellular autotrophs. The former,
the heterotrophs, could ingest organic materials
from nature as food; they were then acted upon
by oxygen and, thus, used for growth and nutrition
of the organism.
Though it cannot be definitely marked, it
appears that the formation of ozone layer in the
atmosphere might have taken place side by side
with the evolution of an atmosphere with free
oxygen. Solar radiations that reached the surface
of the earth would have helped create the ozone
layer. The layer itself would then have prevented
the atmosphere from radiations, thus increasing
its oxygen content. Thick iron deposits of
Precambrian point to this trend of oxygen
increasing in the atmosphere, where ferrous iron
was oxidized into its ferric state. The latter
precipitated on account of its lower solubility,
giving rise to those hematitic-magnetitic deposits.
6.4 Diversification Ensues and
Continues
6.4.1 Multicellularity and
biomineralization
The next significant events in the same course of
evolution would have been the origin of
multicellular heterotroph organisms and
biomineralization, i.e. development of hard parts
on organisms. Both added to the efficiency and
strengthening of the body, i.e. the internal
environment of organisms; multicellularity giving
effect to more coordination and biomineralization
making the body firmer and stronger with hard
parts. Many aspects of this part of organic
evolution, namely early evolution of multicellular
organisms are now becoming evident from recent
studies on fossil Lagerstätten such as the Edicara
or Burgess Shale biota (see Chapter 2 and
Appendix 1). Yet a lot remains unknown and to be
done. One such debated question is about the
mineralized hard parts of organisms.
At one stage, it was considered that the origin
of hard parts in course of organic evolution
enhanced preservation potentialities of organisms.
Thus, the sudden appearance of varied and
numerous organisms with hard parts in Cambrian
fossil record was explained as a mark of their
taphonomic advantage. As they had hard parts, they
were preserved more easily; their predecessors
without hard parts were simply not preserved and,
thus, lost without being recorded.
Broadly true, this idea is amended to some
extent in more recent views. Many palaeontologists
now consider that immediately before Cambrian,
body organization and physiology of organisms
reached quite a complex stage. An important aspect
of this complexity was the exclusion of unwanted
residues of food materials after necessary
ingredients had been acquired from them. In
biology, this process is the excretion. Such excreted
materials are essentially inorganic. To begin with,
they would accumulate on the body itself
 Chapter 6
eventually to harden and form an inorganic i.e.
mineralized hard cover on the body. The cover
added to the safety of the body, as well as acted as
a firm foothold or frame, a shelter, for the softer
fleshy parts. That brought considerable mechanical
advantage in moving different appendages or in
locomotion. Natural selection, thus, favoured
organisms with such harder accumulations on their
body, as they proved better adapted than their
counterparts without hard covers.
6.4.2 Diversification in water:
Vertebrates reign supreme
Origin of life and appearance of organisms took
place in water (may be on land, as some authors
think). Water itself protected the newborn organic
kingdom from different kinds of radiations
reaching the earth’s surface. Even after that, for a
long period, the organic kingdom evolved in water.
It embraced evolution from prockaryotes to
euckaryotic multicellular invertebrates, even
vertebrates, that is, the fishes.
In the earlier parts of this phase, viz. during
Upper Precambrian to Lower Palaeozoic
(Cambrian-Ordovician), heterotrophs appeared to
possess more developed body organization than
autotrophs did. That is, on questions of protecting
body-mass, locomotion, food gathering,
reproduction, etc. heterotrophs proved to have
moved steps ahead of autotrophs. This opened
newer and newer adaptive possibilities before
heterotrophs and during the said span of time,
particularly near the Cambrian-Ordovician
boundary, they went through radiation to find
successful adaptation to different niches of sessile
or vagrant benthic, as well as pelagic mode of living
in aqueous evironments.
Origin of fish-like animals in water in lower
Cambrian is a significant stage in organic
evolution. At this stage, organisms acquired not
just hard protective covering on body, but a well-
developed skeletal framework that provided good
anchorage for different organs of the body, to work
Major Events of History of Life 119
more smoothly and efficiently. It also provided the
entire body some streamlined, adequately firm
shape and structure that helped those organisms to
move freely and smoothly in water. In other words,
those organisms that developed the above
characters could most successfully utilize the
aqueous evironment present before the organisms
of that time. They became the early fish-like
organisms; they were vertebrates, thereby ushering
in the division between invertebrates and
vertebrates in the organic kingdom.
6.4.3 Adaptive radiation, extinction and
advent on land shaped the organic
world through Palaeozoic
Autotrophs and heterotrophs that had evolved
during this earlier part of Palaeozoic were affected
by two mass extinctions, one in late Ordovician
and the other in Upper Devonian. In spite of that,
the evolution of larger taxa (from the level of class
and up) proceeded in older lines even during
middle and upper parts of Palaeozoic (Silurian-
Devonian and Permo-Carboniferous, respectively).
Amidst this, fish-like organisms with jaws and
fins originating from their jawless counterparts
soon became the most successful group in water
of Middle Palaeozoic, assuming numerical
majority as well as extreme diversity.
At this juncture, two hitherto unexplored and,
hence, unutilized habitats opened up before the
organic world. They were land and freshwater
bodies i.e. the continents that had grown to
considerable size. Competition and selection
pressure in water were resolved through adaptation
in these two new environments. Changes that had
been taking place in the organic world paved way
for this adaptation. A few water-dwelling
organisms became inhabitants of freshwater
bodies. Some plants developed vascular tissues in
them and free spores appeared. As a result, there
also appeared continental vascular and non-
vascular plants. On the other hand, the advent of
tetrapods opened adaptation to different niches on
120 Part One: Principles
land independent of water. Hitherto uninhabited,
these niches provided environment without
competition and, thus, there developed rapid and
extensive adaptive radiation. Chapters 5, 18 and
19 give more details of these changes in vertebrates
and plants.
Among invertebrates, insects (Insecta of
phylum Arthropoda) on land with wings and air-
breathing respiratory system, and brachiopods
among benthics, nautiloids and goniatitic
ammonoids of nektic habit became characteristic
of the age. However, the event of far-reaching
consequence, and thus of importance, was the first
appearance of reptiles with amniotic eggs. Like
seeds, these eggs provided an environment within
the eggshell, which was not just well-protected, but
also a fluid with necessary nutrients. This meant,
unlike amphibian counterparts, reptilian eggs could
sustain themselves independent of water. Thus,
reproduction was possible in the drier ambience on
land. Obviously this added a great advantage to those
living on land, making reptiles more successfully
adapted to land than amphibians were.
Global transgression in Permian caused rapid
and varied adaptation of marine organisms, giving
rise to their numerous genera and species occurring
in abundance. But widespread end-Permian
extinction, most severe of the mass extinctions,
exterminated overwhelming majority of Palaeozoic
organisms, animals or plants, land-dwelling or
marine.
6.4.4 Mesozoic ushered in modernization
Triassic, the next period was marked by the
transition from older (Palaeozoic) fauna to the
modern types as well as by the typical
palaeogeographic (a single continent of Pangaea
surrounded by seas) and palaeoclimatic (relatively
arid climate over larger parts of the globe)
developments. Red sandstones and shales with
ferric iron cement developed in continental basins
are considered as signature of this climatic
condition. Organic world, too, faced important
changes. Some of them have been discussed with
reference to vertebrates and plants in their
respective chapters. A few are added here.
New groups of organisms appeared in marine
realm. For example, among benthics, with tabulates
and tetracorals being extinct by the end of Permian,
anthozoans were represented by the new group of
Scleractinia; ceratitic ammonoids replaced
goniatitic groups.
The most significant development perhaps
took place in land vertebrates. Developed pelvic
girdle rendered dinosaurs among reptiles, the
advantage and ability to move and live freely on
land; this along with other attributes established
their superiority on land. Apart from this, the
appearance of placental mammals marked a new
age. These were characterized by growth and
development of the embryo well-protected within
the mother’s womb, until it is adequately grown
and strong enough. This characteristic provided the
vulnerable early ontogenetic stages of individuals
to pass through in a very secured environment; it
was independent of water, where life had begun,
even of the immediate ambience prevailing on land.
With this, their warm-blooded body, improved
quadrupedal posture for still easier locomotion so
far attained by animals — all this made these
animals much better adapted on land. Thus, in
Jurassic, side by side with dinosaurs, mammals
developed on one hand. On the other, there
appeared birds with feathers on their body, giving
it insulation from variation in temperature around.
It was another step for land-dwelling animals
towards overcoming adversaries of nature, i.e.
towards achieving a better adaptation.
There were a few more significant events in
the organic world during the last phases of
Mesozoic, in Jurassic- Cretaceous. After the break-
up of Pangaea had started in Triassic, global
palaeogeography, rather land-sea distribution was
heading towards a shape that ultimately graded into
its present pattern. With that, there were being
established sedimentological regimes on ocean
floors that could be called modern in kind.
There were, thus, some major changes in benthic
 Chapter 6
life in oceans. Among invertebrates there, bivalves
with siphons and endobenthic echinoids with
respiratory tubefeet and petaloid ambulacra
appeared, and finally successfully adapted to and
flourished with endobenthic, particularly
burrowing, mode of living in the new sediment-
laden substrate in place of earlier epibenthics that
were suited to harder rocky substrates (see Chapters
3, 10 and 13). Among the epibenthics, gastropods
with a well-developed canal system marked a new
progress, by which system they not only overcame
their problems in water intake and flush-out (see
Chapter 11 for details), they explored and finally
succeeded in a new, carnivorous predatory method
of feeding. In the nektic mode, ammonites passed
through rapid evolution with varied successful
adaptation. Over and above these developments in
macro-invertebrates, the appearance of calcareous
shell bearing micro-organisms, both benthic and
more so planktic foraminifera, led finally to their
rapid proliferation in Cenozoic. Global
transgressions of middle to late Cretaceous might
have contributed to these changes in marine
invertebrate life. On land, the appearance and
rapid spread of flowering plants, the angiosperms,
expanded the food chain considerably; it led to
the origin and spread of different types of
herbivorous mammals, grazers and browsers, as
well as carnivores feeding on the latter. On the
whole, the organic world started to assume its
present shape. But the end-Cretaceous mass
extinction, which may really have taken place in
a series or steps, again wiped out a large portion
of the then organic world, both on land and in
water. On the contrary, some of the groups which
had appeared earlier, found innumerable
possibilities for adaptation wide open. In
flowering plants, fertilization was further secured
in the protected ambience within flowers which
helped these organisms overcome natural
adversaries to ensure successful reproduction and
thus abundance. Newly developed varied insect
populations took important part in the pollination
process of plants.
Major Events of History of Life 121
Such intimate relationship between plants and
animals were evident in several other cases. The
spread of flowering ground plants, i.e. grasses in
Oligo-Miocene times of Tertiary period, led to the
spread of a new environment of grasslands by the
side of the then forests. This triggered evolution
of browsers (soft plant or leaf-eating organisms)
to grazers with grass-eating habit, and feeding on
them, the emergence and evolution of newer and
newer types of carnivores. Also in the mixed or
chequered environment of grassland and forest,
there took place another significant event of the
organic evolution which led ultimately to the
appearance and evolution of hominids, rather man
(see Chapter 5). With this, for the first time in
organic evolution there appeared an organism,
Homo sapiens sapiens, which not only reached the
highly coordinated body governed by the well-
developed brain, but also it possessed such power
as to exert control over its own internal as well as
external environments.
In plants and invertebrates too, the changes
that took place in the post-Cretaceous time marked
the advent of organisms of modern affinity. Thus,
Tertiary saw the dominance of cryptogams, and
abundance of modern representatives of bivalves,
gastropods, echinoids, scleractinian corals, fishes
and such others.
6.5 Major Mass Extinctions:
Important Component in the
History of Life
Above, we have had a brief summary of how,
step by step, life appeared, diversified and spread
over the earth to assume the present form of the
organic kingdom. This appraisal is never complete
without giving a thought to extinction. Importance
of extinction in evolution have already been
pointed out in section 5.2.4. It had been mentioned
there that extinction has been a parallel event in
the evolution all through, with a few periods or
episodes of mass extinctions present in the
122 Part One: Principles
geological column that were marked by large-scale
or massive extinction. Here we list those major
mass extinctions in the history of life and the
probable causes triggering them.
The earliest of these principal extinctions is
marked at late Precambrian. It terminated the
well-known Ediacaran fauna, which is termed
Vendozoan fauna by Seilacher (1989), distinctly
different from later metazoans. The extinction was
followed by the more recently discovered SSF or
Small Shelly Fossils (Clarkson 1998). It is,
however, considered as produced by a complex set
of factors (Donovan 1989) including widespread
regression, physical stress (restricted circulation
and oxygen deficiency) and biological stress
(increased predation, scavenging and bioturbaton).
The next episode took place in late Cambrian
due to habitat reduction, may be in response to a
rise in sea level. It is best recorded by trilobites and
might have been geologically rapid. The end
Ordovician (end-Ashgillian) extinction was one of
the five more important episodes (the other four
being end-Devonian, end-Permian, late Triassic and
end-Cretaceous) and wiped out nearly 22 per cent
of all families, including those of early vertebrates.
The next major mass extinction took place in
the late Devonian (Frasnian–Famennian
boundary) which destroyed about 21 per cent of
families, vertebrates or invertebrates. The most
severe was the late Permian crisis that wiped out
marine invertebrate families by 57 per cent (~ 95
per cent species disappeared). In the late Triassic
(Norian) 20 per cent and in the late Cretaceous
(Maestrichtian) 15 per cent of families were
extinct, respectively. In the last of these, both land
and marine faunas became extinct.
In Tertiary, a stepwise extinction at Eocene-
Oligocene boundary was associated with the
terminal Eocene cooling event along with changes
in oceanographic circulation (see Chapter 20 for
more details). The last and the more recent
extinction (late Pleistocene) is associated with the
latest phase of the cooling that took place in
Cenozoic, i.e. the Pleistocene glaciation. The
extinction phase followed the glaciation and might
have been accentuated by predation by man.
 Part two
Major Invertebrate
Groups
 7
Phylum Cnidaria
7.1 Introduction
The phylum which was earlier called Coelenterata
has presently been classified into two phyla
(Clarkson 1998). The one called Ctenophora is
represented by extant, i.e. recent animals with its
only fossil representative described from Lower
Devonian of Germany. The other is the phylum
Cnidaria, which may be regarded as the simplest
representative of metazoan or multicelled animals.
Phylum Porifera includes animals, the sponges,
simpler than the cnidarian ones. They are
multicelled and have different kinds of cells, but
those are not organized into tissues. Besides they
do not have any nervous system or nerve cells. So,
they are now being increasingly considered as an
intermediate stage, termed multicelled Parazoa,
between one-celled Protozoa and multicelled
Metazoa (see Factsheet 7.1).
Cnidaria has cells organized into tissues. Body
wall is diploblastic with an outer ectoderm and
an inner endoderm, as well as an intermediate
jelly-like mesoglea. The latter contains nerve mesh,
albeit in a very simple form. This makes cnidarians
more complex in body organization than sponges.
On the other hand, the rest of the metazoans have
triploblastic body wall, certainly one step ahead in
organization.
125
Cnidarian body has a body cavity or enteron
which opens only at the distal end serving as the
mouth. The mouth is surrounded by prehensile
tentacles used for food gathering. The enteron has
a few fleshy partitions called mesenteries generally
arranged radially. Endoderm of the body wall and
mesenteries help in digestive processes; ectoderm
with the help of its hair-like appendages called
pseudopodia help in food catching.
7.2 Cnidaria in Palaeontology:
Importance of Polyp and
Medusa
Cnidarians are aquatic and dominantly marine.
Their fossil record extends from Precambrian
Ediacara biota (see Factsheet Apx.1.1). Present-
day corals (class Anthozoa), hydra (class
Hydrozoa) and jellyfish (class Scyphozoa) are
examples of the phylum. But these groups do not
carry equal importance in palaeontology. The
reason lies in the life cycle of cnidarian animals.
Cnidarian life cycle is characterized by alternation
of generations of different kinds. Two different
types of individuals, polyp and medusa alternate;
polyp reproduces medusa asexually and medusa
reproduces polyp sexually, and so on. A polyp is
126 Part Two: Major Invertebrate Groups

FACTSHEET 7.1
Cnidaria Phylum: Where it Stands in the Organic World

Phylum Earlier Broader Principal Geological Example

known as identity characteristics age

Few phyla Protozoa Protozoa/ Unicellular, Precambrian Foraminifera,

of animal affinity Protoctista cell nucleate, -Recent Radiolaria, etc.
in Kingdom heterotrophic
Protoctista
Porifera Porifera Parazoa Multicellular, Cambrian Sponges
nucleate,cells -Recent
not organized
in tissues, no
nervous system

Cnidaria Coelenterata Metazoa Multicellular, Precambrian Corals, hydra,

diploblastic body- -Recent jellyfish, etc.
wall, nerve-net
in mesogleaa
Ctenophora Coelenterata Metazoa Multicellular, Recent, the Sea-gooseberry,
possibly only fossil comb-jelly
triploblasticb in Lower
Devonian
Other phyla Metazoa Multicellular, Other
triploblastic animals
(a) Some additional characteristics: Body cavity or enteron has a single opening serving both as mouth and anus; no
excretory and circulatory systems; marine, nektic or benthic.
(b) globular body; nektic in habit.

sedentary, apically fixed and ending in a mouth
surrounded by tentacles at the distal end of a
cylindrical body; mesoglea is thin. Medusae are
free-swimming, discoidal in shape with the mouth
surrounded by tentacles, located downwards at the
end of a funnel-shaped projection of the body.
Mesoglea is thick in medusae. These
morphologically different individuals appearing at
different times, thus, serve as examples of
temporal polymorphs.
Cnidaria is subdivided into three classes, viz.
Hydrozoa, Scyphozoa and Anthozoa. All these
three classes range from Precambrian to Recent.
Of them, the primitive or less specialized
cnidarians, the hydrozoans, show this feature of
polymorphism. In more advanced groups one of
the either may be suppressed entirely. Thus, in
Scyphozoa polypoid phase is very reduced and the
class is more characterized by its medusoid phase.
On the other hand, in Anthozoa, medusoid phase
is eliminated entirely. It has a direct bearing in fossil
record, as only polyps have hard skeletal parts and
medusae lack them. This makes scyphozoan fossils
rare in the geological column. However, certain
exceptionally well-preserved records, such as the
Precambrian Ediacara biota presents medusoid
 Chapter 7 Phylum Cnidaria 127

scyphozoans, casting much light on the animal and
its biology. Anthozoans are, however, very
common in certain parts of the geological past and
reveal different important palaeontological aspects
of ancient life. The following discussion will
concentrate on Anthozoa with brief comments on
two other major classes of Cnidaria.
Hydrozoans are polymorphic, with radial,
tetrameral symmetry. They may be solitary or
colonial (see section 7.3.2). They are classified into
six orders of which one is extinct (Spongio-
morphida; early Mesozoic) and another only extant
(Siphonophora). Of the rest four with both fossil
and recent records, Hydroida and Hydrocorallina
are more important.
Hydroids may secrete chitinous external
skeleton and may be solitary or colonial. They are
well-known in Ordovician. Some species of recent
genus Proboscidactyla are symbiotic with
tubular worms (sabellid type). Mesozoic-Tertiary
occurrences of a hydroid-serpulid worm
association suggest a similar symbiotic or
commensal relationship existing in the past.
Hydrocorallines (e.g. Millepora) resemble
corals in appearance, secrete calcareous skeleton,
are colonial in habit and often important
constituents in some recent reefs, thus being
ecologically significant.
Scyphozoans are dominantly free swimming
medusoid with a brief sessile polypoid stage that gives
rise to a number of small medusae. They do not have
skeletons, though they, including jellyfish, have left
fossil record since Precambrian. Fossils occur either
as compressions in fine sediments as sand infillings
of the gut cavity, or as simple resting marks.
Anthozoans are totally polypoid, strictly sessile
and marine, calcareous skelton-secreting
cnidarians. In some groups of anthozoans, the
mineral is calcite, while in one major and extant
group scleractinia, the skeleton is made of
aragonite. Skeletal morphology of anthozoans,
particularly relevant for fossils, is discussed in the
following sections.
7.3 A Framework for Morphology
There are controversies on subdivisions of the
class (Factsheet 7.2). Generally, speaking three
sub-divisions, viz. Tabulata, Rugosa (or Tetra-
corallia) and Scleractinia (Hexacorallia of some
authors) are the major ones among them,
particularly as fossils. The following discussion
embraces all the three, though as and when
required, some separate treatment for individual
groups is also made.

FACTSHEET 7.2
Varied Opinions on Systematics of Major Groups of Anthozoa
Earlier opinion Major group Present opinion

Subclass = TABULATA = Order (Subclass Zoantharia)

? Cambrian-Permian
Subclass = RUGOSA/ TETRACORALLIA = Order (Subclass Zoantharia)
? Cambrian/Ordovician-? Permian/Lr. Triassic
Order = SCLERACTINIA = Order (Subclass Zoantharia)
(Subclass Zoantharia) Mid. Triassic-Recent
Order = HETEROCORALLIA = Order (Subclass Zoantharia)
(Subclass Zoantharia) Up. Devonian-Lr. Carboniferous
(Subclass Octocorallia) OCTOCORALLIA (Subclass Octocorallia)
? Precambrian/Ordovician-Recent
CERIANTIPATHARIA (Subclass Ceriantipatharia)
Recent (doubtful fossil record)
128 Part Two: Major Invertebrate Groups
Each individual anthozoan animal represents
a polyp. It may live singly or along with a few
others being connected with one or more of them
during the lifetime. The former is termed solitary,
while the latter is the colonial mode of living.
(Factsheet 7.3). Respective hard parts are also
referred as solitary, single or simple and colonial
or compound. They are called corallite or
corallum, their usage being strewn again with
some controversies (Factsheet 7.4).
Morphology of corallites can be studied under
the following heads:
1. Shape of corallites and its variation;
arrangement of corallites in a colony.
2. Corallite wall and its varieties.
FACTSHEET 7.3
Living in a Group
Men live in society, a group of individuals with
developed intercommunication and a set of
bindings and rules built thereupon, leading them to
play upon their environments even.
Elephants, for example, live in herds, each
individual a separate entity but communicating with
others, yet subjugation to environment is hardly
overcome.
Echinoids are gregarious, where they live in large
numbers at a place with each individual main-
taining its separate entity and existence.
Some corals live in colonies, each colony being an
organic whole with individuals living together in an
interconnected and interdependent manner.
FACTSHEET 7.4
Corallum-Corallite: Definition and Debate
Corallum/corallite refer to skeleton of
Earlier usage Recent usage
Corallite
A single coral Individuals of a colony
Corallum
Colonial corals Both single and colonial
3. Major features within corallite: septa, tabulae,
dissepiments, axial structure.
4. Other features.
7.3.1 Shape of corallites
The shape of corallites is controlled, firstly by
whether the animal is single or colonial. Each
solitary corallite or each individual corallite of a
colony begins as a low conical calcareous cup
covering the animal at its apical side. It is called
basal disc. As it grows, a number of partitions
called septa develop. The polyp, the basal disc and
its septa, all grow essentially vertically upwards,
as the animal remains attached to the substrate of
the basin. Towards that distal end, each septum
has a margin curved downwards towards the axis
of the corallite. Such upper edges of septa, thus,
create a bowl-shaped depression at the distal end
of the corallite, that is known as calice or calyx.
The main bulk of the polyp sits on it. With further
growth of polyp, more mineral matter is added
along the distal margin of the basal disc around
the animal and the corallite grows accordingly.
Simultaneously, septa and other structures grow
on inside the hollow of the corallite defined by the
added mineral matter, which forms a wall for the
animal and the corallite. The ultimate shape of the
corallite will depend on whether the rate of growth
is greater upwards or sideways. In the former case,
the corallite will assume a tubular or cylindrical
shape and in the latter, the initial conical shape
will be largely retained. There may be a number of
varieties of shape depending on how far vertical
rate exceeds over horizontal rate or vice versa
(Figure 7.1; Factsheet 7.5).
7.3.2 Colony and shape, and
arrangement of corallites
In colonial forms, the shape of corallites depend
on the nature of their arrangement within the
colony. A colony is an organic whole in which a
number of polyps live together in an interconnected
and interdependent manner, each of them, barring
 Chapter 7 Phylum Cnidaria 129

(b)

(a)

(c)

(d)

(f)
(e)
(g)

(h)

(k) (l)
(i) (j)

(m)

(i) (iii)
(ii) (iv)
Fig. 7.1 Cnidaria.
(a) A coral polyp; (b) A medusa (both are sections in life position); (c) to (l) Different shapes of
solitary corallites; (c) Button-shaped; (d) Patellate; (e) Long trochoid; (f) Curved ceratoid; (g) Curved
cylindrical; (h) Ceratoid with talons; (i) Scolecoid; (j) Pyramidal; (k) Turbinate;
(l) Calceoloid; (m) Variation in shape with variation in vertical and horizontal growth rates—(i) horizontal
rate > vertical, (ii) two nearly equal, (iii) and (iv) vertical > horizontal.
130 Part Two: Major Invertebrate Groups

FACTSHEET 7.5 7.3.3 Economy of space and closer

Variation in Corallite Shape
Corallite tubular Corallite conical
(v > h) (h > v)
Axis Axis Apical XSn(circle/ellipse
straight zigzag angle semicircle/polygon)
Tubular Scolecoid > 90° Patellate
Tubular > 40° Trochoid
Calceoloid/turbinate
Prismatic » 30° Ceratoid pyramidal
h: horizontal growth; v: vertical growth;
XSn: cross-section
the first one, having formed from one or the other
member of the colony itself. Members may even
be specialized in functions they perform, some
controlling food-gathering, some others
reproduction, and so on. Whether the animal will
turn out to be single or colonial, is determined by
its mode of reproduction. Anthozoans show two
major modes of reproduction: asexual or budding,
and sexual or fission. The genera characterized by
the first kind form colonies. It should be added
here that colonies are not to be confused with reefs.
Coral reefs are embankments within shallow,
warm, tropical seas in which colonial corals may
be dominant, with or without the presence of a
number of single forms.
Among anthozoans, tabulates of Palaeozoic
age are strictly colonial; corals of another extinct
Palaeozoic group Rugosa (presently many authors
consider both to have ranged into Triassic), as also
of the extant major group Scleractinia include both
solitary and colonial forms. Obviously, it is difficult
to bring out the relationship between reproductive
processes and formation, and growth of colonies
in the case of tabulate and rugose corals.
Scleractinians, on the other hand, develop well-
organized skeletons with fair amount of secretion
of calcium carbonate. They may, thus, provide
more light on the formation and growth of colonies.
interconnection control
arrangement in colonies
Arrangement and organization of polyps and their
corallites in a colony is closely linked on how
efficiently the members are interconnected and
interdependent to live and grow successfully
together. For the first place, for a particular
number of polyps, the less space they require to
live together, the easier will be for them to be
interconnected and interdependent. So,
arrangement of corallites in a corallum of a colony,
depends on two aspects: first, developing closer
interconnections and second, using space
economically. Basically, each corallite of a colony
is either conical or cylindrical with a circular or
elliptical cross-section, same as in solitary corals.
When more and more individuals are produced
from a polyp with such a corallite, they either
remain parallel or branch out. Even if they are
closely compact and are longitudinally in contact
with adjacent ones, there remains some amount of
space unutilized in a colony, leaving gaps in
interconnection. From such a loose organization,
colonies become more compact with more
utilization of space and closer interconnection. It
demands that corallites become prismatic
(or pyramidal) instead of being tubular (or conical),
with cross-section changing from circular/elliptical
to polygonal. But walls existing between two
corallites of even polygonal section stand in the
way of interconnection, especially of the softer
parts of the polyps. Pores in the wall help
connecting adjacent corallites and their polyps; the
development of a single common wall between two
adjacent corallites is a further step towards
removing obstruction to interconnection. The next
step is the removal of wall altogether and then the
merger or confluence of soft and hard parts of
adjacent corals. Figure 7.2 depicts the series and
resultant types.
Variations in the shape of corallites and
arrangement of corallites in a colony are in all
 Chapter 7 Phylum Cnidaria 131

(a)

(i) (ii) (iii) (iv)

(b)

(i) (ii)
(iii) (iv)

(c)

(i) (ii) (iii) (iv)

(d)

Fig. 7.2 Arrangements of corallites in colonies: a series of utilizing space and increasing interconnection.
(a) Corallites tubular, separate, connected by crosstubes (i and ii) or separated by coenenchynal tissues
(iii and iv),
(b) Corallites tubular, in contact in a chain (i and ii) or in a loose bundle (iii and iv),
(c) Corallites prismatic, sharing walls with mural pores (i and ii) or without (iii and iv), (d) Corallites without
walls,
(a), (b), (c), (d) are cross-sections, (i and iii) longitudinal section or view, and (ii and iv) cross-section.

likelihood genetic, a genus or species being These variations appear to be related to the kind
characterized by some shape or some of substrate on which the colony is growing,
arrangement. However, colonies also may be intensity of current, content of suspended fine
massive, encrusting, creeping or foliaceous, etc. sediments, etc.
132 Part Two: Major Invertebrate Groups
7.3.4 Wall
Wall may or may not be present, both in solitary
(e.g. Montlivaltia) or in colonial (e.g. Isastrea)
corals. The wall is generally imperforate, hard
mineralized and grown in continuity with the basal
disc. However, each growth stage is marked by
breaks at the beginning and at the end, represented
by a groove in each case. The surface of the tubular,
conical or polygonal corallites thus contains
parallel growth lines or rings; the thicker the wall
is, the more prominent are the growth lines.
Commonly Rugosa (meaning rough) have thick
walls; tabulates and scleractinians have thinner
walls or none at all, in the latter.
Such sheet-like wall is called epitheca. There
may be other kinds too, found in scleractinian
corals only. These may be septotheca in which
the outer edges of septa are thickened and fuse
with those of adjacent ones; paratheca where septa
do not reach the margin and the space left there is
occupied by dense dissepiments that form the wall
and synapticulotheca in which adjacent
synapticulae fuse with each other to produce the
wall (septa, dissepiments and synapticulae are
discussed later) (Factsheet 7.6).
7.3.5 Internal features
The hollow of the interior of corallite contains a
number of features. Major ones among them are
FACTSHEET 7.6
Variation in Corallite Wall
Wall type Formed
Epitheca In continuation of basal disc;
generally imperforate; in genera
of all major groups
Septotheca By outer thick edges of septa;
found only in scleractinia
Paratheca Of dissepiments, spongy; found
only in scleractinia
Synapti- By joining of synapticulae;
culotheca generally perforate; found only in
scleractinia
classified on their alignment and shape. These
include: (i) Septa which are sheet-like partitions,
longitudinal and vertical, i.e. running along the
length of the corallite from apex to mouth/calyx
and arranged radially or bilaterally, the symmetry
being evident in cross-sections or calyx; these may
extend from the wall to the axis or may stop short
of that length at any end; (ii) Tabulae are also
sheet-like partitions, but transverse to the axis and,
thus, horizontal; they are mutually broadly parallel
and may extend from one wall to another
diametrically opposite or may be shorter; (iii)
Dissepiments are small, inclined, fish scale-like
vescicular structures generally confined near the
margin and with their convexity towards the axis,
and (iv) Axial structure, which is a solid or spongy
linear feature running along the axis of the corallite.
Different genera or species of anthozoans have
different combinations of these features,
characteristic of the genera or species themselves.
7.3.6 Septa
Septa are by far the most important of the four types
mentioned above and, that too, at different
taxonomic levels, serving as their characteristics. A
species has some characteristic septa; whereas the
order Tabulata lacks septa or has only rudimentary
spinose ones. Septa in Rugosa are largely bilaterally
symmetrical, whereas in Scleractinia the symmetry
is radial or biradial (Figure 7.3).
Body cavity or enteron of anthozoans have
mesenteries in pairs. Each mesentery has a kind of
muscle on one side of it. In each pair, the muscles
are attached on the sides facing each other. But
there are two diametrically opposite pairs in which
that is not the case; the muscles occur not on facing,
rather on opposing sides. These are called directive
mesenteries. In any radially symmetrical septal
arrangement, the two pairs of directive mesenteries
together define a bilateral plane of symmetry
passing through them. This signifes a fundamental
or primary bilateral or at least a biradial symmetry
in anthozoans, whatever be its final expression,
radial or bilateral.
 Chapter 7 Phylum Cnidaria 133

(a) (b)
(v)

(i) (ii) (iii) (iv)

(c)

(d) (e) (f)

(g) (h)

(i)
sp

sp s s (k)
(j) (l)

Fig. 7.3 Features of anthozoans.

(a) Six protosepta with radial symmetry and mesenteries, (b) Introduction of later septa, major and
minor in radially symmetrical form, (c) Serial sections to show introduction of septa in bilaterallly
symmetrical forms, (i) Median septum; (ii) Median septum broken into cardinal and counter with two
alar septa; (iii) Two counter-lateral septa added; (iv) and (v) minor septa added in four sectors,
(d) Schematic view for septa and axial structure (columella), (e) Schematic view for tabulae,
(f) Schematic view for dissepiments, (g) Cross-section of a corallite to show septa, dissepiments and
columella, (h) Cross-section of a corallite to show strong dissepiments, weak short septa and axial
structure, (i) Schematic section to show origin of palus (p), (j) Synapticulae (s) between traces of septa
(sp), (k) Mural pores in a tabulate coral, (l) A turbinate coral with well-developed septa.
134 Part Two: Major Invertebrate Groups

The space between the two mesenteries of each Septa vary in other aspects too (Factsheet 7.7).
pair is called entocoel and that between two Of them, variation in structural organization of
adjacent pairs exocoel. Septa form between each septum is more important. Crystals of calcium
mesenteries. In early ontogeny they form within carbonate that make the corallite, be they aragonite
entocoels, but later and in most cases within or calcite, make three kinds of structural units, viz.
exocoels. Accordingly, they are termed entosepta trabecular, fibro-normal and lamellar. A septum
or exosepta. On the basis of the stage of is made up of any combination of these units.
introduction of septa within a corallite, they are However, lamellar units are not found in
variously classified. Thus, prosepta scleractinians. Morphology of septa depends on
(or protosepta) are the first formed septa, six in their constituent units and how they are arranged.
number in Rugosa and Scleractinia; metasepta are Thus, if trabecular units are loosely bound in a
all septa other than prosepta; primary are those septum sheet, the latter becomes perforate; if some
introduced in the first cycle of septa; secondary of those unit project are out of the septal plane,
in the next cycle and tertiary, and so on. All they make spines or carina.
prosepta and a few earlier metasepta are major In spite of the fair amount of variation in
septa which are longer, extending from the wall morphology, external including shape, or internal,
to the axis; rest, the majority, are minor septa, similar adaptation has produced similar type of
shorter in length (in cross-section of the corallite). morphology in many unrelated corals and their
lineages. Such ‘convergence’ in evolution of corals
FACTSHEET 7.7 makes it necessary that some finer structure, not
dependent on ecology and environment, should be
Variation in Septa given weightage in taxonomy. Septal micro-
On when introduced in ontogeny structure, including the three units referred above,
stand a potential candidate for that purpose.
On cycle of On time of On importance However, they are not always thoroughly known
introduction introduction in morphology and understood as yet.
Primary; Prosepta Major In most scleractinia, corallites including their
Secondary; (introduced first) (long in TS) septa are formed of aragonite, whereas in rugosa
Tertiary; Metasepta Minor they are made of calcite. As aragonite is metastable
etc. (introduced later) (short in TS) and is easily changed into calcite, many authors
On structural characteristics believe calcite in rugosa of Palaeozoic age, may
really be diagenetic. But others consider it as
On characters of On morphological
septal laminae in LS variation in TS
primary and unchanged. The debate is yet
unresolved.
Solid/imperforate Straight/sinuous
Perforate Long/short/ 7.3.7 Other structures
Acanthine/spinose Lonsdaleoid
Amplexoid (detached from wall) Structures or features other than septa, viz. tabulae,
(attached to wall, Carinate/spinose dissepiments, epitheca, etc. are formed generally
shortened near axis) Perforate of fibro-normal units in rugosa and trabecular units
Dilated in scleractinia.
(thinner at axial end) Of these, tabulae are horizontal partitions that
Rhopaloid
occur as the major, even the only structure in
(thicker at axial end)
Tabulata; sometimes they are important in rugose
LS: longitudinal section; TS: transverse section genera. Morphologically they may be flat, concave

or convex upwards. Also, viewed in longitudinal
sections, they may extend from one wall to
diametrically opposite wall (called complete) or
from wall to the axis or near it (incomplete)
(Factsheet 7.8).
FACTSHEET 7.8
Variation in Tabulae
Extension Curvature Example
in TS (orally)
Complete: Concave Syringopora
wall to wall Flat Halysites
Incomplete:
wall to axis Convex Favosites
Dissepiments may be elongate or rounded
vescicular; in some genera they make the wall,
being densely packed along the margin. Tabulates
do not have them; rugose and scleractinian genera
and species may contain dissepiment to different
extents, sometimes characteristically. In
Cystiphyllum, a rugose coral, they are the only
structure.
Axial structure is also absent in tabulates and
may be present in some genera of Rugosa and
Scleractinia. Its presence and morphology may be
characteristic for a genus or species. It may be solid
or spongy and may have formed in a number of
ways. Among them, axial ends of septa may fuse
with each other to form axial vortex; small tabulae
may be packed one above the other along the axial
region to give rise to axial column; the axial end
of a septum may be thickened to form a solid or
hollow columella.
In addition to the above four kinds of major
structures within a corallite, there may be a few
others to be found in this or that group. Some of
them are described briefly in the following
paragraphs.
Fossula (pl. -lae) is a depression on the calyx
of the corallite which forms due to stoppage of
growth of a prosepta, with adjacent other septa
growing normally beyond it. It is designated by
Chapter 7 Phylum Cnidaria 135
the name of the prosepta, whose position it
occupies; thus cardinal fossula indicates that the
cardinal septum is suppressed, alar fossula for alar
septum, and so on. Fossula may be characteristic
in some genera, for example, cardinal fossula in
Zaphrentis.
Pali (pl. palus) are structures found at the axial
region of a small group of scleractinians. Normally
septa grow from wall inwards towards the axis.
Pali form when the axial parts of some entosepta
are detached from their marginal parts.
Synapticulae (sing.-la) are small needle-like
connections between two adjacent septa; they
represent bundles of trabeculae that do not lie on
the septal plane or lamina. They are found in
scleractinians.
In some rugose genera there is a lid or
operculum fitting to the calical outline. It shows
growth rings on outer surface and traces of septa
on the inner side. Calceola provides an example.
Rugose genus like Omphyma shows on its surface
roots or talon for fixing the corallite to the
substrate.
7.3.8 Septa and classification of
Anthozoa
Septal varieties have been discussed above. In
addition, variation of septa in rugose and
scleractinia help in their characterization. The two
groups differ considerably in the character of
prosepta, their sequence of introduction and
formation and arrangement of metasepta. On these
basis, the two groups were differentiated from each
other. More recent view, however, attaches lesser
importance to this aspect, though that does not
reduce the significance of septa.
As a corallite preserves all the stages of
ontogeny from the basal disc to the stage the animal
met death and was fossilized, serial transverse
sections provide how the different structures grew
within the corallite at different ontogenetic stages.
Such sections reveal that in rugose corals there
develops one single septum between the two pairs
136 Part Two: Major Invertebrate Groups

(a) (b)

(c)

(d) (e)

Fig. 7.4 Corals.

(a) Button shaped coral with inverted calyx, (b) A thamnasterioid colony in thin section,
(c) The same sample in hand specimen, (d) Scale for both (in mm), (e) A slice of a meandroid colony
(6 to 7 on the scale equals to one inch).

of directive mesenteries. It is called the median
septum and running along a diameter it defines a
bilateral or biradial symmetry. Subsequently, it
stops growing in its axial part and is, thus, divided
into two septa, a cardinal and a counter,
diametrically opposite to each other. At the next
stage, a pair of alar and another of counter-lateral
start growing on the two sides of cardinal and
counter septa, respectively, maintaining a bilateral
symmetry. At this stage, thus, six prosepta are
grouped into four types: cardinal, counter, alar and
counter-laterals. Metasepta or minor septa that start
developing here onwards are introduced at four
points, one in each of the four among the six
regions defined by the six prosepta. The areas
between the counter and counter-laterals do not
contain any later septa. The bilateral symmetry is
maintained until there are numerous septa within
the corallite to change the symmetry to radial. In
summary then, rugose have (a) six prosepta divided
into four types, (b) later septa introduced at four
points, and (c) initial bilateral symmetry is

disturbed to become radial as septa grow in
number. In fact, Tetracorallia, the other name of
Rugosa, is derived from this tetrameral
characteristics of the group.
Scleractinia, too, has six prosepta. But they
are introduced all at one time and being similar in
morphology, give rise to a hexameral radial
symmetry. Later septa are introduced at points,
multiple of 6, viz. 6, 12, etc. in consecutive cycles
and without disturbing the radial symmetry.
Scleractinia was, otherwise, called Hexacorallia
on this hexameral characteristics, though presently
the name is used for a different small group.
In Tabulata, septa are either absent or, when
present, occur in multiples of six as rudimentary,
spine or ridge-like traces on the inner surface of
the wall.
The above differences in septal characters
were considered to indicate fundamental difference
among the groups. They were accordingly ascribed
different systematic positions. Tabulates and
tetracorals were extinct by the end of Palaeozoic,
whereas Scleractinia appeared in Middle Triassic.
Phylogeny envisaged a radiation in Palaeozoic
which produced the two major groups of corals,
given the subclass status, viz. Tabulata and
Tetracorallia (or Rugosa). After they had been
extinct at the end of Paleozoic, Scleractinia
appeared and finally occupied the niche vacated.
However, towards the end of the seventies of
the last century, a few European palaeontologists
presented a different view (see Babin 1980). They
showed:
1. In both Rugosa and Scleractinia, the symmetry
reflected by the earliest septa is biradial or
bilateral. Even in the latter group, the median
septum comes first, as it does in Rugosa.
2. Even other four prosepta in Scleractinia are not
introduced simultaneously; they appear in two
pairs successively, similarly as they do in
Rugosa.
3. A scleractinian family, Caryophyllidae
maintains the bilateral symmetry.
Chapter 7 Phylum Cnidaria 137
4. In both the groups, later septa make a pinnate
arrangement.
5. Even in Scleractinia, septal introduction does
not take place in cyclical order; those apparently
belonging to a cycle appear one after another.
These observations tend to rule out any
fundamental difference between the two groups.
Some authors, thus, prefer to consider Scleractinia
as a more advanced descendant of Tetracorallia
without assigning it any majorly different
systematic status.
Another set of facts may be relevant on this
issue. Living scleractinians present a wide variety
of morphology and different kinds of arrangement
in colonies. It makes one of the most successful
animal group in the warm, shallow seas. About
65 per cent of living corals are colonial and barring
the tabulates, about 80 per cent of Palaeozoic corals
are solitary. Evolution from solitary to colonial
mode, thus seems to have formed a distinct trend
in anthozoans; in fact, Palaeogene occurrences of
the Paris basin bears testimony to this conclusion.
Colonial corals may then be more successful
members of the groups, in their joint, economic
use of nutrients and other resources of their marine
abode. It is suggested that in course of ruguse corals
getting adapted more and more to colonial mode,
there evolved scleractinians with changes also in
other characters like septa. Selection pressure of
the shallow marine niches led to the appearance
and evolution of scleractinians.
7.4 Geological Importance of
Anthozoa
This class of Cnidaria is considerably important
in biostratigraphy, palaeoecology, etc. The two
major extinct groups, Rugosa and Tabulata serve
as important documentary evidence of ancient life
hitherto lost.
Secondly, a few genera and species of all the
three major groups serve as index or guide fossils.
138 Part Two: Major Invertebrate Groups
Calceola sandalina (Middle Devonian: rugose),
Halysites catenularia (Lower Silurian: tabulate)
are the examples. As corals are sessile in habit,
wide geographic distribution of these guide species
or genera point to a meroplanktic stage in the
ontogeny of these animals that are found also in
living forms.
The more acclaimed importance of anthozoans
lie in palaeoecological studies. Scleractinia that
appeared in Middle Triassic provides one of the
most abundant and important constituents of coral
reefs and coral islands in modern seas. These are
found at different points on the earth in the shallow,
warm seas of the present-day tropical belt
(particularly at Australia-New Zealand, Laksha-
dweep-Maldive, Carribean Islands, etc.). These
reefs and islands are carbonate build-ups, main
parts of which are made of accumulations of
skeletons of organisms that live right there. The
prolific growth of the constituent organisms
suggest their successful adaptation to the
environment in which they live and the reefs grow.
Hence, if any carbonate body in a geological
succession may be identified as the then reef, and
if we know the ecology of the constituent
organisms, the conditions of formation of the
ancient reefs may also be inferred.
At present, there are two ecological categories
of scleractinians. Most of the genera and species
can be constituents of coral reefs; they are called
hermatypic corals. A few genera and species, on
the other hand, do not form reefs and live in deeper,
cold waters; they are ahermatypic.
Hermatypic corals live in shallow, warm,
normally saline sea water, clear of fine suspended
sediments. More precise requirements are as
follows:
Depth of water: In very clear water, reefs
may form even at depths of 90 metre. They,
however, grow better at 40–50 metre depth. Most
reef-forming genera and species live most
successfully at depths of 15 metre or less.
Temperature: Reefs form in warm water at
average temperatures between 36°C and 22°C;
25°C–29°C range appears to be the optimum; a
few genera and species may survive for a brief span
at 17°C–18°C.
Latitude dependence: Most coral reefs are
restricted to the tropical belt between 30°N and
30°S latitudes. This is rather due to the suitable
temperature and availability of planktons, food for
corals.
Salinity: Corals are strictly marine. Of them,
hermatypic corals live best within 1 or 2 parts of
salinity of normal marine water of average
35 per cent, salinity; reefs may form, however,
between 27 per cent and 40 per cent.
Clearness of sea water, sunlight and symbiosis
between corals and algae: Coral reefs grow best
in well-lighted sea water. In clear coastal water,
sunlight penetrates upto about 200 metre depth,
though in more turbid water near river mouths with
high amount of suspended fine sediments, sunlight
does not reach that depth. Here the sediments also
clog the hollow space within corallites causing
animals to suffocate and reef growth stalled. In
clearest ocean water, sunlight may penetrate even
beyond 1000 metre.
Restriction of coral reefs to lighted water,
rather the photic zone is more because of symbiosis
between a photosynthetic dinoflagellate
Zooxanthellae (loosely called brown algae) and
reef-forming corals. The former lives in endoderm
cells of coral polyps and through photosynthesis,
consumes carbon dioxide to give off oxygen.
Corals take in that oxygen and give out in turn
carbon dioxide for Zooxanthellae to consume. The
process helps corals to live in proliferation.
Besides, coral polyps and Zooxanthellae
share the relatively small amount of phosphorus
available in sea water and vitally required by
both. The limited supply is recycled alternately by
the two.
In addition, in the shallow depths of photic
zone the amount of different solutes including
calcium carbonate, partial pressure of carbon
dioxide are of such value as to help corals secrete
calcium carbonate more easily as their skeletal

matter. This causes rapid and prolific growth of
coral hard parts that contribute to rapid growth of
the carbonate build-up of reefs.
Currents: Coral reefs need circulation for
their growth. This ensures adequate supply of
nutrients, mainly different kinds of planktons and
other vital requirements such as oxygen,
necessary for corals to live. Moreover, currents
also help in removing suspended sediments,
another favour to corals. However, strong currents
may break down coral skeletons, particularly the
delicate ones.
Exposure to air: Corals can stand only brief
exposure to air and so reefs cannot grow much
above the water surface.
These requirements for coral reefs discussed
above pertain to the modern groups, dominated by
scleractinia. Palaeozoic reefs were formed by
rugose and tabulate corals. It can be assumed that
all these anthozoan groups, extant or extinct, are
and were limited by similar ecological constraints.
But a change in limiting conditions for these
different groups cannot be totally ruled out.
Chapter 7 Phylum Cnidaria 139
Besides, any symbiotic relationship of the kind
modern reef corals have with Zooxanthellae,
cannot be ascertained for Palaeozoic reef-corals
as Zooxanthellae or similar micro-organisms have
lesser preservation potential.
However, as anthozoans have not undergone
any fundamental changes in their history, and rather
evolved with the same body organization and
morphological plan, similar ecological
requirements for extinct and extant groups may be
broadly inferred. On that basis, it is also concluded
that ancient coral reefs of rugose and tabulate corals
were also formed in the then shallow, warm tropical
seas. Distribution maps of ancient coral reefs of
different periods of geological past will, thus,
depict the position and shifting through ages, of
the tropical belts on the surface of the earth. Such
a picture actually tallies with that obtained from
other evidences and theories of continental drift
or rather plate tectonics. It proves the efficacy of
palaeoecological studies on corals and the broad
applicability of the principle of actualism as
discussed in section 3.3.1.
 8
Coiled Shells:
An Introduction
8.1 Introduction
The phylum Bryozoa comes next to the phylum
Cnidaria in regard to complexity of body
organization. Its fossil record is also not really poor.
But we skip over it to discuss a few phyla that are
more important in palaeontology. We start with
Brachiopoda and Mollusca.
The two phyla are considerably different in
fundamental body organization, or in the nature
and disposition of organs and structures of soft
and hard parts of body, so much so that they
demand separate treatment. Even the three major
classes of the phylum Mollusca, viz. Bivalvia
(or Pelecypoda or Lamellibranchia),
Gastropoda and Cephalopoda show wide
variation in morphology and anatomy. But there
are certain considerations for which a common
introduction to the four groups is necessary.
The four groups, viz. Brachiopoda, Bivalvia,
Gastropoda and Cephalopoda, are widely known
in palaeontology as animals possessing calcareous
coiled shells. Barring the subclass Coleoidea of
the class Cephalopoda and a few gastropods, in
all the other members of the two phyla this shell is
external to the body, acting primarily as a protective
140
cover on the latter. In coleoids, the shell is internal
and few like in octopus, as well as some
opisthbranchiate gastropods lack any hard shell.
External or internal, these shells are made of one
or two component parts called valves, which are
relatively simple in structure and serve mainly for
protection and holding the body. They are, thus,
different from what are generally called skeletons,
which, external or internal, have numerous
component parts that are more complex in structure
and organization and are intricately associated with
the soft parts of the body.
However, though basically simple, these
shells are coiled in a vast array of morphological
types. In brachiopods and bivalves, the shell is
bivalved and the coiling is evident only near the
earliest part of ontogeny and the part of the shell
or valve formed at that stage (viz. umbo). But in
gastropods and cephalopods, where the shell is
univalved, i.e. made virtually of a single valve,
the coiling is evident throughout the shell or the
valve. All these four groups are largely marine,
though some bivalves and gastropods are fresh
water-dwelling and at least the pulmonate
gastropods are adapted to air-breathing and are,
thus, terrestrial.

8.2 Molluscan Body Plan and
Variation
Notwithstanding the variation, molluscs are made
on the same basic ground plan of body. It is easier
to understand this organization on the basis of an
example of Neopilina, a genus belonging to the
primitive molluscan class Monoplacophora that
ranges from Cambrian to date. The animal was
found from the deep seas near Denmark. Its body
organization may be considered as the simplest
fundamental type to be found in Mollusca. Hence,
animal of a simpler organization can only be
imagined as the most primitive Mollusca.
This hypothetical archetype (Figure 8.1) would
have a simple cup-like shell on the body, made of
calcium carbonate secreted by a layer of tissue,
called mantle or pallium. Below the mantle, the
body had a mouth at one (anterior) end and anus
at the other, (posterior) end. The latter opened into
a mantle cavity, which contained gills for
respiration. The visceral mass, the fleshy body, is
placed in front of the mantle cavity and extends
below the latter with a flat bottom that helps the
animal for locomotion, the mass itself acting as
foot, in the same manner as is found in gastropods.
All the subsequent mollusca developed on this
basic plan, their variation arising, presumably from
variation in adaptation to different modes of living
and feeding. This may be inferred from
observations on the extant members of different
mollusc groups.
As we know from the present-day
representatives of different classes of molluscan
animals, in two minor classes, Monoplacophora
(Cambrian to Recent) and Polyplacophora (or
Amphineura: Upper Cambrian to Recent), as also
in the major class of Gastropoda (Cambrian to
Recent), the animal uses the visceral mass as foot.
The animals are benthic, slow-moving or
vagrant and are largely deposit feeders. They
have a jaw-like part in the mouth, called radula,
which is used in advanced carnivorous gastropods
for predation and feeding. They also have a head
Chapter 8 Coiled Shells: An Introduction 141
and a nervous system, that help them locate food
materials. They may also develop a siphon system
to get over the disadvantage arising out of the
characteristic torsion in their body (discussed in
more details later).
Bivalves are essentially suspension-feeders
or filterers, living in burrows, blind or closed
downwards away from the sediment-water
interface. They, thus, do not require strong senses
or nerves and neither have head nor jaws. They
are endobenthic as they live in burrows. So, they
do not require any movement; the foot-like
process, which they have, needs only be used to
dig burrows. Gills in these animal also help in
filtering out suspended food material; the mantle
cavity is large enough to store water and they
develop siphons for taking in fresh water and
pouring out its used up foul water. The pallial
sinus in the pallium or mantle is a mark of such
siphons.
Cephalopods are nektic, swimmers, naturally
living on hunting as predators. Their streamlined
bilaterally symmetrical body or shell fit to their
requirement of smooth movement in the fluid
medium; their siphuncle controls buoyancy for
vertical movement through water. They cannot
afford to bear much thick shell adding extra weight;
but then again, to withstand great hydrostatic
pressure and prevent the shell from implosion, the
thin shell needs reinforcement or strengthening
with the help of septa. For locating preys, hunting
and feeding upon them, the animals need well-
organized, sensitive head and brain, tentacles and
sharp, strong jaws. In place of siphons of bivalves
or gastropods, cephalopods develop tubular
hyponome, which eject water in jets to increase
speed. Well-developed head with tentacles is used
as a locomotory aid, helping in propelling
movements.
All this means that, developed on the same
basic ground plan of body, animals belonging to
the three classes, Gastropoda, Bivalvia and
Cephalopoda were adapted to three different niches
(on the substrate, within sediments and in the water,
142 Part Two: Major Invertebrate Groups

(b) Gastropoda
(c) Scaphopoda

(d) Monoplacophora
(e) Amphneura
PHYLUM
MOLLUSCA
(a) Hypothetical
archimollusc

(f) Bivalvia
(g) Cephalopoda

(h) Brachiopoda

PHYLUM BRACHIOPODA

Fig. 8.1 Relationship of molluscan groups with the ancestral hypothetical archimollusc.
Brachiopoda is shown for comparison. For each of the three major classes of Mollusca, viz. Bivalvia,
Gastropoda and Cephalopoda of the phylum Mollusca, the cross-section shows the external shell in
black and features of the body lying inside. Minor groups, viz. Monoplacophora, Polyplacophora
and Scaphopoda, (shell and body not shown) are possibly derived from the same ancestor. (based on
Clarkson 1998).

respectively) with different modes of living and
feeding (vagrant epibenthic deposit feeding for
gastropods; burrowing endobenthic, filter feeding
for bivalves and nektic, pelagic, hunting for
cephalopods) that diverged them into what they
became later and what they are now found as.
Brachiopods belonging to a different phylum
with a different body organization live in
epibenthic, dominantly sessile mode and are,
hence, suspension-feeders. So like bivalves, they
do not have head, jaw, foot, etc.; they have big
mantle cavity; in place of gills they have
lophophores that help them in respiration and food
gathering. Ranging from Cambrian onwards,
brachiopods thus seem to have occupied another
niche and mode of living, to exist along with the
molluscs.
8.3 Shell Growth and Its
Computer Simulation Model
In addition to these, some aspects of shell
morphology throw more interesting lights on
understanding these groups. In all these groups the
first formed shell (or valve) is a sharp conical cup.
Secondly, the shell increases by accretion of shell
material along the margin of the already existing
earlier part (excepting septa in cephalopods and a
few gastropods, which are new parts added to the
existing shell). Had the amount of accretion, that is
the amount of material added, been the same all along
the margin of the circular, elliptical or such other
cross-section of the initial cone, the conical shape
would have been retained all through the ontogeny.
But that is not the case. Different amounts of material
are added to different parts of the margin, so much
so that the shell assumes a coiled appearance.
As mentioned earlier, this coiling is evident
clearly and all along the shell in gastropods and
cephalopods. In brachiopods and bivalves, it is
apparent only in the earliest parts of valves, i.e. in
their umbonal regions. Then again, in cephalopods
the coiled shell is bilaterally symmetrical, the
Chapter 8 Coiled Shells: An Introduction 143
symmetry plane coinciding with the plane at right
angles to the axis of coiling. It means coiling takes
place in that plane, with each whorl (a complete
360 volution or coil) successively going away
from the axis, wound around the earlier whorls.
In gastropods, the shell is asymmetrical, in which
each whorl coiled around the earlier whorl or
whorls is shifted or translated along the axis of
coiling. In result, there is no single common plane
at right angles to the axis to hold all the whorls on
it. However, in a cephalopod or a gastropod shell,
the area of cross-section (called whorl section)
increases at a uniform, relatively small rate, as the
shell grows in size. These attributes of cephalopod
and gastropod shells can be expressed in terms of
three parameters. They are, namely: (1) translation
along the axis (T); (2) rate of increase in the area
of whorl section (W); and (3) the distance of each
whorl from the axis (D). In terms of these
parameters, cephalopod and gastropod shells may
be differentiated as follows. The former shells
have T = 0; in the latter T ¹ 0; in both the cases W
is low and D may vary from small to large
(see Figure 8.2).
Coming to brachiopods and bivalves, both
bivalved with each valve equivalent to the shell of
a gastropod or cephalopod, we have in the former,
shell and valves both bilaterally symmetrical (the
symmetry plane at right angles to the axis of
coiling), i.e. T = 0. But whorl section or cross-
section of the valve (represented by the
commissure) increases very rapidly (W high), while
the distance from the axis or D is low. But in the
bivalve, while shells are generally symmetrical,
valves are asymmetric and inequilateral. The two
other attributes in bivalves are similar to those in
brachiopods. So for bivalves we have T ¹ 0, D is
low and W is high. The four groups can, thus, be
distinguished on the basis of their characteristic
combination of T, D and W. The observations rest
on a computer simulation modelling of coiled shells
undertaken by Raup (1966; 1967; also Raup and
Michelson 1965). Using different values of W
(1–106 ), D (0–1.0) and T (0–4), coiled shells of
144 Part Two: Major Invertebrate Groups

m m

g dp

a
g
(i) a

(i) (ii)

(iii)
(ii)
(a)

(iii) (iv)

(b)

(c)
Fig. 8.2 Minor mollusc groups.
(a) (i)–(iii) Monoplacophora; (i) ventral view, (ii) dorsal view, (iii) lateral view; (b) (i)–(iv) Polyplacophora
(Amphineura); (i) dorsal view, (ii) ventral view, (iii) dorsal and (iv) ventral views; (c) Scaphopoda (m:
mouth, g: gills, dp: dorsal plates, a: anus). (Based on diagrams in Clarkson 1998, Lehmann and Hillmer
1980.)

different shapes were simulated. It was found that
only a small amount of these shapes were attained
by shells occurring in nature. But at the same time
the four groups occupied distinct parts or domains
of the reconstructed shapes.
8.4 Univalved and Bivalved Shell
The study has other bearing on understanding
coiled shell morphology. As mentioned, cephalopod
and gastropod shells increase in cross-section at

uniform, but relatively low rate. It signifies that the
opening of the shell, which is the aperture in reality,
remains relatively small in comparison to the main
shell. Thus, if and when the animal takes refuge
into the shell for protection, the shell can be closed
with the help of a small lid (opeculum) to fit the
aperture. The two, the main shell and the
operculum, are so different in size that the shell
may best be termed univalved.
On the other hand, in brachiopods and bivalves,
the rate of growth of whorl section being very high,
the corresponding opening of the shell becomes too
big for any size of the shell. It, thus, turns out to be
unsafe and ineffective in maintaining the main
function of the shell, namely, protection of the
animal. It then demands development or secretion
of a second component of the shell, equal or nearly
equal to the existing one and lying opposite to it on
the other side of the body, to effectively cover the
latter. The shell, thus, becomes bivalved.
8.5 Orientation and Symmetry of
Bivalved Shells
In bivalves, this bivalved shell is equivalved, though
each valve itself is asymmetrical or inequilateral,
whereas in brachiopods it is inequivalved, with both
shell and the constituent valves bilaterally
symmetrical or equilateral. This difference owes
its origin to the difference in modes of living of the
two groups. Most of the bivalves are slow-moving
burrowers. The sediment in which they make
burrows is a viscous mass on the two sides of the
animal. To penetrate it smoothly, bivalves require
a streamlined bilaterally symmetrical body and
shell. To the front is the dead end of the burrow,
while to the back is the open end at the sediment-
water interface. The animal draws in water, for
food, nutrient and oxygen or discharges it through
opening on this posterior side. The environment
and activities are different at anterior and posterior
ends. This accounts for inequilaterality or
asymmetry of the valves. The posterior side tends
to be larger in these bivalves, because it is
responsible for most of the vital activities of the
Chapter 8 Coiled Shells: An Introduction 145
animal with respective to the organs placed on this
side. On the other hand, the two valves perform
the same function of burrowing and face similar
environments. Hence, they become similar and
mirror image of one another.
Majority of brachiopods and some bivalves
are sessile. In such a case, one of the valves, which
is generally the lower, is the abode and holds the
animal in it. As the main valve, it thus becomes
larger. The other valve, upper or not, has only to
act as a lid on the valve fixed to the substrate. It
can afford to be smaller. Since the attachment
starts early in ontogeny, the animal has less chance
to grow freely on that side; the beak, the earliest
part of a valve or the umbo around it lie here. It
grows freely and maximum only opposite to it.
The sessile animal has no front (anterior) and back
(posterior), as such, as there is no locomotion. But
anterior-posterior concept makes a sense, when
we consider them in regard to the direction in
which the animal grows freely or otherwise. Thus,
in brachiopods, the beak (or the umbo) lies at the
posterior, the opposite end being termed anterior.
But the environments to the right or left of the
shell (or valves) are similar, defining a bilateral
symmetry between them. Inequivalved, equilateral
nature of brachiopod shells are thus explained.
Sessile bivalves are also inequivalved, but their
valves maintain the asymmetry, more intrinsic in
bivalves.
The above discussion highlights the facts that
even though there are fundamental differences
between the two phyla of Brachiopoda and
Mollusca, there are significant similarities. Their
adaptation to different niches with different modes
of living and feeding brings in morphological
differences. In addition, differences in the rate and
pattern of growth of body and shell, when judged
in conjunction with the effects of adaptation,
provide a common framework to understand
morphology and its variation in these two phyla,
rather in the four groups referred above. Details
may be judged on this basic framework.
The whole discussion and some more
information have been summarized in Factsheet 8.1.
 FACTSHEET 8.1

146
Unity and Diversity in Four Multicellular, Invertebrate, Coiled Shell-Bearing Animal Groups

PHYLUM BRACHIOPODA MOLLUSCA

CLASS BIVALVIA GASTROPODA CEPHALOPODA

Biological Eukaryotic, multicellular, invertebrate metazoa

identity

Geological age Lr.Cambrian-Recent Ordovician-Recent Cambrian-Recent Up.Cambrian-Recent

Major Class: Articulata, A few orders Order: Archaeogastropoda; Subclasses; a few in
divisions Inarticulata/ Recently Caenogastropoda; Pulmonata zoic+ Nautiloidea,
added subphyla(a) Ammonoidea, Coleoidea

BODY
Body position Body cavity (Bcv) Visceral mass (Vsm) Vsm with gut Vsm with gut
Part Two: Major Invertebrate Groups

posterior with gut anterior anterior posterior

Mantle Cavity MC anteriorly placed MC posterior below Vsm MC anterior above head MC anterior below Vsm
(MC)

Mouth (M) M on the wall anterior, M anterior M anterior M anterior

between Bcv and MC

Anus (A) A in inarticulates, A posterior A anterior, above * mouth A anterior, below * mouth
absent in articulates

Head and jaw No jaw in head/M No head or jaw Well formed head and jaw ** Well formed head and jaw **

Respiration Lophophore in MC Gills in MC help respiration One group of gastropods has ‘lungs’,
helps in respiration i.e. respiratory cells on mantle walls

(
a) Phylum Brachiopoda
Subphyla Linguliformea ü
ý Inarticulata Rhynchonelliformea } Articulata * See Page 147 for related feature
Craniformea þ ** See Page 148 for related feature

(Cont...)
 FACTSHEET 8.1 (Cont...)
Unity and Diversity in Four Multicellular, Invertebrate, Coiled Shell-Bearing Animal Groups

PHYLUM BRACHIOPODA MOLLUSCA

CLASS BIVALVIA GASTROPODA CEPHALOPODA
Shell
Characters
Position, Shell external; covers the body; bivalved; pedicle (P) Generally external, absent or internal in some;
component, and brachial (B) in brachiopods; right and left in houses the body or supports it in internal types;
utility bivalves univalved; completely/partially empty.
Symmetry Shell and body bilaterally symmetrical Both asymmetrical Both bilaterally symmetrical
Symmetry Across the body, Along the body and None due to an early *Vertical in life position
shell and valves between the valves ontogenetic twist*
Coling Coiling of shell/valves evident only at umbo Coiling distinct all along the shell; uncoiled /straight in some;
planispiral conispiral generally coni-/trochospiral planispiral

VALVES
Symmetry Each valve equilateral/ Each valve inequilateral / Single valve, asymmetrical Single valve, symmetrical
symmetrical asymmetrical
Position Ventral-dorsal Right-left w.r.t body Virtually dorsal Surrounds body except at
(above/below the body) anterior oral end
Chapter 8

Opening Muscles open/close Muscles close, ligaments Aperture may be closed

and closing the shell open the shell by operculum by muscles
Some major Hinge in articulates; Hinge with teeth and sockets Septa may be present in Both septa and siphuncle
characteristics P-valve has teeth, in the same valve; very long shells;
B-valve sockets; dentition asymmetric; no siphuncle
articulation by muscles some edentate
in inarticulates

GROWTH Shell/valve starts as a shallow cup/cone; grows by accretion, i.e. by addition of hard mineral substances of shell
pattern material along the margin; septa in cephalopods are added structure.

Growth Translation along the axis of coiling T ; Whorl expansion W; Distance of the generating curve from the axis D
Coiled Shells: An Introduction

parameters {H-High; L-Low}

T=0: W-H: D-L T¹0:W-H:D-L T ¹ÿÿ0ÿ:ÿW - L : D - L to H Tÿ=ÿ0ÿ:ÿW - L : D - L to H
(Cont...)
147
 FACTSHEET 8.1 (Cont...)

148
Unity and Diversity in Four Multicellular, Invertebrate, Coiled Shell-Bearing Animal Groups

ECOLOGY
Habitat, mode of living, feeding habit
Brachiopods: Marine; Suspension feeder; Benthic,epi/endo-; Fixosessile: Plani-rhizopedun culate/encrusting;
Liberosessile: Ambitopic (freelying); Infaunal (burrower); quasiinfaunal; interstitial
Bivalves: Marine-fresh water; Filter-feeder, some deposit feeder; Generally endobenthic; shallow/deep burrower;
Epibenthic: byssate/cemented/freelying/boring/cavity dwelling in rock/wood; Pelagic (swimmer: temporary)
Gastropods: Marine-fresh water; some land-dweller; Deposit-feeder: Also filter-feeder; Epibenthic: sluggish vagrant; crawler, grazers;
also sedentary/parasitic/predatory**
Cephalopods: Marine; Predator, i.e. hunter**/carnivorous,** hunter/scavenger; Pelagic, fast swimmers; predatory; a few slow epibenthic
Morphology related to mode of living
Brachiopods: Pedicle/spines for attachment; lophophore/brachidium for food-gathering/respiration
Bivalves: Siphon for water for food and respiration (pallial sinus its mark on shell)
Gastropods: Septa in long shells; no siphuncle; slitband/canals for water for gills
Part Two: Major Invertebrate Groups

Cephalopods: Septa for stronger shell (hydrostatic advantage): siphuncle for buoyancy control (hydrodynamic advantage)
ADDITIONAL INFORMATION
Hard part composition
Brachiopods: Calcite in articulates; chitinophosphatic in inarticulates
Bivalves: Calcite and/or aragonite plus conchiolin
Gastropods: Mixed/layered aragonite; outer conchiolin
Cephalopods: Aragonite: odd parts calcitic (e.g. belemnite guard)
Periods of importance
Brachiopods: Inarticulates in Cambrian; Articulates in Permo-Carboniferous; many in Ordovician/late Mesozoic
Bivalves: Mesozoic to Recent; rudists in Cretaceous
Gastropods: Archaeogastropod in Ordovician; Caenogastropod
post-Palaeozoic; particularly Tertiary-Recent
Cephalopods: Nautiloids in Ordo-Silurian; Ammonoids in Caronif,
(goniatites); Triassic (ceratites) Jurassic (ammonites); coleoids late Mesozoic-Recent
Special features
Brachiopods: Simple basic design, but great diversity of forms and lifestyle on filter-feeding; homeomorphy as well; adaptability in articulates;
conservatism in inarticulates
Bivalves: Shell form and mode of living closely related; successful burrowers; almost total sensory deprivation due to sedentary lifestyle
Gastropods: Torsion governs biology and shell morphology; homeomorphy; many fossil species not natural
Cephalopods: Efficient buoyancy regulation and attitude correction; dimorphism; heteromorphism; good stratigraphic control; evolutionary
pattern distinct and well-studied.
 9
9.1 Introduction
Brachiopods are eukaryotic, multicellular,
invertebrate metazoan animals. They are strictly
marine and benthic. The phylum, they belong to, is
poorly represented in the Recent times, though at
different points in the geological past
(for instance, in Permo-Carboniferous) they evolved
rapidly and abundantly, to be represented by a large
number of genera and species of varied morphology.
In the introductory discussion (Chapter 8), it has
been indicated that brachiopods are dominantly
fixosessile epibenthic, though some may be free-
lying or ambitopic while there are also endobenthic
or quasi-endobenthic forms to be found in the
phylum (there are more variations to be discussed
later). The animals are suspension-feeders and lack
head, jaw or foot. They bear external shell, which is
bivalved, inequivalved, yet equilateral, hence
bilaterally symmetrical, across the shell or the valves.
The morphology of these brachiopod shells
can be considered in the following frame:
1. Position, shape, symmetry, dimension,
orientation, etc. of the shell and valves.
2. Hinge and associated features or their
alternatives.
3. Features inside the shell.
4. Features on the surface of the shell.
149
Brachiopoda
9.2 The Valves
In bivalved, inequivalved shells of brachiopod, the
two valves are placed, broadly speaking, dorso-
ventrally with respect to the body. To work smoothly
during lifetime, that is, to open and close the shell
for food-gathering and other activities, the valves
require to be attached to each other along some line.
In that case, the shell opens at the opposite end. The
said line, thus, acts as a hinge (of the type found in,
say, a briefcase), itself being called the hinge line.
The symmetry is per-pendicular to this hinge.This
kind of hinged or articulated shell is found in
articulate brachiopods (earlier Articulata was
considered as one of the two subclasses, the other
being Inarticulata; for present opinion, see
Factsheet 8.1); inarticulate brachiopods lack any
hinge in their shells. The valves are then held
together with the help of muscles only.
Formerly, the two valves were called ventral
and dorsal according to their position with respect
to the body. But it was subsequently found that in
a large number of articulate brachiopods, the
animal undergoes a torsion during its embryonic
stage. This torsion changes the ventro-dorsal
position of the valves, whereby the hitherto ventral
valve becomes dorsal in subsequent stages of
ontogeny and vice versa. The terms, thus, lose
150 Part Two: Major Invertebrate Groups

significance. Hence, in later studies the ventral features, viz. hinge, an external feature and bra-
valve is referred to as pedicle valve and the dorsal chidium, an internal one. Both hinge and
valve as brachial. The former, which is also the brachidium have already been referred in section
larger valve, has the pedicle attached to it, using 9.2. To add here, brachidium is a spring-like
which the shell remains fixed. To the latter, the skeletal structure that acts as the frame for
smaller valve, is attached the brachidium, an lophophore of brachiopods. Formerly, it was
important internal structure of the shell. thought that the animal can throw the lophophore
out of the shell for the purpose of food gathering
9.3 Appearance and Measures and respiration. But later it has been concluded
that brachiopod shells do not open as much as to
The geometrical shape of the equilateral valves of throw lophophore and brachidium out of it. Rather
most brachiopod shells may be defined on two it draws in water through slightly gaping valves
criteria: (i) commissure and (ii) lateral profile. creating current with the help of numerous fine
Commissure is the line along which the two valves hair-like projections on the surface of the
are in contact in a closed shell. Lateral profile is lophophore. Nevertheless, the changed status does
drawn or conceived along the plane of symmetry, not reduce the importance of lophophore or
which is also perpendicular to the plane of brachidium in the vital activities of brachiopod
commissure; different types of lateral profile are animals.
listed in Factsheet 9.1. The geometry of these two The shape and alignment of brachidium vary
mutually orthogonal closed curves, thus gives the in different genera being characteristic for them.
three-dimensional shape of the shell (Figure 9.1). They not only determine the shape of the brachial
Variation in the shape of brachiopods shells is valve to which it is attached; the shape of the
shown in Figure 9.1. The three dimensions of the pedicle valve is also determined by the shape of
shell, length, breadth and thickness, also depend brachidium. Figure 9.2 shows a few examples of
on these two curves; commissure and lateral how shell-shape is determined by hinge and
profile. brachidium. It should be mentioned here that the
The latter two are external characters of the shape of some brachiopod shells or their valves is
shell and are themselves defined by two other also controlled by ecological constraints. Thus,
burrowers like Lingula have a slightly convex shell
with a rectangular commissure that acts like a razor
FACTSHEET 9.1 to move through soft muddy sediments. On the
Brachiopod Shells in Lateral Profile
other hand, genus Hercocestria is cemented by the
larger valve, which has an aberrant tubular shape,
Lateral Pedicle Brachial with the smaller valve set as a lid on it.
Profile Valve Valve
Biconvex Convex Convex 9.3.1 Orientation
Ventriconvex More convex Less convex
Dorsiconvex Less convex More convex For reasons already mentioned in section 9.2,
Concavoconvex Convex Concave brachiopod valves are called pedicle and brachial,
Convexoconvcave Concave Convex in place of the former ventral or dorsal designation.
Resupinate 1 Convex (at umbo) Concave In fact, many pedicle-bearing brachiopods are held
Concave (other end)Convex with the commissure and two valves in a vertical
Resupinate 2 Concave (at umbo) Convex position, making ventro-dorsal terminology still
Convex (other end) Concave redundant.
 Chapter 9 Brachiopoda 151

1
2a

5 5 (iii) (iv)
4 (ii) (v)
(i) (a)

6
7 (ii)
(i) 6
2b

(iii) (iv)
(b)

(i) (ii) (i) (ii)

(c) (d)

(i) (ii) (ii)

(i)
(e) (f)

Fig. 9.1 Variation in brachiopod shells shown in views, viz. (i) pedicle, (ii) brachial, (iii) umbonal,
(iv) anterior and (v) lateral.
Commissure Profile Ornaments Other
(a) Subcircular Biconvex Concentric Large foramen
(b) Triangular Biconvex Concentric and radial Strong median ornament
(c) Semicircular Resupinate Mainly radial
(d) Subcircular Convexoplane Concentric and radial
(e) Semielliptical Concavoconvex Concentric and radial
(f) Semielliptical Convexoconcave Concentric and radial
Number index: (1) Foramen; (2a) Non-strophic hinge; (2b) Strophic hinge; (3) Posterior; (4) Anterior
(5) Concentric ornament; (6) Median ridge; (7) Median sinus; and (8) Radial ornament.
152 Part Two: Major Invertebrate Groups

(a)
(b)

(c) (d)

(e)

Fig. 9.2 Brachidium and shell shape in brachiopods.

(a) Transverse biconvex shell, outwardly directed long brachidium, (b) Longitudinal biconvex shell,
low outwardly directed brachidium, (c) Slight transverse concavoconvex shell, low brachidium directed
towards pedicle valve, (d) Transverse biconvex shell, low brachidium directed towards pedicle valve,
(e) Subcircular biconvex, simple brachidium.

It has also been indicated in section 8.5 that as
the sessile animals do not move, the anterior-
posterior direction cannot be designated normally.
The animal is hinged and is also attached to the
substrate from early ontogeny onwards and grows
minimum there. It grows freely and the maximum
amount of material is accreted to the shell, in
diametrically opposite direction along the plane of
symmetry. The latter in which the valves grow
freely is then called anterior and the hinge and
pedicle (or beak or umbo, explained later) mark
the posterior. On the two sides of the symmetry-
plane, accretion is the same and gradually increases
from posterior to the anterior on either side.
The three dimensions, length, breadth or width
and thickness or depth are measured mutually
orthogonally, anterior-posterior maximum
dimension along the plane of symmetry being the
length and breadth perpendicular to it in the plane
of commissure (Figure 9.1).
The shape of brachiopod shells become
important in respect to the phenomenon of
homeomorphy, i.e. similar external morphology
attained by different unrelated genera or species of
the same geological time or of different times.
However, as it demands more elaboration,
homeomorphy will be discussed later.
 Chapter 9 Brachiopoda 153

2
2
3

5 4

1
1 3

Fig. 9.3 Muscle attachment of two valves in brachiopod shells.

Different features and modifications. For inflated biconvex shells thin tendon or raised platform in
place of long muscles, or short muscles for flatter shells are special modifications.
Number index: (1) Adductor muscle; (2) Diductor muscle; (3) Pedicle;
(4) Cardinal process; (5) teeth and socket;

9.4 Brachiopod Hinge
As mentioned, articulate brachiopods have a hinge
structure, while inarticulates do not. The hinge not
only holds the two valves permanently, the valves
operate, that is, open and close about it. The hinge
marks posterior of the shell.
Brachiopods with or without hinge show a few
more features at or around the posterior margin.
Some of them need introduction, before getting
into details of hinge, others will be discussed in
due course.
All kinds of brachiopod shells start to grow
from an initial small cup. The apex of that cup is
referred as beak. A small to large area around the
beak is abruptly more convex (or concave in the
cases, e.g. brachial valve of Productus or
Rafinesquina) than the rest of the valve. This area
is called umbo or umbone. Its curvature (convex
or concave), prominence (highly curved, moderate
or flat), etc. are often characteristic of genera. Besides,
as the brachiopod shell is inequivalved, the umbones
of pedicle and brachial valves are not the same in
curvature or prominence. Thus, in Productus, in
contrast to the low, concave brachial umbo, the
pedicle umbo is highly convex and incurved.
More primitive brachiopod hinge is called
strophic hinge. In this case, both the valves have
straight margins along the hinge and these two
straight lines lie in contact with each other to make
the strophic hinge. The hinge line itself is straight,
and acts as the hinge axis. In most of the instances
the hinge line is equal to the breadth of the shell
or valve. To hold the two valves together, there
are teeth and sockets on the hinge; the former lies
only in the pedicle valve and the latter in the
brachial valve, fitting into the teeth of the pedicle
valve. As the two valves are in contact along the
whole length of the hinge, the teeth and sockets
need not be very large or strong to keep the valves
together.
Early articulate brachiopod genera or species
and some later ones have strophic hinges. Barring
them, most articulate brachiopods have a non-
strophic hinge. In this type, there is no real hinge
line in either of the two valves. They have curved
margins near the hinge and are hinged only at
two points that act as two fulcra (a loose
comparison of this type may be made with
skylights, where the windowpane is hinged at two
fulcra on the frame). On the pedicle valve, there
are two teeth at these two points of hinge and the
brachial valve has two sockets fitting into the teeth.
As the valves are hinged only at those two points,
the teeth and sockets need to be strong, large and
curved to operate effectively. After the animal dies,
these teeth and sockets of non-strophic hinge hold
the valves strongly together, whereas in strophic
brachiopods, the small teeth and sockets along the
straight hinge line fail to keep the valves together.
This is why fossils of non-strophic forms are
preserved more with closed shells, whereas in
strophic brachiopods, the valves are more often
preserved separately in fossils.
154 Part Two: Major Invertebrate Groups
In both articulate and inarticulate forms, the
shell opens or closes only with the help of muscles.
Fossils do not preserve muscles, but there are
muscle scars on the interior of the shell. Their
position also brings about some changes in the
exterior of the shell, particularly of umbo. We will
deal with them later.
Interarea is one of the external features on
the posterior margin. It is a flat triangular area
between the beak and the hinge line of a valve.
Palintrope is a similar, but curved surface between
the beak and the hinge line. Generally, the term
interarea is used in the case of strophic forms with
straight hinge line. Palintrope is then reserved for
non-strophic forms with curved hinge line. There
is, however, difference of opinion on these terms.
As in the case of umbones, interarea or
palintrope, if and when present, are not similar in
the two valves. It may be present only in the pedicle
valve or, if present on both, is larger on the pedicle
valve. This is an important feature on which a closed
brachiopod shell may be differentiated from a
closed bivalve shell. In the latter, the interarea of
the two valves are generally similar, as the shell is
equivalved.
9.5 Pedicle and Its Opening
Majority of brachiopods are pedunculate,
attached to the substrate with the help of pedicle.
In plenipedunculate forms, this pedicle is a
strong or tough rod-like structure, with a hard
covering and softer tissues within. It is resistant
to a fair intensity of current. Pedicle is attached
to the interior of the larger valve and comes out
through an opening called pedicle opening,
which may be confined to the pedicle valve (e.g.
Terebratula) or shared by both valves (e.g. in
Strophomena). In the latter case, however, the
larger portion of the opening is shared by the larger
or pedicle valve. There are brachiopods, in which
the pedicle is a brush or root-like in structure,
comparable to byssus of bivalves that are pushed
into oozes for anchorage or help attach the shell
to dead shells on the substrate. These are called
rhizo-pedunculate (e.g. Chilidonophora, a living
terebratulid).
The pedicle opening is basically triangular with
apex towards the beak and base towards the hinge
line. It is called delthyrium. The pedicle itself has
a circular cross-section. So, when it emerges
through the triangular delthyrium, there may be a
number of alternative cases. If the pedicle is too
thin for the delthyrium, that is the circular cross-
section is too small for the triangle, it requires some
support to keep the appendage steady while
emerging through the delthyrium. Some amount of
shell material is secreted to cover up the remaining
gap within the delthyrium, either in the form of
one plate (deltidium) or more than one, generally
two plates (deltidial plates). If dethyrial triangle
is smaller than the pedicle cross-section, there
develops a smaller triangular opening on the
brachial valve. Pedicle then emerges through the
diamond-shaped opening shared by both the valves.
The opening on the brachial valve is called
notothyrium and any cover on it, similar to
deltidium on the pedicle valve are called chilidium
or chilidial plates (see Figure 9.4).
In genera like Magellania (a recent
terebratulid), delthyrium though covered by
deltidial plates bears a large circular opening at the
top. Pedicle emerges through this, called pedicle
foramen or simply foramen.
9.6 Internal Features
9.6.1 Brachiopod musculature
Brachiopod shells open or close with the help of
muscles. Different kinds of muscles perform
different functions. They leave scars on the interior
of the shell, whose shape and size are determined
by the function the concerned muscle performs,
and how thick the shell or valve material is.
The scars, thus, help identify the muscles.
Musculature is different in articulates and
inarticulates; it is also genus or species specific.
 Chapter 9 Brachiopoda 155

6
1
2
2 9
3
4 3 10

7 (ii) (iii)
(i)

8
11

(iv) 5
(a) (v)

1 2
3

(i) (ii)

4 5

(iii)
(iv)
(b)
Fig. 9.4 Brachiopod shells.
(a) (i) Primary and secondary shell layers in brachiopod shells; (ii) Impunctate shell; (iii ) Pseudopunctate
shell; (iv) Endopunctate shell; (v) ‘Inarticulate’ shell
Index: (1) Periostracum; (2) Primary shell layer; (3) Secondary shell layer; (4) Epithelium (cellular);
(5) Alternate phosphatic and organic matter in ‘inarticulate’; (6) Calcite fibres; (7) Generative zone;
(8) Caecum giving way to punctae; (9) Taleolae; and (10) Endospine.
(b) Pedicle opening
Index: (1) Open delthyrium; (2) Foramen; (3) Deltidium; (4) Deltidial plates; and (5) Delthyrium and
notothyrium.

Muscles work only by contraction. Thus, in
articulate brachiopods, the shell closes when
adductor muscles contract. They leave two scars
on the pedicle valve and four on the brachial valve,
as each adductor muscle is divided into two before
reaching the brachial valve. Since the two valves
are more permanently attached at the hinge or near
the posterior margin, the shell closes or opens at
the anterior end and so the adductor muscles need
to be placed to the front of the hinge (Figure 9.3),
at right angles to the valves. Adductor muscles may
be of two kinds: quick adductors work as reflex to
sudden incidents, catch adductors can hold the
valves firmly for a longer time.
The shell opens at the contraction of diductor
muscles. It is attached to the pedicle valve anterior
to the hinge and just outside the adductors, but in
the brachial valve the attachment lies to the
posterior of the hinge. In many articulate
brachiopods, there is a cardinal process in the
brachial valve, hard and calcareous. The diductor
muscle ends at this cardinal process. When
diductor muscles contract, the adductors relax; the
cardinal process below the brachial beak is tucked
into the gap below the pedicle umbo. Thus, at the
posterior end of the shell the two beaks are drawn
towards each other as a result of which the anterior
ends are pulled apart to open the shell. Adductor
156 Part Two: Major Invertebrate Groups
muscles are then put in tension and they contract
to release it. This closes the shell.
In inarticulate brachiopods, the two valves
close by adductor muscles. There are a single
posterior muscle and a pair of anterior ones. When
they relax, the valves slightly move apart. They
come closer to close the shell as the muscles
contract.
Adjustor muscles form a third kind.
Pedunculate forms bear this muscle. It is attached
to the pedicle and can move the latter. By that it
helps the shell or the valves to change their
position. This is particularly important to help the
animal move in the direction of current in the water
to collect suspended food material for the filter-
feeding animal.
In inarticulates, there are also different types
of large and well-developed oblique muscles to
rotate, shear or slide the valves against one another.
Together they make the musculature strong in
inarticulates.
In some articulates and craniiform
inarticulates, muscles are not directly attached to
the valves. Rather there is a sort of muscle platform
on which the muscles rest. In a few forms, where
the valves are highly convex, leaving a large space
inside them, muscles are reduced and joined instead
by thin, yet strong tendons.
9.6.2 Lophophore and brachidium
Lophophore and brachidium make another system
of internal structures in brachiopods. Placed within
the mantle cavity, lophophore is the main organ of
brachiopods for food gathering and respiration. In
many articulate groups, a hard, mineralized bra-
chidium provides a skeletal framework for
lophophore. Both brachidium and lophophore are
loop-like or coiled-like spring and can be moved
with the help of muscles. On its surface,
lophophore has numerous hair-like cilia that
constantly move to create movement in the ambient
water. This provides the animal with fresh water
for respiration, as also helps the animal discharge
the excess or used-up water. The point will be
further discussed in relation to surface ornaments
of brachiopods; here it may be added that the issue
defines one importance of lophophore. Another
importance of lophophore-brachidium couple lies
in their controlling the shape of shells as discussed
in section 9.3.
9.7 Mineralogy and
Microstructure of Shells
Before going to surface sculpture (structure/
ornaments: use is optional), a brief treatment on
mineralogical composition and microstructure of
brachiopod shells may be helpful. Details of
accretory growth pattern of brachiopod shells are
better understood on this knowledge of shell
mineralogy and structure.
Calcareous shell of articulate brachiopods has
three layers. The outermost layer is a thin
periostracum, made of proteinaceous material. In
living animals this layer is covered by a gelatinous
sheath of mucopolysaccaride. It is the first element
to form and it protects the growing edge of the
shell. Both protein and gelatin being organic
compounds have least preservation potential in
fossils. The next inner layer is called the primary
layer. Made of rather structureless crystalline
calcite, the layer is of constant thickness. It is
succeeded inwards by the secondary layer made
of fine fibrous calcites, stacked regularly and each
with a trapezium-shaped cross-section. It is
secreted throughout the life, is the thickest below
the umbones and gradually thins down towards the
margin. Along the margin of the mantle of
brachiopod body, there is a generative zone with
shell-secreting cells. These secrete the shell layers,
first, the gelatinous sheath and then successively
the three layers, from periostracum to the
secondary layer.
Different groups of living and fossil articulate
brachiopods differ in details of this general plan
of development and constitution of shell materials.

Variations are often diagnostic at higher taxonomic
levels, viz. orders. Besides, calcareous shells of
articulate (rhynchonelliform) and even craniiform
inarticulate brachiopods are made of calcite. This
makes them more stable as fossils than shells of
another common bivalved fossil group of Bivalvia.
In the latter, shells are made of metastable mineral,
aragonite and, hence, are often lost to leave only
the molds and casts.
Among inarticulates, craniiforms (e.g. genus
Crania) have calcareous shells in which all the
layers found in articulates occur, though with
different importance and microstructure. Relatively
little is known about shell formation and
compositional variation in linguliform
inarticulates. Two principal types of microstructure
are found. In one, the primary layer is made of an
admixture of chitin and calcium phosphate (e.g. in
genus Discinisca); in a second type those two
constituents are interlayered (as in Lingula).
9.8 Punctation of Shells
Punctae make one of the important characteristics
of microstructure of brachiopod shells. In many
articulate and calcareous inarticulate shells, fine
hollow tubular features are found to run across the
shell layers. They extend from the body up to the
periostracum. During lifetime, these tubes are
occupied by extensions from the mantle. Made of
organic compounds such as protein, glycogen, etc.
the materials within punctae are used to help
respiration and to store nutrients. There are also
some ‘toxic’ compounds that lend to the animal a
sort of protection from predators. Shells with such
punctae are called endopunctate, or simply
punctate. In fossils, where the periostracum layer
is generally absent, these punctae are seen on the
surface as very fine circular depressions.
Brachiopods without punctae in shells are called
impunctate. In certain others, like strophomenid
brachiopods, the shell layer is traversed by a set of
irregular calcite rods (taleolae), which on the shell
Chapter 9 Brachiopoda 157
surface looks like punctae. These shells are termed
pseudopunctate, though taleolae has nothing to
do with punctation (Figure 9.4).
In brachiopod history, a few groups (e.g. orders
such as Pentamerida, Atrypida, Athyridida) are
typically impunctate. A few orders are only
punctate (e.g. Spiriferinida, Terebratulida). But in
many others (Orthida, Spiriferida, etc.) punctation
is found to have evolved in course of evolution of
some impunctate lineages. It is because of this that
Cooper’s scheme of classification of brachiopods,
once much used, has now been done away with.
9.9 Surface Features
Articulate and inarticulate brachiopods also differ
in regard to surface features of the shells. The latter
have generally simpler surface. But simple or
complex, whatever be the surface features, the
variation in shape and surface of brachiopod shells
are limited to fewer types. Brachiopod animals
themselves are limited to a relatively small range
of size; they can not neither be too big nor too
small. Naturally, the shells secreted by such
animals are limited in their surface area. All the
variations in surface features are controlled by
these small ranges of the volume of the animal body
and the area of its surface. Combinations cannot
be too many.
Shell surface in brachiopods shows three kinds
of features (as indicated elsewhere, structures,
sculptures or ornaments are alternative to features,
connoting the same), viz. (i) concentric, (ii) radial,
and (iii) median. Concentric features are generally
grooves developed concentrically (rather
confocally) about the beak. In a few cases, as in
Productus, concentric features may be represented
by coarse ribs. Radial features diverge away from
the beak; median feature is actually a particular
radial feature developed in the plane of symmetry.
Both these types are represented by fine or coarse
ribs, costae, costellae or plications. These surface
features are diagnostic of genera and species.
158 Part Two: Major Invertebrate Groups
Median feature is commonly a sinus or sulcus in
the pedicle valve and a ridge or fold in the brachial
valve, fitting to the corresponding feature of the
pedicle valve. Clarkson (1998), however, uses fold
for the feature on the pedicle valve and sulcus for
that on the brachial valve.
Concentric features represent growth stages of
the valves. Radial type has different significance.
The amount of shell material that is added to each
earlier stage of growth by accretion has four vector
components, viz. horizontal, vertical, radial and
forward (i.e. towards anterior). Along the symmetry
plane, however, radial and forward become one
and the same. There is no radial ornament to
develop, if the four vectors maintain a balance.
The margin of the shell, represented by a concentric
feature, remains a smooth curved line
(rectimarginate). But if the vertical component is
greater, the surface becomes folded or crenulated.
The more the vertical growth, the more pronounced
would be the folds on the margin.
Now since surface features are genera or
species specific, the question that arises is: why is
it that some forms have radial features, others do
not. As said, the area of the shell surface is
determined by the volume of the body. If for any
inherent or genetic reason, not known as yet, the
material secreted by a genus (or a species) and,
thus, the area of the shell surface proportionate to
that amount of material, is greater than the surface
area of the body (it means that the rate of growth
of shell overcomes the rate of growth of the body),
the extra material can only be accommodated in a
vertical growth of the surface. It gives rise to radial
features.
There is also a functional importance of radial
features. When the shell opens for taking in water
for feeding and respiration, the two valves are
separated from each other. For a shell of a
particular breadth (i.e. size of the shell and body
too), the length of the margin of a smooth-surface
shell will be less than that of a shell with radial
features on the surface. This, in turn, signifies that
for the equal amount of separation of valves, i.e.
equal amount of opening of the shell, the radially
ornamented shell will take in more water proving
more efficiency. In other words, such shells will
require slight opening of shells to take in adequate
water. Besides, such a slight opening will allow
only water and finer suspended material to pass
through. Larger particles, detrimental to the
animal, can only pass through the hinges of the
folds of radial features. But very fine setae or
spine-like elements located at those points prevent
such particles from entering the body. This way
radial ornaments also help the animal in filtering
out unwanted particles. Factsheet 9.2 provides a
concrete case study of functional morphology of
brachiopod fossil.
FACTSHEET 9.2
A Case Study of Brachiopod
Functional Morphology
Cryptorhynchia is a Middle Jurassic genus from
Kachchh.
The genus is strongly ribbed on surface:
multicostate brachiopods indicate adaptation to
shallow, agitated carbonate shelves (Almeras
1987). It has medium sized foramen, a moderately
biconvex shell and a weakly incurved umbo that
indicate epifaunal life mode.
It is associated with coral skeletal banks, sponge
meadows and oolitic barrier bars that provided a
hard substrate.
Epibionts like oysters and serpulid annelids are
attached to Cryptorhynchia shell random over
pedicle valve, but never across the commissure or
in the posterior part of brachial valve, suggesting
pre-mortem infestation; as the organism rested on
the substrate with the umbo directed downward.
This position elevates the median fold and projects
it horizontally towards the current direction. It also
prevents epibionts from reaching the umbonal part
of lower, brachial valve. This mode is functional in
a crowded environment (as preferred by gregarious
mytilid bivalves).
Summing up, it is concluded that Cryptorhynchia
lived in nests on shallow, unstable carbonate shelf
(Mukherjee et al., 2002)

External morphology of shells, including
shape and surface features, thus, depends on the
volume of the body and the area of the surface.
Both these measures have limited range of
variation in brachiopods. And so it is found in the
history of the phylum that genera of the same age
or different ages, often develop similar external
morphology. This is called homeo-morphy, which
may be isochronous or heterochronous
(see Factsheet 9.3).
FACTSHEET 9.3
Homeomorphy in Brachiopods
Similarity in external morphology (viz. shape,
surface features) found in unrelated genera is called
homeomorphy.
Example:
Productorthis of Middle Ordovician;
Order Orthida
and
Dictyoclostus of Upper Carboniferous;
Order Strophomenida;
Two genera similar in semielliptical commissure,
concavoconvex lateral profile, long, but very narrow
interarea and fine radial costae; different in
punctation, Productorthis being impunctate and
Dictyoclostus pseudopunctate.
Following two kinds of homeomorphy are
recognized:
Isochronous, i.e. of the same age : common in
Jurassic
Heterochronous, i.e. of different ages.
Median feature that lies on the symmetry is
deep in some genera (e.g. Rhynchonella). It is also
associated with much plicated anterior and lateral
margins. In all likelihood, water enters these shells
from the lateral margins, bathes the brachidium
and comes out as foul water through the median
sulcus.
Surface features of a different kind are spines,
hollow or solid. In some genera (e.g. Productus),
spines occur on pedicle valve; in some others, they
lie along the posterior margin of the hinge area
(interarea) (e.g. in Chonetes). In the first case, the
Chapter 9 Brachiopoda 159
spines, acting like spikes of snow-shoes, probably
helped the shell to float in viscous sediment in the
quasi-infaunal mode of living of the genus. In the
second, the spines are interpreted as a means to
regulate the position of the shell amidst currents.
In genera like Acanthothyris, spines are hollow and
have extensions of mantle material within them.
They may have acted as sensory organs in face of
adversities or enemy.
9.10 Additional Information
Beyond Morphology
9.10.1 Ecology and palaeoecology
Commonly known as typical fixosessile
epibenthics, brachiopods, however, show quite a
wide range of adaptations. Most of the articulates
(rhynchonelliforms) and craniiform inarticulates
are fixosessile that remain attached throughout life.
Of them there may be a few types. In majority
cases, a simple rod-like pedicle (or peduncle)
serves for attachment (plenipedunculate); in a few
others, the pedicle ends in a bunch of root-like
processes which help attachment (rhizo-
pedunculate) (see section 9.5). In another type,
long, thin pedicle may hold the shell floating in
water, much like a flying kite in air, to add special
advantage to filter feeding. Fixosessile forms may
also be encrusting/ cemented. In genera like
Crania, where pedicle is absent, the pedicle valve
is cemented; some productids are cemented only
in early parts of ontogeny, whereupon they are
cemented only at the umbonal region. Some genera
are attached by spines, discussed in section 9.9.
Liberosessile or free lying forms (also called
ambitopic) include those in which pedicle opening
is closed (many strophomenids, some spiriferids
and orthids). They could survive only where
burrowing as organisms were absent or sparse.
Linguliform inarticulates are burrowing in
which Lingula shows a unique method of
burrowing, as described in Factsheet 9.4.
160 Part Two: Major Invertebrate Groups
FACTSHEET 9.4
Unique Method of Burrowing in
Linguliform Inarticulates
Glottidia, a genus, furnishes the example
The animal enters the sediment anterior end first,
pedicle trailing behind. It moves downwards
through rotary movements of the valves through
sediments. Sediments are made softer by water
ejected from the body through periodic closing of
valves. Some lateral setae throw sand, made sticky,
upwards. After some length of the burrow, the
animal makes a u-turn and burrows upwards. It
then assumes the right position, with anterior
opening towards the water and pedicle directed
inwards into the burrow.
Quasi-infaunal habit has no recent example;
fossils like some productids were adapted to this
habit (see section 9.9).
Some small brachiopods are interstitial that
live in marine sands.
9.10.2 Classification
Schemes of classification of different groups of
organisms often require drastic changes on account
of newer facts or newer understanding. This is
well-exemplified with brachiopods. As mentioned
in section 9.8, at one stage punctation was
considered very vital for the group and, thus, was
given almost exclusive importance. But later that
attempt was revised for reasons discussed in the
said section. At that time, Muir-Wood (1955)
emphasized on a scheme that took account of a set
of characters, not just one. He also suggested
building up a scheme from generic level to higher
units. Subsequently, new significant finds from
Kirghizstan and Antarctica on early brachiopods
have added important data. Besides, Williams
et al., 1996 provided a computer-based cladistic
analysis that laid a foundation consistent with the
stratigraphic records (Clarkson 1998). Present
system, highlighted in this book too, is based on
all these changed opinions. It includes three
subphyla: Linguliformea, Craniiformea and
Rhynchonelliformea. The first has two classes, one
of them with four orders and the other with one.
The second subphylum has three orders. These two
were earlier included in Class Inarticulata.
Rhynchonelliformea, equivalent to the earlier
Articulata, is by far the most abundant and varied.
It includes five classes, three of which have only
one order each. Strophomenata is an extinct class
of Middle Cambrian to Triassic age with two orders
including strophomenids and productids. The fifth
class Rhynchonellata ranges from Cambrian to
Recent and includes nine orders, viz. orthids,
pentamerids, spiriferids, atrypids, athyridids,
terebratulids, etc.
Majority of brachiopod orders and families can
be identified on the basis of external morphology
alone, but to diagnose genus or species it may be
necessary to use features, such as lophophore,
muscle scars, hinge, cardinal processes. Problems
creep in as brachidium is not preserved in many
fossil groups.
To study these internal features, it may be
required to take help of special methods. When
the calcitic shell is preserved and cleaned from
the matrix, internal features may be studied from
valves separated from each other. In the case of
firmly closed shells, serial sections may be
grinded successively, with each successive ground
face photographed or drawn or otherwise
replicated, before regrinding. Wax models or
computer modelling arrived from these serial
sections may provide the entire internal structure
reconstructed.
On application of different modern concepts
and methods developed in recent years, we may
mention one such example on brachiopods. It
relates to bringing out examples of punctuational
model of speciation on the basis of species of
Cryptorhynchia and Kutchithyris from Kachchh
(Mukherjee, Bardhan and Ghosh 2002, Mukherjee,
et al., 2003)

9.10.3 Affinity and brief history
Some authorities believe brachiopods to be
polyphyletic, a ‘clade of organization’ which
includes other lophophore-bearing animals,
namely, bryozoans and phoronids. Even the
chitinophosphatic shell composition of linguliform
inarticulates is considered by some authors to
indicate closer relationship of these brachiopods
to bryozoans and phoronids, than to other
brachiopods (Clarkson 1998).
The earliest brachiopods are now said to occur
some distance above the base of Cambrian. But
even at that point, they are diverse. The first real
burst or radiation of the group may have taken place
Chapter 9 Brachiopoda 161
in early Ordovician to be followed by late
Ordovician glaciation and dwindling of brachiopod
fauna. Late Palaeozoic, i.e. Permo-carboniferous
witnessed another major radiation of the phylum
that produced well-known productids, spiriferids,
etc. including large aberrant coral-like
representatives of them. End-permian extinction
wiped out these groups. A few groups (such as
terebratulids and rhynchonellids) continued into
Mesozoic and Cenozoic, but they often assumed
importance. Paucity of post-Mesozoic brachiopods
is ascribed sometimes to the emergence and spread
of predatory starfish and carnivorous gastropods,
dreaded enemies of sessile epibenthics,
brachiopods and also bivalves.
 10
Bivalvia (Mollusca)
10.1 Introduction
The class Bivalvia (see Factsheet 8.1 for alternative
names of the class) that includes a kind of
multicellular, eukaryotic, invertebrate, aquatic
animals, is one of the most important divisions of
the phylum Mollusca. It ranges from Ordovician
to Recent. In course of its evolution, the group
adapted itself to different environments from
marine to freshwater, with varied mode of living
on or within the substrate (epi-/endobenthic) and
even in water (pelagic), and specializing in a kind
of suspension-feeding with the help of siphon.
As a result, at least at certain points in their history,
in some periods of the geological past as also in
the recent epoch, the group has proved itself as
one of the most successful animal-group in water,
judged from their abundance, variation and
importance. In the same vein, it is also considered
as one of the important groups in palaeontology.
10.2 Morphological Variation:
Adaptation, the Cause
Like brachiopods, bivalves secrete an external
bivalved shell, which is though equivalved and
inequilateral (for details of differences see Chapter
162
8 and discussions to follow). As in the case of
brachiopods, in bivalves too, shape and size
(surface area) of the shell and some other characters
depend on volume and surface area of the body,
relative growth of the body and the shell. So,
variation of shape and external morphology of
bivalve shells are likely to fall within a short range,
just as it was with brachiopods. In fact, Raup’s
theoretical study of shell forms (mentioned in
section 8.3) reveal that in nature, bivalves assume
only a small section of theoretically possible
shapes. But different kinds of adaptation has added
more variation in bivalve shells, as compared to
variation in brachiopods. So, bivalve morphology
is better appreciated when judged on the anvil of
habits-habitats and adaptations of these animals.
10.3 Shape, Dimensions and
Orientation
Since brachiopods and bivalves both have bivalved
shells their morphology has many common aspects
and, thus, can be discussed in the same frame.
The two groups differ primarily in their mode
of living. Whereas most brachiopods are sessile,
and epibenthic, the majority of bivalves are
vagrant, endobenthic organisms. A bivalve thus has

locomotion during lifetime, which means anterior-
posterior direction makes sense in it. It moves
forward to make burrow, hence the dead end of
the burrow is to its front; the open end towards
sediment-water interface lying to the back or
posterior of the animal. Secondly, bivalves have
to make burrows in soft, viscous sediments, for
which purpose they have to bear some such
streamlined body and shell that may favour smooth
passage through a fluid medium in an erect position
of the body, dorsal side up and ventral downwards.
Within a burrow, with the front and back as defined
earlier, the animal faces the same environment to
its two sides, right and left. Hence, the body and
shell each maintains a broad bilateral symmetry
with respect to these two sides and along the
anterior-posterior direction; the valves placed as
right and left valves on two sides of the symmetry
plane. This makes bivalve shells equivalved
(on right and left), inequilateral (antero-
posteriorly), whereas brachiopod shells are
inequivalved (larger, pedicle and smaller, brachial),
equilateral (also see section 8.5).
Brachiopod shells open towards anterior,
hence the hinge lies at or near the posterior end,
the symmetry plane placed at right angles to it. In
the erect position of bivalves within the burrow,
the hinge must lie on the dorsal side in the
symmetry plane itself, the shell opening towards
the lower, ventral side.
The shape of the shell or valves of bivalves is
determined by geometry of the commissure, as it
is with brachiopods. It may, thus, be referred as
circular, ellitical, trapezium, rectangular, oval, etc.
But, since the two valves are similar, the lateral
profile is always biconvex and, thus, largely non-
diagnostic. Length, height and thickness of the
shell are designated as shown in Figure 10.1.
10.4 Umbo and Beak
As parts of shells, the beak (the sharp apical part
of valve that is also the earliest formed part) and
umbo (abruptly elevated region on the shell
Chapter 10 Bivalvia (Mollusca) 163
surface around beak) refer to the same structures
as those of brachiopods. But, since the two valves
are similar in bivalves, their umbones are similarly
curved or coiled. As discussed, in brachiopods,
this is not the case and the two umbones may be
widely different there. Besides, in bivalves there
cannot be any case of umbo of one valve
overlapping that of another. They lie on two sides
of the symmetry plane, adjacent or not adjacent
and are never overlapping. This is because there
is no non-strophic type of hinge to be found in
bivalves. Moreover, in brachiopods umbones are
orthogonal to the hinge line, whereas in bivalves
they are inclined to the hinge. Most commonly
they are inclined towards anterior (called
prosogyral), as more shell material is added to
the commissure on its posterior side. In a few
genera such as Trigonia, Glycimeris, etc.
umbones are coiled towards posterior
(opisthogyral), though in these cases too the shell
material is added more posteriorly on the
commissure. In some other genera, e.g. Pecten,
Spondylus the valves are equilateral with the shell
material added equally on the anterior and
posterior sides of the commissure and, hence, the
umbones are coiled orthogonally (orthogyral).
Coiling of umbo, thus, appears to be a feature
controlled by particularities of growth.
10.5 Hinge and Dentition
Having a bivalved shell, bivalves require and, thus,
have hinge in their shells, placed at or near the
dorsal margin (in contrast to posterior margin in
brachiopods), along which the two valves are more
permanently attached to each other during lifetime
and about which the shell opens or closes.
Whereas hinge is present only in articulate
brachiopods, it is there in most bivalves, weak or
strong though it may be.
Besides, in brachiopod hinge teeth occur on
the pedicle valve and socket on the brachial valve.
In bivalves, however, the same valve may contain
teeth and sockets alternately placed; the other
164 Part Two: Major Invertebrate Groups

1 10 (a) Internal view of a left valve

2 (anterior to the right,)
3 (b) External view of the same,
4 (c) Dorsal view of the closed shell
11 showing equivalved nature,
(a) 5 (d) Ventral view with the valves kept
(b)
slightly open,
7
6 (e) Posterior view of a brachiopod
12 shell showing inequivalved nature
(compare with (c)),
(f) Lateral profile of the same
(c)
(d) brachiopod shell,
8 (g) Adductor muscle and ligament in
9
a closed shell,
(h) External ligament,
(e) (f) (i) Internal ligament ((g) to (i) are
sections through beaks),
(j) Sessile epibenthic (as in rudistids),
13 (k) Temporary attached,
13 (l) Nektic,
(g) (m) Partially endobenthic,
(h) (n) Sessile epibenthic (as in oyster
(i) banks),
(o) Byssally attached,
(j) (p) Shallow burrower,
(q) Deep burrower,
Number index:
(l) (k)
(o) 1 Beak
(m) 2 Cardinal teeth/socket;
(n)
3 Lateral teeth;
4 Adductor scars;
(p) 5 Pallial line;
(p)
(q) 6 Pallial sinus;
7 Concentric growth rings;
8 Lunule;
(q) 9 Escutcheon;
10 Length;
11 Height;
12 Dentition;
13 Ligament;
Fig. 10.1 Bivalve features: morphology-ecology.
valve will then contain corresponding sockets and which is not symmetrical about the symmetry
teeth respectively, to fit into those of the former plane of the animal. Dentition is considerably
valve. In fact, this teeth and socket system, called varied in bivalvia and is characteristic at different
dentition, is the only structure in bivalve shells taxonomic levels.

Bivalve hinge line, as in brachiopod, may be
straight or curved. But there is no hinge type in
bivalves comparable to the non-strophic hinge of
brachiopods that act on two fulcra. Even in bivalves
with a curved hinge, the valves are attached along
the whole of the hinge line.
10.6 Hinge Plate and Hinge Area
Excepting in a few genera or species (e.g. Pecten,
Mytilus, Ostrea, etc.), there is an essentially flat
triangular area between the beak and the hinge line.
This part of the valve, called hinge plate, lies,
during lifetime, either vertically on the symmetry
plane or is inclined towards the latter in the
opposite direction from that of the main valve. In
the first case, the beak, the hinge plate and the
curved hinge line all lie on the symmetry plane.
The two hinge plates of the right and left valves,
respectively, are thus in contact with each other.
On the other hand, when the hinge plate is inclined,
the two such plates of the two valves, meet each
other along a straight line that itself acts as the
hinge line. The outer triangular surface between
the beak and the hinge line is then called hinge
area (equivalent to brachiopod interarea).
Dentition of bivalves occur on the hinge plate.
For curved-hinge forms where the hinge plates of
the two valves are vertical, teeth and sockets occur
on the plate surface. For forms with straight hinge
line and hinge area, they are cut on the thickness
of the hinge plate, as shown in Figure 10.2 .
A few genera do not have any hinge plate; in
Pecten the hinge line is straight; teeth and sockets
occur as tiny thickenings or groovings on the valve
interior on two sides of the beak; in Mytilus, too,
the hinge line is straight, but teeth are small spine-
like projections below the beak (sockets are
corresponding depressions). In genera like Ostrea,
Alectryonia, etc. where the shell is foliaceous and
hence thick, the hinge area is a simple depression
below beak ending in a curved hinge line. These
genera are also teethless or edentate, in which case
Chapter 10 Bivalvia (Mollusca) 165
valves are held together only by stronger muscles
and ligament.
Hinge line and hinge area in brachiopods and
bivalves, or even within the bivalves themselves,
are thus only analogous structures, morphological
features related to the growth of valves.
10.7 Variations in Dentition
As mentioned, dentition of bivalves, i.e. teeth and
sockets occur on the vertical portion of the hinge
plates of the two valves, which are in contact with
each other when the shell is closed. In curved-hinge
forms, it is the triangular surface of the hinge plate
of each valve between the respective beak and
hinge line, whereas in forms with straight hinge
line and hinge area, it is the thickness of the hinge
plate, as shown in Figure 10.3.
As in strophic brachiopods, in bivalve shells
with straight hinge line and hinge area, teeth and
sockets are numerous, yet small, broadly
undifferentiated in shape and size, and located all
along the hinge line at right angles to the latter.
Such a dentition is effective, so long as the hinge
operates about the whole length of the hinge line.
With curved hinge line, the effective length is
smaller and that demands fewer but stronger teeth.
To accommodate them in the space available on
the hinge plate, they are shorter and stouter below
the beak, radiating downwards from the latter,
while on the lateral sides they become slender and
longer running parallel to the margin. In effect,
teeth and sockets are, thus, differentiated into two
types, cardinal, below the beaks and laterals,
parallel to the margin.
Studies on dentition consider undifferentiated
type as more primitive, while the differentiated
teeth and sockets evolved subsequently. Between
the two there are varied combinations which differ
in number of teeth and sockets, their shape and
size as well as position in respect to beak, hinge
line and margin. These types are defined below.
Before that, it may be added that over and above
166 Part Two: Major Invertebrate Groups

(i) (i)
(i)

(b) (ii)
(a) (ii)

(c) (ii)

(d)

(f)
(e)

(g)
(h)

(i)
Fig. 10.2 Bivalve shells: shape and ornament.
(a) Arca; (b) Trigonia; (c) Spondylus; (d) Venus; (e) Hippurites; (f) Glycimeris; (g) Mytilus; (h) Mya;
(i) Alectryonia. a(i), b(i) and c(i) are external view, whereas a(ii), b(ii), c(ii), (d), (e), (f), (g), (i) are
internal view.
Note: Dimyarian shells in (a), (b), (c), (d), (f), (g), (h) Monomyrian shells in (i)
Taxodont dentition in (a), (f) Pallial sinus in (d), (h)
Heterodont dentition in (d) Strongly inequivalved shell in (e)
Schizodont dentition in (b) Inequivalved shell in (i)
Isodont dentition in (c) Strongly inequilateral shell in (g)
 Chapter 10 Bivalvia (Mollusca) 167

(a) (b) (c)

(d) (e)

(g) (h) (f)

(i) (j) (k)

(i) (ii) (m)

(l)

Fig. 10.3 Bivalve features: valve interior and dentition; shape and ornaments.
(a) Heterodont; (b) and (e) Taxodont; (c) Desmodont; (d) Schizodont; (f) Edentate; (g) and (i) alate
shells; (h) Dysodont; (i) and (l) Shell with radial ornaments dominant; (j) Closed equivalved shell
with short heart-shaped lunule to the anterior of beak and long escutcheon to the posterior; (k) A
single right valve with concentric growth rings on the surface; (l) Closed inequivalved shell, the left
valve in front and the right valve, respectively; (m) razor-shaped deep burrowing form. (scale in mm).
168 Part Two: Major Invertebrate Groups
the general characteristics of dentition, as indicated
above, there are two more fundamentally
significant dentition types. One has already been
mentioned, i.e. edentate that characterizes a few
genera and species. The other type of dentiton,
called pachydont, has few but strong rough or
rugose, large teeth or sockets. Both these strikingly
different types are found in such bivalves which
are sessile and epibenthic and hence are
inequivalved, may be equilateral-like brachiopods
or cylindroconical-like corals. In the former, the
oysters or genera such as Ostrea, Alectryonia,
Gryphea, Exogyra, etc., strong muscle and
ligament help articulation of edentate shells. In the
second case of rudistid bivalves (e.g. Hippurites
or Radiolites, which formed prominent reef-like
structures in Cretaceous) the sturdy pachydont
dentition provided strong articulation.
The major types of bivalve dentition (Figures
10.1, 10.2, 10.3) are as follows:
1. Taxodont: This type of dentition is
characterized by numerous, yet small teeth and
sockets, broadly undifferentiated in shape and
size, and located all along the hinge line
generally and nearly at right angles (sometimes
radially) to the latter. Genera such as Arca and
Glycimeris, etc. have this kind of dentition.
Taxodont dentition of the genus Nucula is
another variation. Besides, some genera of
Ordovician age have a palaeotaxodont type of
dentition, in which anterior teeth are larger and
raised. May be, they helped protect the fleshy
foot-like process that emerged towards front
for the purpose of burrowing.
2. Dysodont: It is a rather weakly developed
type of dentition that has very few, small and
weak teeth or sockets; they are relatively
simple spine-like as in Mytilus or are formed
due to low thickening or folding of shell
material as in Pecten.
3. Isodont: This type is characterized by two
teeth or sockets, generally large, similar in
shape and size, strong and placed on two sides
of a central ligamental pit (explained later),
for example, in Spondylus.
4. Schizodont: This type also has large and
rough teeth, as in Trigonia, Unio, etc.
5. Heterodont: Majority of Tertiary and recent
genera, but even some ancient Ordovician
bivalves are heterodont, in which dentition is
the most differentiated into cardinals below the
beak with or without laterals anterior and
posterior to the former (e.g. Cyrena with
laterals; Venus without laterals). Palaeo-
heterodont or actinodont are varied precursors
of this type of dentition.
Heterodont dentition is described in a system
introduced by Bernard and Munier Chalmas.
In it, major teeth are marked with numbers and
sockets with dash ‘—’ , starting from centre
and increasing sideways with odd numbers
used for right valve and even ones for left
valve. Anterior and posterior are distinguished
by marking the numbers with ‘a’ and ‘b’.
6. Pachydont: Made of very large, heavy and
blunt teeth, this type of dentition is found in
rudists, which are sessile and attached to hard
substrate (e.g. Hippurites, Radiolites).
7. Desmodont: This type of dentition has very
small teeth or none at all. A kind of ridge at
the end of hinge line may serve for teeth
(e.g. Crassatella, Mya). In Mya, there is a
process lying below the beak and consisting
of a spoon-shaped structure on left valve and
an inverted bowl-shaped depression just below
the beak on right valve for internal ligament.
(Figure 10.2.H). This is known as
chondrophore.
10.8 Adductor Muscles and
Ligament
10.8.1 Opening and closing of shell
Though ligament occurs in bivalves near the hinge
line or umbo and, thus, had to be referred in

connection with dentition, it is better compre-
hended when judged along with adductor
muscles. While the latter help the shell to close,
ligament acts in opening it. Both of these belong
to the non-mineralized, i.e. soft parts of the body,
which leave their marks on the shell.
Opening and closing of bivalve shells take
place in the following way. About midway between
the dorsal margin or hinge and the ventral margin,
there lie the adductor muscles running across the
symmetry plane and attached to the two valves.
As natural response, the muscles contract to pull
the two valves near each other. The shell is closed
by that. On the other hand, ligament lies between
the two umbones at the dorsal margin. The more
the shell is closed, the wider the two umbones move
from each other. It causes ligament to stretch and
thus to develop its tendency to contract and release
the stress created by stretching. As the ligament
regains its position, the umbones come closer,
while the valves move apart at the ventral margin,
thus opening the shell. Adductor muscles are then
stretched, tending to contract again to its
equilibrium. When that happens, the whole process
is repeated.
This process explains why bivalve shells are
seldom found in closed condition and why shells
of recently dead bivalve animals in which fleshy
parts have already decayed and been lost, are found
gaping, being held still together along the hinge.
As adductor muscles are lost and with that, also
the pull on the valves to close the shell, ligament
throws the valve open. Organic material of the
ligament takes more time to decay. Till then the
ligament, in association with the hinge and its
dentition, holds the valves together. With time,
ligament dries and becomes brittle. Any small
pressure then splits the ligament, whereby the
valves fall apart.
10.8.2 Variations in adductor muscles
Valves are inequilateral in bivalves. They are
generally more elongated along the anterior-
posteriorly lying symmetry plane. Hence, to close
Chapter 10 Bivalvia (Mollusca) 169
such a shell most efficiently, a system of paired
adductor muscles, one at the anterior end and the
other at the posterior, is best suited. In additon, in
majority of bivalves, the posterior portion of the
body and shell are bigger than the anterior
counterparts. Hence, the posterior muscle requires
to pull stronger and so must be bigger. The more
the beak is placed near the anterior end of the hinge
line, the smaller and less important becomes the
anterior muscle in such shells. On the other hand,
as the shell becomes equilateral in sessile forms
like Ostrea, Pecten, one single adductor muscle
placed near the centre becomes sufficient and
efficient.
Muscles are lost in fossils. Their scars remain
on the inner side of the valves. Accordingly as the
muscles are two or more unequal or equal, shells
may be :
1. Monomyarian, with a single scar at the centre
or slightly posterior (e.g. Ostrea, Pecten); and
2. Dimyarian, with two adductor scars, one
anterior the other posterior.
The latter has two types:
(a) Isomyarian with two scars similar, found
generally in slightly inequilateral shells
(e.g. Cyrena), or
(b) Anisomyarian with two scars dissimilar in
size as well as shape, the posterior one
being bigger (e.g. Mya, Mytilus).
10.8.3 Variations in ligament
Ligament which lies at the dorsal margin of the
shell between or near the two umbones, shows
some variation. Fossils do not have ligament, but
features which held the ligament are preserved and
are often characteristic of genera.
In majority of bivalves, the ligament is external
to the shell and extends both to the anterior and to
the posterior of the beaks. With posterior portion
of the shell being larger in most bivalves, ligament,
too, has larger share posterior to beak (called
opisthodetic; e.g. in Cyrena, Mytilus). In some
genera, with larger anterior part of shell, ligament
170 Part Two: Major Invertebrate Groups
is also placed dominantly in front of beaks
(prosodetic; e.g. in Nucula). However, in many
genera, ligament lies internally in the shell, below
the beaks (amphidetic; e.g. Pecten, Spondylus, Mya).
Such ligament is also different in their working.
When the shell is closed, this kind of ligament
(called resilium) is pressed between the two
umbones. To release this pressure, it needs to
expand. As it does, it pushes the two umbones
apart. This makes the shell open at the ventral
margin, putting the adductor muscle under stretch.
The latter contracts and closes the shell, thereby
subjecting the internal ligament again under
compression. Generally, resilium lies in a triangular
pit, shallow in Pecten, deep in Spondylus. The pit
is referred as resilifer. In genus Mya or other
desmodonts, it may lie in a specialized structure
called chondrophore (see Figures 10.1, 10.2, 10.3).
10.8.4 Pallial line and sinus
Mineral matter of bivalve shells is secreted by the
mantle or pallium of the body. But at any stage of
growth the shell is a little bigger than the mantle
area. The mantle is attached to the shell along a
line slightly inside of the ventral and lateral
margins. This line is called the pallial line. It runs
broadly parallel to the margin from one adductor
muscle scar to another in dimyarian forms. In
monomyarian shells, this line lies ventral to the
adductor scar. A portion of the mantle extends
ventrally beyond this line; but it is no longer
attached to the shell. Rather it hangs freely between
the two valves.
When the pallial line is a continuous curved
line from one scar to another, it is referred as entire
and the shell then is integripalliate. But in many
genera, this line has a break near the posterior
adductor scar, where it swerves inwards and then
marking an indentation, curves back to the original
course. The indentation is called pallial sinus and
the shell sinupalliate. The sinus may be weak or
strong, sharp, angular (as in Venus) or broad,
rounded (as in Mya) Figures 10.2 D, H in all cases
being characteristic of genera; but whatever be the
shape or prominence, it is invariably placed
towards the posterior scar. This characteristic
position is explained when we look at what the
sinus is formed for.
10.8.5 Significance of pallial sinus and
pedal scar
It has already been said that bivalves are
dominantly vagrant, endobenthic that make
burrows in soft sediments. For this the animal
requires two vital actions: first, making the burrow,
and second, acquiring oxygen and nutrients from
water behind it, when it is inside the burrow. To
make burrow, it moves forward; so it then needs
an organ or appendage near its anterior margin that
can remove the sediments lying in front of it. In
reality there is an axe-like fleshy process in the
animal that acts as a foot (hence, the name
pelecypoda: pelecys, axe). The animal presses upon
it to move forward and also can use it to remove
sediment in front of it. Sometimes, this foot leaves
a distinct scar on the internal surface of the shell,
immediately below the anterior adductor scar. It is
known as pedal scar and is quite discernible in
genera like Unio.
On the other hand, while within the burrow,
the animal has the sediment-water interface and,
thus, water itself at its back. It thus demands an
organ meant for drawing in water. This is a tubular
process, which serves the purpose. It draws in fresh
water for its feeding and respiration, as well as
throws out the foul water after use. It also keeps
the burrow clean of pollution. In some genera this
siphon system is more developed; there are two
tubes, one inhalant and the other exhalant. Pallial
sinus marks that part of the body, where the siphon
is pushed into the mantle. Naturally, it cannot but
occur in the posterior part of the pallial line or shell.
Some deep-burrowing genera such as Mya or
Mactra, etc. have a big siphon. To accommodate
it, their shells remain gaping at the posterior end,
when they are closed.
 Chapter 10 Bivalvia (Mollusca) 171

10.8.6 Surface ornaments classification of bivalves based on adaptation.

Like brachiopods, bivalve shells also have mainly
two types of ornaments: radial and concentric
(Figures 10.1, 10.2, 10.3). But there is no median,
radial ornament, as the bilateral symmetry plane
does not run across the valves; it is between the
two valves (see Factsheet 10.1).
Though both the groups of animals are
suspension-feeders or filter-feeders, they differ in
the mode of living. It will be apparent from the
discussion above (section 10.8.5) that bivalves do
not require their radial ornaments for drawing in
of water. However, as in brachiopods, in bivalves
too, different types of ornaments develop from
differential rates of growth of the body and its shell.
A few ornaments like spines are formed as typical
adaptive developments.
10.9 Bivalve Adaptation:
Functional Morphology of
Shells
Varied adaptation of bivalves was known since
long, as is indicated from the early scheme of
Functional morphological analyses of these
adaptations are more recent additions. Ideas have
developed, even changed in course of addition of
more data from newer and newer findings
particularly based on observations of recent
bivalves and their mode of living. At one time, the
siphon was interpreted as an organ used for a
backward propulsion of jets of water to add a
forward thrust to the animal swimming in water.
But observations that most bivalves were
endobenthic and used their siphon for suspension
feeding, completely changed the idea.
Functional significance of morphological
features such as adductor muscle scars, pallial
sinus, ligament, etc. has already been pointed out
in the above sections. A few other necessary
discussions will be made in this section.
Some of the features of bivalve shells such as
hinge, dentition, mineralogy and microstructure of
shells, etc. appear to be less influenced by
adaptation. They are, thus, used more for syste-
matics and classification.
On the other hand, shell forms, adductor scars,
pallial line and sinus, ornaments, etc. are affected

FACTSHEET 10.1
Bivalve Ornaments: Common Examples

Ornaments Characteristics Examples Probable significance

Only concentric Generally fine grooves Cyrena, Venus Growth at different stages of
ontogeny
Concentric in distinct As above; foliations Ostrea Sessile, benthic, cemented
foliaceous shell distinct
Only concentric Coarse and prominent Mytilus Secretion increases at certain
ornaments in thin shell stages for reasons unknown
Dominantly radial Radial ornaments more Arca Shell secretion overcomes
prominent growth of body
Radial and concentric Radial or concentric Glycimeris Shell secretion and growth of
equally prominent none prominent body at par
Concentric in anterior part, Both prominent Trigonia May be associated with
radial in posterior with/ burrowing habit
without a rib in between
172 Part Two: Major Invertebrate Groups

by habits and habitats. They are better suited for
functional morphological analysis.
10.9.1 Recent eco-morphotypes:
Endobenthic types
Present-day bivalves show a few major eco-
morphotypes, each adapted to some particular mode
of living and, thus, acquiring certain morphological
characteristics (Clarkson 1998). Factsheet 10.2 and
Figure 10.1 show them. They clearly point out a
varied adaptation for bivalves. It includes both
epibenthics and endobenthics as also some pelagics,
viz. nektics; benthics prefer different types of
substrates, hard rocky or soft, sediment-laden; there
are sessile or vagrant or free-lying, cemented or
byssate bivalves, as also burrowers, borers or
cavity-dwellers. For most of these types we can
find, either quite a few examples or persistent
occurrences, both suggesting successful adaptation
of these animals to their respective mode of living.
However, the most general, abundant and, hence,
the most successful adaptation is the endobenthic,
burrowing mode of living.
Though varied in adaptation, bivalve animals
are suspension-feeders or filter-feeders. A few
genera (e.g. some genera of Nuculacea) are
deposit-feeders. Morphological variations in
bivalve shells depend on these two factors: mode
of living and feeding habit.
Thus, shallow burrowing forms have
equivalved shells, with a circular or trigonal
commissure whose length and height are nearly
equal, anterior-posterior axis is virtually parallel
to the hinge axis and at right angles to the ventral-
dorsal axis; these are generally isomyarian and
integripalliate; on surface they may bear typical
ornaments, sometimes different in anterior and
posterior sides. Shells may be gaping at either or
both ends, for foot and siphon, respectively. Deep
burrowing forms are, on the other hand, razor
shaped or tubular with an elongated commissure
in which the length is much greater than the height;
the three axes, viz. anterior-posterior, hinge and
dorsal-ventral are, however, similarly disposed as
they are in shallow burrowing forms. But siphon
in deep-borrowers are generally longer and, thus,
leave a more distinct and larger pallial sinus. These

FACTSHEET 10.2
Bivalve Mode of Living

ENDOBENTHIC Burrowing Shallow burrowing Cerastoderma, Venus, Tellina, Trigonia, etc.

Deep burrowing Mya, Ensis, Solen, Phacoides, etc.

Boring Pholas, Teredo, Lithophaga, etc.

Nestling Hiatella, etc.
EPIBENTHIC Sessile byssate Mytilus, Modiolus Pteria, Lima, etc.
Byssate, temporary Lima,Pecten, Arca(?), etc.
swimming
Sessile cemented Attached by Ostrea, Gryphaea, Alectryonia, etc.
left valve
Attached by Spondylus, Plicatula, Hippurites,
right valve Hippuritella, Radiolites, etc.

NEKTIC/ Free-lying Gryphaea, Pecten, etc.

NEKTOPLANKTIC Swimming Posidonia, etc.

forms are normally strongly asymmetrical and
anisomyarian. Since they require little movement
inside the burrow, they tend to have weak dentition,
internal ligament, often placed in chondrophores;
but the shell is tough and sharp to assist burrowing
and the shells are gaping at both ends. However,
Phacoides, a deep burrower, has a circular
commissure like shallow burrowing genera.
Borers, too, have thin, tough and sharp shells that
can cut through rocks, wood or such other hard
substances. Sometimes, as in Pholas, the shell has
spines or thickened ends at the anterior. There are
genera which are nestling, that is, which live in
existing holes in the substrate.
10.9.2 Epibenthic types
Epibenthic bivalves are majorily sessile and are,
thus, quite similar to brachiopods in some respects.
Shells are inequivalved and less inequilateral. In
one type, the lower valve, also the larger, is
cemented to the substratum; in a few others, spines
serve for anchoring. In some genera, the animal
has a feature that helps in anchoring, as pedicle
does in brachiopods. Such forms are, however,
equivalved. In these forms, the foot-like process
of the body secretes a bundle of fibrous colagen-
protein. The bundle is known as byssus. It forms
as a sticky mass, but soon it becomes solid and
holds the shell attached to the bottom. Byssus
comes out of the shell through a small or large
indentation (accordingly called a byssal notch or
sinus) on the lateral margin of the shell. Since
byssus itself is secreted by the foot, it cannot but
occur on the anterior side. These shells often have
a narrow or acute anterior end, making them
strongly asymmetrical, with an elongated triangular
commissure and highly unequal adductor scars, the
anterior one being very small. Anterior-posterior
axis is at a low angle with the hinge axis. The
sagittal plane between the two valves, that contains
the commissure, is kept vertical during lifetime
with the ventral part of the shell flattened to hold
the shell upright, maintaining bilateral symmetry
about the sagittal plane. In a few genera
Chapter 10 Bivalvia (Mollusca) 173
(e.g. Pteria), an ear-like prolongation at the
posterior end of the hinge line controls orientation
of the sagittal plane.
In cemented or otherwise fixed sessile forms
and in free-lying genera, the sagittal plane lies in a
horizontal position. The valve which lies below
this plane is attached to the bottom, is generally
distinctly larger and thicker, with the animal
residing in it. The upper and smaller valve acts as
a lid. The shell is monomyarian and the adductor
is generally prominent and central in position. In
respect to bivalve body, the lower valve may be
either the right or the left valve.
Ostrea is an important example of the latter,
i.e. left valve attached. The genus has no dentition,
but strong ligament and adductor muscle instead.
As the shell grows attached to the substratum, it
grows in layers, making the shell foliaceous and
its shape often conforming to the irregularities of
the base. Sometimes, the surface has well-formed
radial ribs, obviously to accommodate more shell
material than the surface area of the body, and
having some bearing with intaking of water as in
brachiopods.
Genera with right valve attached may be
grouped into two types themselves. In the first, the
lower valve is attached with the help of spines, as
in Spondylus. The shell is inequivalved,
monomyarian and isodont. In the second type
found in the extinct family Rudistidae, the lower
valve grows vertically and, thus, becomes coral-
like, tubular or horn-shaped. Hippurites is a typical
example. However, in Chama, another genus of
the same family, the shell is rather bowl-shaped.
In both the cases, the shell is thick, rough on
surface, monomyarian and characterized by
pachydont dentition.
Pecten, Gryphaea, etc. are genera that lie freely
on the substrate. Of these, the latter is attached when
young; on maturity it becomes free-lying. Pecten, on
the other hand, can make a sort of saltatory movement,
when attacked; it flaps the two valves forcefully to
swim out of danger. These genera are also
inequivalved, monomyarian and weak in dentition.
174 Part Two: Major Invertebrate Groups

Pecten or Lima also have a byssus each. Lima lives
more attached to the bottom, than swimming.
Posidonia is a genus that is typically nektic.
Factsheet 10.2 summarizes the list of bivalve
ecomorphotypes.
10.9.3 Evolution of adaptation
Different groups of bivalves have typical
adaptations. At the same time, adaptations change
or evolve with time too. A case in point is observed
in marine mussels of the family Mytilidae. These
are sessile byssate forms, dominantly epibyssate.
They attach to hard substrate, but some form
clumps or banks of soft bottoms, being attached to
debris of dead shells or even to one another
themselves. Some mytilid species are also
endobyssate, living partly or entirely buried in soft
sediments, attaching to large sedimentary particles
or such other objects available. Epibyssate forms
have a flat ventral part of the shell to keep the shell
stable; in endobyssate forms the ventral portion is
sharp, and like the hull of a boat. There are other
morphological differences too. From anatomical
and palaeoecological studies, it is inferred that
endobenthic habit is more primitive condition from
which the epibenthic condition are derived during
the course of evolution.
Thus, endobyssate species (of Modiolus) were
more typical of Palaeozoic, whereas epibyssate
species appeared later (e.g. Mytilus). Factsheet 10.3
shows the morphological differences between
endobyssate and epibyssate forms.
Adaptations of bivalves discussed above, are
known from living genera and species.
Conclusions on the mode of living and feeding
habit of ancient organisms may be drawn from
fossils, comparing their morphology with those
of known extant equivalents. However, in
palaeoecological studies of bivalves, such
functional morphological analyses must be
augmented with other evidences viz. association,
covering sediments, etc.
10.10 Some Other Aspects
A few other aspects are relevant in a discussion on
bivalve morphology. One of them is the gills.
Though they are not preserved in fossils, gills are
considered important in bivalve classification.
In fact, the name Lamellibranchia for the class
suggests the importance attached to these. There
are four types, the two major types are Eulame-
llibranch and Filibranch; Protobranch is the
primitive and less developed type, and Septibranch
is a specialized development associated with
adaptation to boring in hard rocks or wood.
Two other features to consider are the shell
composition and microstructure. They include
mineralogical composition of shell, its overall
organization, crystal habit and texture, etc. Six
different combinations are observed, all under
microscope (see Factsheet 10.4). The types
appeared at different phylogenetic stages of the
group. At the same time, at least in one case or

FACTSHEET 10.3
Evolution of Adaptation, Case Study from Mytilids

Criterion Venter Posterior byssal Age

retractor muscle
Endobyssate Like hull of boat ..pulls obliquely downward; Since Middle Palaeozoic
anchors shell upright
Epibyssate Flattened for stability ..pulls at right angles to venter; Since Mesozoic
holds shell firm against substratum
 Chapter 10 Bivalvia (Mollusca) 175

two, the shell microstructure seems to be related
to the mode of living. For example, tough and
hard crossed lamellar type (see Factsheet 10.4) is
found more in deep burrowing or boring forms.
Shell mineralogy of bivalves has another
importance to palaeontologists. The most imporant
and dominant constituent of bivalve shells is
aragonite. Since aragonite is metastable, it is
unlikely to be preserved in ancient (Palaeozoic or
early Mesozoic age) bivalve fossils; it is lost either
by dissolution or by replacement to calcite. In the
first case, the fossil is preserved as a mould,
external or internal. In the second case, the body
fossil may lose finer details of shell morphology
during replacement of aragonite to calcite. This
may also be a reason for relative paucity of bivalve
fossils in Palaeozoic, as compared to fossils of
articulate brachiopods of the same age with calcitic
hard parts. Relative dominance of Palaeozoic
brachiopods is explained as indicative of better
adaptation and, thus, successful radiation of the
latter. But simple taphonomic advantage of that
group may have acted as a major cause. It requires
more study to settle the point.

FACTSHEET 10.4
Primary Shell Microstructure Types in Bivalves

Simple prismatic: Columnar polygonal calcite or aragonite prisms

Composite prismatic: Tiny radiating acicular crystals
Nacreous: Tabular sheets of aragonite resembling a brick wall (in cross-section), usually found in
middle and inner shell layers
Foliated: Lath-like calcite crystals arranged in sheets
Crossed-lamellar: Normally aragonitic, made of closely spaced lamellae, each made of thin stacked plates
of aragonite, those of adjacent lamellae being inclined in opposite directions to one
another. (There is a complex crossed lamellar variety, which has intergrowths of
blocks of crystals set in four principal orientations.)
Homogeneous: With small granular anhedral crystals
 11
Gastropoda (Mollusca)
11.1 Introduction
Gastropoda (the animal uses gastric mass, the
fleshy body itself as pod or foot; hence the name)
is another class of the phylum Mollusca, and
includes, like Bivalvia, a kind of multicellular,
eukaryotic, invertebrate, majorly aquatic animals.
There are, however, gastropods which are
terrestrial in habit, that is, can live on land and are,
thus, air-breathing. The class appeared slightly
earlier than the bivalves, in Lower Cambrian and
continues till date. In the long course of its
evolution, the group adapted itself to different
environments from marine to freshwater, as also
to land with accompanying morphological
variations. Like Bivalvia, the class Gastropoda also
had their widest and strongest adaptive radiation
in Cenozoic era. At the same time, in both the cases,
the shell is made mainly of aragonite. In result,
gastropod shells face every possibility of being lost
in fossils. Hence, the absence of gastropods in a
fossil record, may not necessarily indicate their
non-existence or extinction. It may simply be the
taphonomic disadvantage that might have stood in
the way of gastropods fossils being preserved in
sufficient number.
These similarities notwithstanding, the two
classes of animals have important differences
176
between them. Firstly, adaptive modifications in
gastropods are not as well brought out as they
are in the case of bivalves. Secondly, gastropod
shells are distinctly and totally coiled; only few
gastropod genera do not have coiled shells. In this
respect, gastropods are similar to cephalopods,
animals of a third major class of Mollusca. Study
and description of shell morphology of these two
groups, thus, need be done in a different way.
Brachiopods and bivalves have bivalved shells,
often with hinge. In gastropods and cephalopods,
shells are largely univalved, in which the second
component, the operculum is a small lid much
smaller than the main valve. There is no hinge
either in these shells. Moreover, the bivalved shell
of a brachiopod or bivalve, totally covers the body.
The opening in each valve along the commissure
(equivalent to aperture of gastropod or cephalopod
shell) is exactly covered by the second valve, so
that there is no final opening in the shell, when
closed. In gastropod or cephalopod shell, on the
other hand, the valve (or for that matter the shell)
is closed at one end and open at the other in an
aperture. It provides the passage through which
the animal comes out of the shell to perform its
vital activities or withdraws into the latter for
protection during lifetime. Thirdly, there being no
question of repeatedly opening or closing the shell,

musculature of gastropods or cephalopods is again
of a different kind. Surface features on these shells
are also aligned with coiling and are, thus, different
from surface features of brachiopod-bivalve shells.
For these reasons, the format for the study of
morphology is changed into the following:
1. Shape of shells, type of coiling, dimensions,
symmetry, orientation, etc.
2. Aperture and associated features
3. Internal structures
4. Surface features
11.2 Two Important Characteristics
of Body
A discussion on morphology of gastropods should
begin with a mention of a few characteristics of
their body. It has been indicated in Chapter 8 that
gastropod body undergoes a torsion in an early
ontogenic stage. On account of this, a part of the
body is twisted through 180° in respect to the
other. In basic molluscan organization, mouth and
anus are placed at the opposite ends of the body.
But the torsion in gastropod body brings them to
the same end, anus being placed above the mouth.
This creates problem in the functioning of both
the openings. Different groups of gastropods
develop different means to overcome this
difficulty, which brings in some corresponding
changes in shells too. Details of this will be
discussed later.
The second important characteristic is not
directly related to shell morphology, though it is
significant. Most of the gastropods have gills for
respiration. In one subclass, Pulmonata, however,
such gills are abolished. These animals have
numerous blood-circulating vessels in the mantle
wall, with the help of which the animals acquire
oxygen directly from water or air. In the first case,
they act like a gill, though different from the gills
which other aquatic gastropods normally have. In
pulmonates which take oxygen from air, the said
Chapter 11 Gastropoda (Mollusca) 177
process acts like a lung. Pulmonata is one of the
successful groups of gastropods in the recent
times. A majority of this group are terrestrial.
Obviously for these land-dwelling organisms,
lung-like respiratory apparatus that can acquire
oxygen from air, serves for a better adaptation. A
number of pulmonates are, however, water-
dwelling and they bear the gills referred above.
But these gills, though acting towards the same
end of respiration, are different in structure and
mode of functioning from the gills of other
gastropods. In evolutionary sequence, these
pulmonate gills represent an advanced type of
respiratory apparatus over the more primitive type
of normal gills of gastropods.
Apparently trivial, the phenomenon points to
an important aspect of evolution. Evolutionary
changes are characteristically irreversible. Thus,
an organism at an advanced stage of evolution
cannot acquire exactly the same conditions or
characters of more primitive or ancestral stages.
This is why the concept of degeneration is no
longer accepted in evolutionary studies. In this case
of gastropods, the gills of water-dwelling members
of the group evolved into a sequence of two
advanced stages: first, a lung-like process that
helped some pulmonates to successfully adapt to
land-dwelling, air-breathing habit. From them
evolved some other pulmonates which went back
to water-dwelling mode of living with the newly
developed process acting as an advanced gill.
The sequence, gills to lung and then to gills again,
does not represent any degeneration, for the
advanced gills are never the same primitive ones
as aquatic gastropods normally have; these are
completely different organs.
11.3 Morphology: Coiled Shells
The first morphological characteristics of gastropod
shells to note is the coiled nature of the shell. In
most gastropods, this coiling is conispiral (or
trochospiral) (e.g. in Turritella; see Figure 11.1).
178 Part Two: Major Invertebrate Groups

(c)

(a) (b)

(d) (e) (f)

Fig. 11.1 Gastropod coiling.

(a), (b) Trochospiral/Conispiral convolute with single whorl visible from outside; (a) Shell surface
partially broken to show the interior, Cypraea, (c) Planispiral convolute: Bellerophon, (d)
Pseudoplanispiral: Euomphalus, (e) Trochospiral with earlier whorls visible from outside surface with
a slit-band, aperture with a slit Pleurotomaria, (f) Aberrant non-coiled gastropod; Vermetus.
The line denotes the axis of coiling; mark the absence of symmetry plane in (d), though it looks like
planispiral.
Such coiling does not have any symmetry; as coiling (Figure 11.1; e.g. Euomphalus, an extinct form; and
proceeds and whorls are added, there is also a Planorbis, which lives even today).
translation along the axis of coiling. As a result, the The sense of direction of coiling does not
whorls never lie in any single plane. pertain to planispiral coiling. On the other hand,
In a few gastropod genera, found only as fossils conispiral forms may be left- or right-handed. To
(e.g. Bellerophon; Figure 11.1: their inclusion in the ascertain this, the shell is held apex upwards and
class is, however, questioned by some authors) it is the aperture towards the observer. The position of
otherwise, where the whorls lie in a plane that defines the aperture, to the right or left, determines the
a bilateral symmetry. Such bilaterally symmetrical respective type of coiling. Alternatively, when the
coiling is called planispiral. Besides, in a few other coiling is clockwise, viewed from apex, it is right-
genera, either extinct or extant, there is so little handed or dextral; when it is anticlockwise, it is left-
translation along the axis, that the shell looks very handed or sinistral. Sinistral coiling is rarer; Physa
much like the planispiral forms, but without any is a significant Indian example; in a very few cases,
symmetry. Such coiling is pseudoplanispiral, to be dextral and sinistral types are found equally
identified only by the absence of symmetry abundant in populations of the same species.

11.4 Compactness of Coiling
Gastropod shells may have few or many whorls
(a whorl is a complete 360 volution or coil). The
animal resides in the last whorl of the shell (also
called body whorl); the rest of the shell is vacuum
and called the spire. In loose coiling, a greater
number of earlier whorls are seen from outside. On
the other hand, coiling becomes more compact with
each succeeding whorl increasingly overlapping the
earlier whorls. In such cases, earlier whorls are
found only partially from outside and may be totally
covered by the last whorl (i.e. by each succeeding
whorl at any stage of growth). Different conditions
are designated by different terms, usage being
controversial to some extent. Presently, two terms
are preferred: evolute, where earlier whorls are
fairly, distinctly and considerably visible; involute,
where they are completely or nearly completely
covered by the last whorl. However, different
degrees of compactness of coiling are often found
suitable for identification at generic level. Factsheet
11.1 gives a comparison between the recent usage
and an older one that may be found useful for the
purpose.
11.5 Orientation, Dimension and
Shape
Gastropods crawl on its body holding their shells
above the body, slightly leaning towards back.
FACTSHEET 11.1
Compactness of Coiling (Applies Both to Gastropods and Cephalopods)
Types of compactness Characteristics
Evolute Loose whorls not in contact
Advolute Whorls adjacent and in contact
Involute Each whorl partially overlaps earlier whorl
Convolute Each whorl overlaps earlier whorls
completely
Chapter 11 Gastropoda (Mollusca) 179
Thus, the apex points towards the posterior and
the aperture is at the anterior end of the shell. At
any point on the shell, the anterior-posterior
direction is determined with respect to this frame.
More conventionally posterior and anterior are
referred as apical/proximal and oral/distal. (see
Figure 11.2).
Since dimensions are meant to indicate some
or other aspect of shape, for gastropod shells the
length or breadth bear no significance. Instead,
dimensions are expressed as (a) spiral angle and
(b) relative length (or height) of spire and body-
whorl. The first is measured as an angle subtended
between the two diametrically opposite tangents
that touch the whorls of spire and in the second
the lengths are measured along the axis of coiling
or translation. These two are main parameters that
define different types of gastropod shells; third is
the curvature of the shell or whorl surface (convex
or flat, rarely concave). Figure 11.3 shows a few
variants of gastropod shell-shapes.
In planispiral gastropods, dimensions include
the maximum diameter of the shell and the
maximum thickness measured at right angles to it.
The closer the two, the more rounded or spherical
the shell becomes. Shell assumes flattened, discoid
shape as diameter increases over thickness.
Gastropod genera have characteristic shapes.
However, similar shapes have repeatedly come
up in different genera or species of different
times. If similar kinds of adaptation caused this
and, if so, what were those adaptations, these
Recent usage
Earlier whorls completely or partially
visible from outside
(evolute)
Only one whorl visible from outside
(involute)
180 Part Two: Major Invertebrate Groups

2 (c)

4
5
3
6

8
7 (b)
9

10 11
13

(d) (c)
12
(a)

7 and 8
(f)

8
(e)

Fig. 11.2 Gastropod shell, different features.

Number index: 1 Adapical/posterior, 2 Apical angle, 3 Spire, 4 Shelf, 5 Suture, 6 Ramp,
7 Umbilicus (open in (f)), 8 Columella (solid in (e), hollow in (f)), 9 Body whorl, 10 Columellar fold
on inner lip, 11 Outer lip, 12 Abapical/anterior, 13 Aperture.

are the questions yet unresolved. A few instances
may be relevant in this regard. For instance, in
Silurian succession of Nova Scotia, sedimentary
rocks of different environments are found to contain
gastropod genera of different types of shapes. Thus,
hard, rocky or soft, sediment-laden substrates of
open shallow seas, soft substrates of partially
restricted lagoonal lakes at the land-sea margin and
such other environments—each bear gastropods of
characteristic shapes.
11.6 Functional Morphology of
Gastropod Aperture
Even though functional or adaptive significance of
gastropod shell-shapes are still unknown, the
functional morphology of gastropod aperture is
fairly well brought out. In this regard, torsion in
gastropod body or shell and its relation with the
mode of living and feeding habits assume great
relevance.
 Chapter 11 Gastropoda (Mollusca) 181

(a) (b)

(c) (d)

(e) (f) (g)

(h)

(i) (j) (k) (l)

(m) (n) (o)

Fig. 11.3 Gastropod shells: variations in shapes, canals and ornaments (scale in mm).
182 Part Two: Major Invertebrate Groups
Gastropods are mainly benthic; majority of
them are vagrant, some sessile. Sessile forms are
filter-feeders like brachiopods; vagrant gastropods
are deposit-feeders or hunting predators. They feed
on algae or other plants growing on wet substratum
of basins or other smaller organisms living there.
Hunting gastropods prey upon larger organisms
(viz. bivalves), drilling through their shells or
prying open their closed shell.
In either sessile or vagrant mode, the aperture
of gastropod shell is always important. Located at
the distal or anterior end of the shell, it is the only
passage through which the animal can bring its
body out of the shell or withdraw it within. Both
the head and the mouth of the animal are situated
at this end.
Torsion in early ontogeny produces two
effects. The shell, which in molluscan basic plan
lies on the body, is brought towards posterior end
of the body, on account of the torsion. This may
have proved advantageous for relatively free
movement of the animal, because it is easier for
the animal to bear the weight of the shell, shifted
towards posterior from its position immediately
above the head or the body. But at the same time,
torsion brings the anus towards the anterior end to
be placed above the mouth. This causes foul water
coming out of the body to flow above the mouth.
In such a case, there always remains a possibility
of its being drawn into and, thus, mixing up with
fresh water flowing into the mouth for food and
respiration. The resulting pollution is definitely
harmful for the animal.
Gastropods seem to have tried a number of
adaptations to confront the problem. Thus, in one
subclass Opisthobranchiata there is a detorison in
some genera. It brings the anus back to its posterior
position.
In an order Archaeogastropoda of another
subclass Prosobranchiata, there is often a small
hole (called trema) at or near the apical end or a
deep reentrant or cut (called slit) on the outer lip
of the aperture. As the shell grows the slit changes
its position continuously, thus marking a depressed
slit-band (also called selenizone) on the outer
surface. Both trema and slit are used to act as
passage for expelling foul water.
Another order, Caenogastropoda, of Proso-
branchiata, takes to adaptation to an advanced
means. Many genera of this order develop an organ
called siphon. It is tubular and can suck in fresh
water. Foul water is thrown out through aperture
via a different route. The fleshy siphon is
accommodated in an additional growth on the
apertural margin of the shell, known as the siphonal
canal. It may be long or short as the siphon is
(see Factsheet 11.2). In some genera, for example
in Cypraea, there are two siphonal canals, the
anterior for intake of water and the posterior one
for expelling foul water.
Siphon has found some other uses too. Many
caenogastropod genera are endobenthic, living in
burrows they make. Their siphon is used as
proboscis for respiration or finding out food. In
fact, carnivorous hunting caenogastropods use this
siphon for locating preys.
A third subclass of Gastropoda, viz. Pulmonata,
finds a different solution to the pollution due to
torsion. It has already been mentioned that they
develop numerous tiny blood-circulating parts in
their mantle wall. These help the animal acquire
oxygen directly from air, to make it land-dwelling.
Those pulmonates that continue to live in water,
develop a few folds in their mantle, which act as
gills and help them get oxygen directly from
water. So, these gastropods do not face the
problem of developing any special structure for
expulsion of foul water and, hence, do not have
any corres-ponding feature in their shells. See
Factsheet 11.2 for different features of gastropod
aperture.
Of the four divisions referred above, Archaeo-
gastropoda is the most ancient and primitive. The
other three evolved at different times in different
courses of evolution. They adapted to different
means of solving the problem of intaking of fresh
water and outpouring of foul water, which brought
in corresponding changes in their shell morphology

FACTSHEET 11.2
Features of Gastropod Aperture
Geometry of apertural opening
circular in Turritella
curved slit-like in Cypraea
straight slit-like in Conus
Outer and inner lips
inner dentate, outer smooth in Nerita
both lips smooth simple in Turritella
outer with a slit in Bellerophon, Pleurotomaria, etc.
Canals
absent in Turritella
short, twisted anterior in Cerithium
short twisted both anterior and posterior in Cypraea
too. Changes caused varied morphology,
characteristic at genus and species levels. At the
same time, the analysis of their functional
significance help understand their living and
feeding habits. Thus, archaeogastropod genus
Bellerophon has slit-band, a rather primitive means
of expelling foul water. It may have lived, as
present-day archaeogastropods do, in clear water
on hard substratum. Since such a rocky substratum
does not hold rich flora on it, the genus was more
likely to be filter-feeder, feeding upon phyto-
planktons in the water. On the other hand, the long
anterior canal of Fusus, a caenogastropod points
to their probable carnivorous predatory habit.
11.7 Internal Structures
Among other features that often help identify
genera or species, there are columella and
umbilicus. Columella is a rod-like structure that
runs along the axis of a conispiral shell. As a
conispiral shell is formed by continuous growth
and coiling of an initial cone, the outer surface of
the shell is formed by the outer margin of the cross-
section of the cone (which is really the outer lip of
the aperture). On the other hand, the corresponding
inner margin (i.e. the inner lip of the aperture),
which is coiled along the axis of the shell itself,
Chapter 11 Gastropoda (Mollusca) 183
semi-circular in Natica
crescentic in Bellerophon
rhombic in Trochus, etc.
inner lip with folds (columellar) in Conus
both dentate, outer infolded in Cypraea
long anterior in Fusus, Murex
abrupt anterior in Conus
describes an inner cone, narrow and variously
disposed. This cone is termed columella. If this
cone is tightly coiled, it makes a more or less solid
rod-like columella; when the inner cone is rather
loose, it defines a hollow columella. It is not seen
from outside. But at the point it meets the base of
the shell, it defines a depression or opening
accordingly as the columella is solid or hollow.
The latter is called umbilicus. A shell without
umbilicus is known as anomphalous (e.g. in
Turritella); shell with umbilical opening is
phaneromphalous (e.g. Natica); in a third case,
the umbilicus may be filled in by callus, a
secondary shell material and the shell is then called
cryptomphalous (e.g. in Nerita). In planispiral
shells there is no columella, since the inner surface
of the whorl section does not describe any cone;
in these shells umbilicus is the depression on two
sides of the shell at or around the axis of coiling
(see Figure 11.2).
Most gastropod shells are void in their earlier
parts, i.e. the spire. But in some genera
(e.g. Vermetus) there is an important variation.
This genus lives in coral or organic reefs and are
sessile in habit. Its shell increases rapidly and
since it is sessile and attached to the substratum,
the shell grows towards the aperture or distally.
In course of this growth, the shell simply makes
the easiest route, swerving around any obstacle
184 Part Two: Major Invertebrate Groups

that it may come across. As a result the final shell
is a long winding narrow cone, much longer than
the body. The animal resides just near the aperture
and the rest is void and, thus, fragile. The genus
develops some transverse partitions within the
shell near its apex and for some distance from it,
to add some strength to the shell. These are
analogous to the septa of cephalopods. Septa are
characteristic in cephalopods (see Chapter 12 for
further details); commonly gastropods do not have
septa. But the rare instance of Vermetus suggests
that, at least to this extent, septa are not unique
to cephalopods. The structure that is never present
in gastropods, but occur in cephalopods is the
siphuncle, which is thus more diagnostic for the
latter class. Gastropods being benthic do not face
the problem of maintaining a buyoant shell. Since
cephalopods are nektic, maintaining buoyancy is
vital for them and it is the siphuncle that helps
them in this process. The point will be discussed
again in Chapter 12.
11.8 Surface Ornaments
Gastropod shells show a wide range of surface
ornaments, often characteristic at generic or
specific level. (See Factsheet 11.3 and Figures 11.4
and 11.4) In summary, ornaments may be
apertural, generally grooves in this case, following
the margin of aperture and marking growth stages
(as growth lines do in bivalve or brachiopod
shells). They may, otherwise, be spiral, i.e. along
the coiling (for example, spiral ribs in Turritella,
or rows of spines in Cerithium) or radial or
longitudinal, being disposed longitudinally from
apex to apertures [grooves in Fusus; varices (sing.
varix), as in Murex].

FACTSHEET 11.3
Surface Ornaments of Gastropods and Cephalopods
Ornament types by position and alignment
GASTROPODA (mainly conispiral) CEPHALOPODA (mainly planispiral)
1. SPIRAL: with coiling 1. SPIRAL: with coiling
2. AXIAL/RADIAL: through apex and base, 2. RADIAL: radiating from centre
longitudinal
3. APERTURAL: parallel to aperture 3. APERTURAL: parallel to aperture
4. PERIPHERAL: along the venter/periphery
Ornament elements
GASTROPODS:
1. Rib : Turritella; costae; node, Cerithium
2. Rib : Fusus; spine; varices, Murex
3. Groove : Natica; Turritella
CEPHALOPODS:
1. Node/tubercle (1 row Stephanoceras; 2 rows Scaphites)
2. Rib (Simple Ceratites; bifurcating near venter Perisphinctes; bifurcating near umbilicus Macrocephalites;
bi/trifurcating once/more Scaphites
3. Groove Nautilus (found only when the shell is preserved)
4. Smooth Nautilus; crossed by ribs Acanthoceras; Carinate, i.e. with keel (carina); ropy in Amaltheus;
sulcate, i.e. with groove or sulcus in Hoplites; Bisulcate-carinate in Hildoglochiceras
 Chapter 11 Gastropoda (Mollusca) 185

Fig. 11.4 Gastropod shells: columella and armaments (scale in mm).

 12
Cephalopoda (Mollusca)
12.1 Introduction
Cephalopoda (cephalon means head; pod means
foot; the animal uses its head as foot to propel its
body) is one of the major classes of the phylum
Mollusca. Animals are exclusively marine
represented today by Nautilus, Sepia, octopus, etc.
Since long, the class assumed importance to
palaeontologists. It found extensive use in
classification and correlation of marine successions
of Mesozoic era. Its genera and species have also
been effectively used in studying and knowing
organic evolution, including its varied aspects. Of
late, studies on the mode of living and functional
morphology of these organisms have also brought
forth newer interesting conclusions. The present
chapter briefly summarizes these different aspects
about this class of animals.
12.2 Cephalopod Groups and
Their Shells
As mentioned in Chapter 11, cephalopod shells have
much similarities with gastropod shells.
In both the cases, shells are made of aragonite and
are distinctly coiled. But whereas majority of
gastropods are conispiral, cephalopods show wide
186
variations. Of the seven subclasses of the class
Cephalopoda, recognized by some authorities
(see Factsheet 12.1) only three, viz. Ammonoidea,
Nautiloidea and Coleoidea show planispiral genera
and species. Even very few nautiloid and coleoid
genera have such shells; quite a few ammonoid
genera are also not planispiral. In fact, judged as a
whole, most of the Palaeozoic cephalopods and
extant coleoids do not have any coiled shells at all.
Their shells are simple cones, long or short, straight
or curved (Figure 12.2). Some ammonoid genera
have conispiral shells and some others have peculiar,
aberrant shell shapes (heteromorphic). They may be
coiled in one part of the shell, straight or curved
conical in the rest; or they may be partially coiled
and the rest detached or uncoiled from it (Figure
12.3). However, cephalopod morphology may be
discussed considering planispiral coiling as the
most general type. As mentioned, majority of
ammonoids, palaeontologically the most important
group, have such shells, as also Nautilus, a typical
recent form (Figure 12.1).
12.3 Shape of Shells
Different aspects of shapes of coiled shells pertain
to cephalopods, as they do to gastropods
 Chapter 12 Cephalopoda (Mollusca) 187

FACTSHEET 12.1
Key to Cephalopod Groups
1. Internal Features
Subdivisions Siphuncle Cameral deposit
Subclasses Position Septal neck Connecting ring Siphuncular deposit
Orthoceratoidea Central Orthochoanitic Narrow Endosiphuncular Enough,
type may be present well formed
Actinoceratoidea Central Retrochoanitic Inflated Endosiphuncular Generally
type well formed present
Endoceratoidea Ventral Retrochoanitic Wide and Endosiphuncular Generally
well-built type may be present present
Nautiloidea Generally central, Orthochoanitic / Varied, though Simple, if Present in
also ventral Retrochoanitic simple present at all older genera
Bactritoidea Ventral Prochoanitic Narrow and No deposit Not present
simple
Ammonoidea Generally Generally Narrow No deposit Not present
ventral, dorsal prochoanitic,
in one group retrochoanitic
in one group
Coleoidea Ventral Retrochoanitic Narrow No deposit Not present
NB: Gills four in number in Nautiloidea; two in Coleoidea, Protoconch bulbous in Bactritoidea, Ammonoidea and Coleoidea

2. Shape, Surface, Suture

Subdivisions
Subclasses Shell shape Surface ornaments Suture
Orthoceratoidea Straight/slightly Colour bands Orthoceratitic, straight
curved cone or grooves
Actinoceratoidea Large straight cone Simple Orthoceratitic, straight
with blunt, rounded apex
Endoceratoidea Largest straight cone Simple Orthoceratitic, straight
(<9 m) in Palaeozoic
Nautiloidea Earlier straight/curved cone, Simple Straight orthoceratitic
later loose to compact planispiral to wavy nautilitic
Bactritoidea Straight/slightly curved cone Simple Straight
Ammonoidea Majorily planispiral, also of Simple to complex Goniatitic, Ceratitic,
different shapes Ammonitic, Pseudoceratitic
Coleoidea Straight cone, sometimes coiled Simple, often coloured Straight
(Cont...)
188 Part Two: Major Invertebrate Groups

FACTSHEET 12.1 (Cont...)

3. Age and Relationships
Subdivisions
Subclasses Geological age Phylogenic relationship Generic example
Orthoceratoidea Up. Cambrian-Triassic Ancestor to other groups Orthoceras, Michelinoceras
Actinoceratoidea Ordovician-Carboniferous Actinoceras
Endoceratoidea Ordovician-Silurian Endoceras
Nautiloidea Ordovician-Recent Nautilus, Lituites, Hercoglossa
Bactritoidea Silurian-Devonian Ancestor to ammonoids Bactritites
Ammonoidea Devonian-Cretaceous Goniatites, Ceratites, Scaphites
Coleoidea Devonian-Recent Belemnites, Sepia, Octopus

(see section 11.4). However, in planispiral shells
(where whorls lie in a plane perpendicular to the
axis of coiling) dextral or sinistral coiling becomes
irrelevant, because the part of the shell on either
side of the plane of coiling (which is also the plane
of symmetry) is a mirror image of the other. This
variation in coiling becomes relevant only with
conispiral shells, as in gastropods.
In regard to compactness, the varieties are
judged, as discussed in section 11.4. Also the shape
of shells is determined in the same manner as with
gastropods. The following variations in cepha-
lopods may only be added:
1. Conispiral shells in cephalopods show much
less variation in shape; they are turret shaped.
2. Based on diameter and thickness, planispiral
shells may be of the following types:
(a) Sphaerocone: diameter~thickness;
spherical in shape, e.g. Nautilus;
(b) C a d i c o n e : d i a m e t e r > t h i c k n e s s ;
slightly inflated, e.g. Macrocephalites;
(c) Planulate: diameter>>thickness; thick-
and parallel-sided disc-shaped, e.g.
Ceratites;
(d) Oxycone: diameter>>>thickness: thin-
and parallel-sided disc-shaped, e.g.
Amaltheus, Phylloceras.
A fifth variety, in fact included in
planulate and oxycone types, may be
distinguished. It is called Serpenticone,
where numerous whorls are visible from
outside, as in Perisphinctes.
3. Major types of non-coiled shells are as follows:
(a) Longicone: long cone with sharp apex
(b) Brevicone: short cone, apex sharp, yet
the shell is bulky in shape
(c) Orthocone: straight cone
(d) Cyrtocone: curved cone
12.4 Aperture, Columella,
Umbilicus and Ornaments
As do gastropods, cephalopod animals too interact
with its environment, coming out through aperture.
However, since cephalopod body does not confront
any problem of mixing of fresh and foul water as
in gastropods, it does not develop any siphon or
such features on the aperture and, hence, shows
lesser variations in that respect.
In cephalopod shells, the height of aperture
(measured in the plane of symmetry) and its width
(measured perpendicular to height) determine the
shape of aperture. The ratio of the two is often
characteristic of genera and species. For height <
width, the aperture is called depressed as in
Stephanoceras, and for width < height, it is
compressed as in Phylloceras. In addition, with
more and more compact coiling part of the later
whorl may be impressed on the immediately earlier
 Chapter 12 Cephalopoda (Mollusca) 189

6
20
7
21
11
8

2 19
10
1 3
3 4 18
(b) (c) (d)
(a)
9 16
16 9

14

13
(i) (ii)
(e)
(ii) (i)
(F)
15

17

(ii)
(g)
Fig. 12.1 Cephalopod morphology.
(a) Section (along the plane of symmetry of a planispiral shell) of a nautiloid animal with central
siphuncle, (b) Similar section of an ammonoid shell and its hypothetical body,
(c), (d) Section perpendicular to the plane of symmetry, (e), (f), (g) A few representative varieties of
ornaments (i) Apertural view, (ii) Axial/lateral view
Number index:
1 Mouth, 2 Anus, 3 Tentacles,
4 Gills, 5 Chamber/camera, 6 Siphuncle,
7 Septa, 8 Venter, 9 Suture,
10 Aperture, 11 Impressed zone, 12 Umbilicus
13 Radial ribs, 14 Bifurcation points, 15 Corrugated venter,
16 Peripheral ornament, 17 Lappet, 18 Whorl width,
19 Whorl height, 20 Umbilicus, 21 Shell diameter
190 Part Two: Major Invertebrate Groups

(a) (b)

(i) (i) (i) (c)

(g) (h) (i)

(ii) (ii) (ii)

(d) (e) (f)

Fig. 12.2 Cephalopod features.

(a) to (c) Non-coiled early cephalopods;
(a) Brevicone, cyrtocone, Oncoceras,
(b) Brevicone, Gomphoceras,
(c) Longicone, orthocone, Orthoceras (sectioned in lower/apical part),
(d) to (f) Siphuncular deposits (i) Longitudinal section; (ii) Cross-section)
(d) Annulosiphonate, (e) Actinosiphonate
(f) Endoconal, (g) to (i) Major types of siphuncle;
(g) Orthochoanitic,
(h) Crytochoanitic,
(i) Holochoanitic,
(for (g) to (i) all longitudinal sections; darker lines show shell, septa and septal necks; lighter ones
connecting rings).
whorl, forming weak or strong impressed zones Surface ornaments of cephalopods are most
on two sides of the symmetry plane. Impressed varied and complex in the subclass Ammonoidea,
zones are strong in Nautilus or Macrocephalites, acquired during the evolution of the group (Figures
weak in Perisphinctes. 12.1, 12.4). In other subclasses, too, there might
Planispiral shells do not have any columella. have been evolutionary changes in surface
Here umbilicus is defined as the portion of the shell ornaments. Factsheet 11.3 presents a comparative
(as seen on the two sides) other than the body whorl. view of gastropod and cephalopod ornaments.
Conispiral cephalopods have a small depressed Bardhan and Halder (2000) also reported ornaments
umbilicus at the end of the axis beside the aperture. in Paracenoceras, Jurassic nautiloid and their
Columella is either absent or non-diagnostic. evolutionary significance.
 Chapter 12 Cephalopoda (Mollusca) 191

(d)
(a) (b) (c)

(ii)
(i)
(e)

(f)
(g)

(ii)
(i)
(h)

Fig. 12.3 Hydrodynamics of cephalopod shells, heteromorphy and dimorphism.

(a) to (d) Position of centre of gravity (cross) and centre of buyoancy (dot) in different forms of
cephalopod shells; stippled part body chamber, e(i) An orthocone without cameral deposits and with
the animal (with all its fleshy mass) in it tends to swing up the apical portion: g (centre of gravity),
b (centre of buoyancy), 1 and 2 successive positions; (ii) The same orthoconic cephalopod can float
horizontal when cameral deposits in apical chambers counterpose the body weight of the animal,
(f), (g) Heteromorph shells of ammonoids, (h) Sexual dimorphism in ammonoids; (i) macroconch,
(ii) microconch.
192 Part Two: Major Invertebrate Groups

(a) (b)

(c)

(e) (f)

(d)

(g) (h)

Fig. 12.4 Cephalopod shells and fossils.

(a) and (b) A recent shell of Nautilus showing convolute coiling, septa, central siphuncle; (a) Lateral
view, (b) Apertural, (c) Longitudinal section of a fossil of a straight shell (Orthoceras) with septa,
chambers and siphuncle, (d) Fossil of a heteromorph ammonoid with straight shell (viz. Baculites),
(e) and (f) Equatorial/sagittal section; (e) of a shell and a broken 3D; (f) shell (both fossils) of cephalopod
showing septa and chambers; in (f), it is evident how septa emerges from the inner surface of the
shell, leaving no trace on the outer surface; only in internal moulds that septa leave their traces as
sutures, (g) Helically coiled heteromorph ammonoid (Turrilites), (h) Strongly ornamented heteromorph
ammonoid (Scaphites: broken in the last whorl).

12.5 Internal Structures
It has been indicated earlier that gastropods and
cephalopods differ fundamentally in internal
structures of their shells (section 11.7). However,
when judged not just on mere presence or absence
of certain features, but in terms of the mode of
living, feeding habit and associated aspects, the
difference may be realized not only just as what
they are, but also why they have been so.
Both gastropods and cephalopods live in water,
though a few gastropods may live on land.
Gastropods are vagrant, benthic, whereas
cephalopods are and were nektic. Hence, gastropod
body and shell must be able to withstand the
hydrostatic pressure on them. Cephalopods, on the
other hand, may need to adjust themselves to
different depths facing different hydrostatic
pressure. At the same time, they have to move
relatively fast and smoothly to catch preys and
themselves avoid predator enemies. These cause
them hydrodynamic problems.
There have been some studies on how
cephalopods meet with these problems. These
have helped obtain a fairly clear picture on how
the different internal structures of cephalopod
shells are related to these problems and on how
they are related to the mode of living of these
organisms. But before that, we may need some
more details about the body and shell of these
animals.
12.6 Septa, Suture, Camera
In its basic shape, a cephalopod shell is a cone
made of aragonite (some organic compound
conchiolin is admixed with this mineral). As the
shell grows with the growth of body, it assumes a
coiled shape by virtue of differential addition in
different parts of the shell. This cephalopod shell
contains a series of nearly tranverse partitions
which divide the shell into a corresponding series
of chambers or camera (see Figure 12.1).
Chapter 12 Cephalopoda (Mollusca) 193
The partitions are called septa (sing. septum).
They extend inwards into the shell from the inner
parts of the wall and so they are not to be found
from outside. Each septum leaves a mark of the
line along which it leaves on the wall; this line, a
groove, is called suture. On an internal mould
where the shell is lost, suture is found on the outer
surface of the mould. These septa and suture
distinguish the subclasses, even genera and species
of cephalopods on one hand, and the class itself
from Gastropoda on the other. Only in benthic
gastropods with very long shell, the apical part is
partitioned by some septa-like partitions
(section 11.7), whereas septa are universal in
cephalopods.
The animal resides in the last or body chamber
thus formed by the septa. It is the largest of all
chambers though the most insecured. After the
animal dies and its body decays, this chamber is
emptied and is, thus, easily broken. Gastropod
animal, on the contrary, resides far deeper inside
its shell. In cephalopod shell, the portion other than
the body chamber is called phragmocone.
Chambers of phragmocone is partially filled by
fluids.
12.7 Siphuncle Draws the Main
and Fundamental Difference
with Gastropods
Siphuncle is one of the most important internal
features in cephalopod shells (Figure 12.1). As it
is found from living Nautilus, it is a tubular
structure that starts from the last septum or rather
from behind the animal body in the last chamber.
It then runs across all septa piercing through them
up to the first chamber or protoconch. During the
lifetime, it is filled with living material,
surrounding which there is a horny tube made of
conchiolin fibres and then another tube of aragonite
cystals. Both the horny and aragonite tubes are
porous, allowing passage of fluids, to and fro.
In some well-preserved ammonoid fossils,
194 Part Two: Major Invertebrate Groups
siphuncle is found to be made of calcium
phosphate, though in the earlier parts of the shell
the material is calcium carbonate (the presence of
any organic matter is not known).
Foramen is the perforation through which the
siphuncle passes. The latter actually consists of two
parts: (i) septal neck, which is a bent portion of
the septum itself and is, thus, strong and non-
porous, and (ii) connecting ring, which runs
between two septa across the void chamber and is,
thus, fragile (Figures 12.2, 12.4).
Siphuncle is the structure that distinguishes
cephalopods from gastropods, since it is never
present in the latter group.
12.8 Predatory Habit and
Developed Brain
To deal more with siphuncle, some other issues
need be considered. Cephalopods are of predatory
habit; they hunt their preys with the help of their
tentacles around mouth. In addition to radula with
teeth, they have strong jaws in mouth. They take
water in for respiration, i.e. for oxygen through a
narrow, slit-like passage and eject foul water from
the body through a tubular passage called
hyponome. Water is thrown out in jets; the thrust
involved help the animal move in a sort of jumpy
motion. For hunting, they require a fairly developed
and organized brain, and sharp sensory organ like
eyes. With these they can control their movement,
at least to some extent, with changes in ambience
or for necessity of hunting.
12.9 Movement Control in
Cephalopods
Cephalopods require two kinds of controls in
their movement; (i) to float or move horizontally
in water in the most effective posture maintaining
a neutral buoyancy, and (ii) to move vertically in
water. Obviously fossils do not provide any direct
information on these points. For that we require
studies on living animals. Observations on
particularly two genera Nautilus (subclass
Nautiloidea) and Spirula (subclass Coleoidea) have
provided significant information about the nature
of movement of cephalopods and its relation with
the body or shell.
In chambers of phragmocone, the gaseous
fluid is present in less than 1 atmospheric pressure
(in bladders of fishes, air occurs at several times
larger pressure; fishes sink or rise by increasing
or releasing this pressure, respectively).
Moreover, in earlier chambers of phragmocone
this pressure is greater. Later chambers tend to
contain some liquid in place of gaseous material.
At any stage of ontogeny, the animal resides in
the last chamber being seated on the last-formed
septum. As the body grows in size, newer shell
material is secreted around the body to
accommodate it. At some stage of growth, the
body leaves the septum and moves forward
towards the aperture. The space between the
septum and the body is then filled up with a body
fluid (liquid) acting like a cushion. Soon after, a
new septum starts growing behind the body, thus
adding a new chamber to the phragmocone. The
newly-formed chamber is at first filled up with
the liquid already existing. After the septum
becomes sufficiently strong, the fluid is gradually
extracted through the permeable connecting ring
of the siphuncle. The void formed is slowly filled
up with a gaseous substance. This explains why
earlier chambers are filled with gas and later ones
with liquid. The animal can continue to extract
or introduce liquid from or into the chambers. It
takes place through osmosis, whereby not only
the volume of liquid is changed, but also the liquid
which is left behind in the chambers loses solutes
and, thus, becomes lighter. Salts extracted are
deposited in the pores of siphuncles. There is,
thus, a change in density, though very slowly,
which is utilized by the animal in controlling its
buoyancy and vertical movement.

12.10 Contradiction between
Weight and Buoyancy:
Stable Posture of Swimming
Animals
The question that may come out: how could the
animal keep its posture right during these activities
or when it is swimming.
Both Nautilus and Spirula have planispirally
coiled shells, though it is external and convolute
(involute in present terminology) in Nautilus and
internal and advolute (or evolute) in Spirula.
An object floating in a fluid medium is acted
upon by two forces. One is the gravity, which pulls
the object downwards acting on the centre of
gravity. Second is the buoyancy, which thrusts the
object upwards, acting on the centre of buoyancy.
The position of these points, in animals with
planispiral shells depends on the compactness of
coiling. The more the two points are separated from
each other, the more stable the shell is or rather
the animal in floating or swimming with the body
and its bilateral symmetry plane kept vertical. This
is the neutral buoyancy that is controlled largely
by the shape of the shells (Figure 12.3).
12.11 Pressure at Depth and
Strength of Shell
The next questions that need be sorted out are: upto
what depth and how cephalopod shells (in this case
Nautilus or Spirula) can stand the hydrostatic
pressure without imploding and how do these
information help palaeontological studies of
cephalopods.
It has been found that Nautilus does have
such a strong shell as to descend down to
maximum 600 metre depth, though judged from
natural and laboratory observations, it appears that
it is not impossible for the animal to go upto even
785 metre.
Cephalopods have fewer extant re-
presentatives, though with a rich fossil record.
So, palaeontological significance of these
Chapter 12 Cephalopoda (Mollusca) 195
information and studies have much bearing for
understanding the group.
12.12 Shell Shape and Posture
As discussed, there are two basic morphotypes of
cephalopod shells. In one, the shell is not coiled;
it is a straight or curved cone that are found mainly
in Palaeozoic representatives of the class, though
later, at different times there have arisen some
genera or species with this kind of shells. The other
is coiled. Palaeozoic non-coiled forms have simple,
aperturally concave septa, straight sutures,
generally no ornament, though in some orthoconic
genera, the dorsal part bears colour bands. These
bands might have been used for camouflaging the
animal, in the same manner as dorsal markings or
bands in fishes are used. It means these orthoconic
forms could swim keeping their straight shells
horizontal, dorsal side up. But since the last
chamber contained the heavy fleshy mass of the
animal, they could not have normally kept the shell
horizontal, if the centre of gravity and centre of
buoyancy would have been wide apart from each
other. The much lighter apical part would then be
thrown up by buoyancy and the heavier apertural
part drawn downwards by gravity to make the shell
vertical in the water. These Palaeozoic forms,
however, show considerable development of
cameral deposits of calcium carbonate inside
chambers of the apical portion of the shells. These
deposits are again more common on the ventral
surface of shell interior. It, thus, seems logical that
these cameral deposits acted as counter-balancing
weight as against the apertural fleshy mass to keep
the shell horizontal, bringing the centres of gravity
and buoyancy nearer to each other as required to
maintain this posture (Figure 12.3).
12.13 Case of Ammonoids
Barring some genera, called heteromorphs, with
aberrant shell-shapes, most ammonoids are
196 Part Two: Major Invertebrate Groups
planispiral. The former include some non-coiled
cones, but more commonly they show peculiar
shapes of shells: partially coiled, otherwise straight
or curved conical, or even conispiral, etc. Lower
Devonian Hunsruckschiefer formation of Germany
presents fossils of Bactritioidea (considered
ancestor to ammonoids) and Ammonoidea, that
help bring out the changes from straight shells
through loose coiled to compact coiled types, a
trend that might have been followed later in the
subclass itself. It was inferred therefrom that early
ammonoids or bactritoids were swimmers with
their body and shell held horizontal. As coiling
assumed importance and more and more genera
became planispirally coiled, the animals took up
the posture now assumed by Nautilus or Spirula,
in which the symmetry plane is held vertical,
aperture downwards. Intermediate loosely coiled
genera of Devonian might have been less efficient
swimmers or even sluggish bottom-dwellers.
Heteromorphs that are found in the later parts
of ammonoid history, might have had the similar
postures while swimming. It means then, the
aberrant-shaped cephalopods did not suggest any
degenerative stage, as held earlier. They were
adapted as well to swimming as the normal
planispiral forms were.
Similar type of chambered phragmocone with
siphuncle present in the recent Nautilus or even
Palaeozoic nautiloids to ammonoids, prompt us to
conclude that they had similar kind of movement
as hydrodynamic object in water with similar
constraints and advantages. But as it will be evident
from the Factsheet 12.1, there were also many
differences between nautiloids and ammonoids.
Of them, the strength of siphuncle is related to
hydrodynamic problems confronting ammonoid
shells. As ambient hydrostatic pressure increases
with depth, siphuncle in the shell interior becomes
susceptible to implosion. Based on observations
on Nautilus, the strength of siphuncle under
pressure to prevent it from implosion is expressed
by (h/r × 100) where h is the thickness of shell
wall and r is the radius of siphuncular tube.
In Nautilus it is high, near (~10-19) or suitable for
living at depth of < 600 metre. Further it is found
that in Mesozoic ammonoid orders, Lytoceratida
and Phylloceratida, this value is slightly less than
that for nautiloids, suggesting they were suitable
for depths ~450 metre. In later ammonoids, the
value is much less, between 3 and 6.5, or for depths
much less, ~100 metre. Since these later forms
were descendants of lytoceratids and phyllo-
ceratids, it may be concluded that in course of their
evolution, ammonoids went through a change in
adaptation from deeper to shallower waters, at least
in some lineages.
Ammonoids also differed from nautiloids in
suture patterns they had. However, the significance
of septa or suture is still a debated issue.
12.14 Ammonoid Suture and
Heteromorphy
There are several types of sutures found in
cephalopods (viz. orthoceratitic, nautilitic,
goniatitic, ceratitic and ammonitic). Of them,
orthoceratitic suture, a smooth, straight line when
unfolded and nautilitic, a simple wavy suture are
found in the nautiloid and other earlier
subclasses. Sutures of ammonoids are the most
varied and complex among these types (see
Factsheets 12.1 and 12.2). But a complex suture
does not necessarily mean that the corresponding
septal partition that hangs inside the shell is
similarly or equally complex. In fact, that portion
of the partition is simple, convex or concave. But
such simple partitions are attached to the interior
of the shell wall along simple or complex lines,
known as sutures. Complexity entails
subdivisions of larger or broader curves into
several smaller ones that ultimately produce fine
frills. Curves in suture that are convex towards
aperture are called saddles and those concave are
lobes. Factsheet 12.2 and Figure 12.5 show how
ammonoid sutures change towards more and
more complex types.
 Chapter 12 Cephalopoda (Mollusca) 197

FACTSHEET 12.2
Variation and Evolution of Suture in Ammonoidea

Primary suture Suture during maturity Occurs in Order Geological age

Type/no. of lobes Broader type Character

Trilobate3 Goniatitic Smooth saddle, Goniatitida Devonian-Permian

angular lobe
Quadrilobate4 Ceratitic Smooth saddle, denatate, Ceratitida and Mainly Triassic
fine folded lobes Phylloceratida
Quinquelobate5 Ammonitic Finer lobes in larger Few genera of Triassic
smooth saddles Ceratitida
Quinquelobate5 Ammonitic Finer lobes in larger Phylloceratida, Jurassic
smooth saddles Ammonitida and most -Cretaceous
genera of Lytoceratida
Hexalobate6 Ammonitic Finer lobes in larger Few genera of Cretaceous
smooth saddles Lytoceratida
Quadrilobate4 Pseudoceratitic Simpler like ceratitic Heteromorph genera of Cretaceous
suture Ancyloceratida

Naturally, a smooth straight suture will have a
shorter length than a highly frilled suture. It means
corresponding septum of the latter type will have
a longer contact with the wall. This signifies that
the more complex the suture is or was, the greater
supporting strength is or was added to the wall.
Hence, such forms are more likely to withstand
more hydrostatic pressure at greater depths. But
laboratory studies have indicated that there is a limit
to this, beyond which further frilling does not add
any more strength to the wall (Clarkson 1998).
Besides, as discussed earlier, lytoceratids and
phylloceratids with a greater strength of siphuncle
were inhabitants of deeper waters than their
descendant ammonitid; but suture in the two former
groups were less complex than the suture of
ammonites. This has prompted palaeontologists to
seek for alternative explanation of the significance
of septa and suture. Conclusion is yet awaited.
Below a brief note is added to highlight the
opinions.
1. Cephalopod body and shell have an episodic
growth. As indicated, at each stage the body
moves forward in the last chamber, its
attachment to the septum is temporarily lost,
to be regained at the next stage. It is suggested
that the increased area of a frilled septum helps
body or its muscles to find firm hold to the
former.
2. Increased area of attachment of septa with the
wall or length of suture might have helped
distribute the ambient presssure to make it
bearable.
3. At the beginning septa are thin sheets of
organic material. They are then mineralized.
Before mineralization, equal pressure on two
sides of each septum is likely to cause the sheet
to wrinkle, which are retained after
mineralization. In that case, function of the
septa towards adding a supporting strength to
wall is really an added advantage.
Before concluding the discussion on septa and
suture, mention must be made of simplification of
suture from the complex ammonitic type to those
of the order Ancyloceratida. Suture of this order
is termed pseudoceratitic and this change
from ammonitic to pseudoceratitic suture was
considered as an example of degeneration.
198 Part Two: Major Invertebrate Groups

E L U I 4¢ E L U2 U1 I (5) E L U 2 U 1U 3 (6)

VI (i)
(f)

V (h) E L U 2U 1 I (5)
(g)

IV E L U2 I (4)

(e)
III
(d)
(c)
II (b) E L I (3)

I (a)

(ii)
(iii)

(i)

(iv)

(v)

Fig. 12.5 Ammonoid phylogeny and evolution of primary suture.

Periods: I–Devonian, II–Carboniferous, III–Permian, IV–Traissic, V–Jurassic, V–Cretaceous
Major groups: (a) Anarcestina/Ar, (b) Clymeniina/Cl, (c) Goniatitina/Go, (d) Prolecanitina,
(e) Ceratitina/Ce, (f) Ammonitina/Am, (g) Phylloceratina/Ph, (h) Lytoceratina/Ly, (i) Ancyloceratina /An
Primary lobes: E (External/ventral), L (Lateral), U (Umbilical), I (Internal/dorsal)
Primary suture: 3 Trilobate (in Go), 4 Quadrilobate (in Ce, also in Ph, Am, Ly), 5 Pentalobate (in Am)
6 Hexalobate (in Ly), 4¢ Quadrilobate (in heteromorphs of Ancyloceratida)
Cephalopod sutures: (i) orthoceratitic, (ii) nautilitic, (iii) goniatitic, (iv) ceratitic, (v) ammontic

Ancyllocerids are characterized by heteromorphs.
As mentioned, these heteromorphs might have
been well-adapted to the swimming mode of living.
Thus, they could have avoided high pressures at
depth, if necessary by swimming out of that
environment. It would not require for them to
develop complex suture to support their shell wall
under pressure. So, the case may have nothing to
do with degeneration; it marked only a special kind
of combination to a specific adaptation.
Though conclusions on many debates are still
awaited, it is clear from the above that siphuncle
and septa, the two internal structures of cephalopod
shells, are in all likelihood linked up with their
adaptation to nektic mode of living. This is where
these animals are fundamentally different from
gastropods and, hence, there are the ubiquitous
presence of septa and siphuncle in cephalopods
and their variations.
12.15 Dimorphism in Ammonoids
Several species of ammonoids show marked
paired variation in size, along with certain other
morphological differences. This phenomenon is
interpreted as reflection of sexual dimorphism,
i.e. two kinds of morphology in males and
females, respectively (Figure 12.3). This was
substantiated as far back as in 1960s indepen-
dently by Callomon and Makowski (Lehmann and
Hillmer 1980; Clarkson 1998) on the basis of
fossil evidences. Earlier suggestions of
dimorphism, dating back from 1869 (vide
Waagen) lacked strong evidences. Dimorphic
species, as now accepted by many pala-
eontologists, have macroconchs, thought to be
shells of females, several times larger than
microconchs, supposedly shells of males.
Dimorphism is evident only in adult parts of
shells, in which growth had ceased. The characters
significant in this regard are crowding or
approximating of sutures due to diminishing
growth rates, different surface ornaments on the
final body chamber, slight uncoiling of the body
Chapter 12 Cephalopoda (Mollusca) 199
chamber from the rest of the shell or special
modifications in the mouth or peristome. In some
forms, the microconchs are characterized by
lappets, i.e. simple or necked spatulate projection
of edge of aperture of body chamber or the margin
of the mouth, acting as some constriction on the
latter.
The issue of dimorphism is beset with
controversy. For example, it is often uncertain as
to how to recognize them. Since dimorphic pairs
are found in the beds of the same age, it is held
more likely not to indicate variation of morphology
in different, though related species. In the latter
case, differences are expected to be time controlled.
But even then, care must be taken, particularly for
examples, in which the two forms of the same pair
do not occur together.
Secondly, what should be the fate of already
described separate species that are now found to
be dimorphic pairs? Should they continue to bear
the different names as different morphospecies or
should they be now designated with the same name,
signifying that they belong to the same biological
species? The point is not yet settled.
A recently reported Indian example of
ammonoid dimorphism is obtained from
Kheraiceras cosmopolitum, from Lower Callovian
Chari Formation of Kachchh. Here, the macro- and
microconchs resemble each other except in size
and in terminal constriction in aperture of
microconch. Besides a ‘parallel’ evolution in
micro- and macroconchs is inferred, that “improves
the understanding of evolutionary trends involving
complex heterochronic processes” (Bardhan et al.,
1994). Dimorphism is also reported in Jurassic
nautiloids of Kachchh (Bardhan Halder and Jana
1994) and in Placenticeras from Upper Cretaceous
Bagh Beds (Ganguly and Bardhan 1993).
12.16 Ammonoid
Palaeobiogeography
Though ammonoids appeared in Devonian and
underwent an adaptive radiation of goniatitic forms
200 Part Two: Major Invertebrate Groups
(e.g. Goniatitida and Clymeniida) in Permo-
Carboniferous, they became really important
geologically in Mesozoic.
The end-Permian extinction, the severest of
all mass-extinctions in the earth’s history, affected
ammonoids like many other groups. Only a few
genera (e.g. Xenaspis and Xenodiscus) are known
to have survived the Permian crisis. However,
Lower Triassic ammonoids were prolific being
represented by more than hundred genera of
Ceratitida. The radiation appears to stem from
Ophiceras, a descendant of Xenodiscus.
Palaeogeography during this period was also
interesting. Permian land masses had a complex
geography, with a number of high mountain ranges
affecting wind pattern. The ocean separating
Europe from Asia was closing during late Permian
along the Ural Mountains to form the
supercontinent Pangaea. The bulk of the
continental crust formed one vast continent above
sea level. The Tethyan seaway existed as an
embayment of the deep sea projecting from the east
into the equatorial Pangaea much in the positions
of the present-day Mediterranean.
Towards the end of Triassic, fragmentation of
Pangaea started in the Tethyan region. It became a
deep, narrow arm separating the present-day
southern Europe from Africa. Separation of North
and South America and the Gondwanaland
continents followed suit.
On this palaeogeographic background,
biogeographic distribution of Triassic organisms
was undifferentiated both in sea and on land. The
situation persisted in the main till Jurassic when
there developed two biogeographic provinces of
marine life in Europe, viz. the Tethyan and the
Boreal realms. Coral reefs and limestones were
restricted to the Tethyan realm suggesting a
warm tropical condition there. The Boreal was
colder, much similar to some siliciclastic-
dominated subtropical waters without reefs
existing today adjacent to tropical reef-studded,
carbonate-depositing areas in eastern coasts of
North America. Temperature gradient was perhaps
gentler than today and there were neither well-
marked climatic belts.
The two realms were distinctly differentiated
in Lower Jurassic with largely endemic 5
ammonoid genera in the Boreal and about 14
genera in the Tethyan. Higher up in the column
differentiation was reduced, with genera
attaining wider geographic distribution. For
instance, Spiroceras, a Jurassic heteromorph,
became a good index fossil in Europe and also
in America, West Asia, Madagascar and India.
Marine oscillations or renewed transgressions in
Middle Jurassic may be the cause of reduction
of differentiation of realms. Physical barriers,
depth variation of sea, sedimentary facies
(carbonate/ siliciclastic) and organic factors such
as inter- and intraspecific competition may have
minor roles to play. Some authors consider water
temperature to be the cause; but the absence of
marked climatic belts as revealed by
contemporary plant fossils goes against this
suggestion. The Boreal was more homogeneous
and distinct in Upper Jurassic; the Tethyan realm
was marked by local provincialities, viz.
Mediterranean, Himalayan (Indo-Pacific: e.g.
Spiti Shale and Kachchh fauna with Kossmatia,
Virgatosphinctes, etc. traceable into Indonesia
and Australasia), Ethiopian, etc.
Cretaceous global geography was marked by
continued fragmentation of Pangaea and drifting
apart of different continental masses (in the
present-day terms). Climates were warm, though
temperature changed in different ways at different
places, as it could be revealed from oxygen isotope
and plant evidences. Sea level stood higher in
relation to most land areas than it had earlier.
Northern land connnections were, thus, breached.
The tropical Tethyan realm remained.
Cretaceous ammonoids include many
cosmopolitan genera, even species. It means,
though there were three broad provinces, viz.
Boreal, intermediate and Tethyan (tropical), they
were well-connected too. In the third, pseudo-
ceratitic ammonites were adapted to tropical shelf

seas, the reasons being not clear. Intimate faunal
relations existed between both sides of the Atlantic.
Close affinity was also there between the neritic
sea faunas in South Africa, Madagascar,
South India and West and North Australia, or
between areas around the North Pacific. The latter
was different from the interior North American
province, suggesting a circum-North Pacific
orogenic system between the two. There were free
communications between circum-Pacific and
circum-Indian Ocean, with Japan and Madagascar
having some similarities.
Distribution of ammonoids was, thus, largely
controlled by palaeobiogeographic factors.
However, life history, mode of life and post-
mortem floating, etc. might also have been active,
at least in more local scale (see Hallam 1973,
Stanley 1989, 1993 for more details).
12.17 Ammonoid Palaeoecology
Palaeoecological studies of ammonoids have
certain inherent difficulties. As it has been
indicated above (section 12.13), ammonoids with
planispiral or other kinds of shells were adapted
to a mobile mode of life in a fluid medium, i.e. to
an active efficient nektic habit, similar to that of
present-day nautiloids and coleoids. It means the
environment in which they lived, differed from
the environment in which they were fossilized.
Like other cephalopods, ammonoids too had
argonitic shells which readily dissolved during
lithification and diagenesis. It was for this reason
that ammonoids are obtained commonly as
moulds, mostly internal, but also external in cases.
There are ample evidences that such moulds of
ammonoid fossils contain pyritiferous sediments.
Ammonids also occur in black shales (e.g. in Spiti
Shales of Jurassic). These point to the fact that
ammonoid remains were often buried in anoxic
condition of deeper waters also suggesting that
they were open ocean dwellers. Values of h/r
× 100 (h: thickness of wall; r: radius of the tube),
Chapter 12 Cephalopoda (Mollusca) 201
a measurement of strength, for phylloceratids and
lytoceratids come closer to those of present-day
Nautilus pointing to their preference of deeper
water milieu. The same values for ammonitida
are low and so they might have been shallower
water forms.
In this connection it may be mentioned that
throughout Palaeozoic and Mesozoic, the oxygen
level of sea water was lower, particularly at
middle or greater depths, than they are today. This
is because, in the absence of polar ice caps, there
was no vertical oceanic circulation. On account
of this vertical circulation, present-day cooler
polar water moves downwards and across latitude
towards equator, where it warms up and swells
upwards. The process also largely homogenizes
the oxygen level of sea water of different depths.
In Palaeozoic and Mesozoic, this homogenization
was absent and oxygen level was, thus, lower at
middle and greater depths. Organisms like
ammonoids which were adapted to these depths
must have been also adapted to this low oxygen
environment.
It was previously held that heteromorphs were
benthic and crawling in habit. But now it has been
concluded on sufficient considerations that
heteromorphs were not degenerate, neither adapted
to a benthic, crawling habit. The following points
are added to the issue of functional morphological
significance of ammonoid features.
Among the heteromorphs there were different
kinds. For example, helical shells of Turrilites were
well-adapted to a vertical movement with vibrations
or oscillations with apex pointing upwards. Large
highly ornate planulate Acanthoceras probably
lived a sluggish nektobenthic life. Long-bodied
serpenticones, some spherocones and a few other
heteromorphs could have been passive drifter.
Oxycones and streamlined planulates were mobile
nektopelagic of deeper water and were most likely
to be active predators.
Factsheet 12.3 provides some generic
examples of ecomorphotypes.
202 Part Two: Major Invertebrate Groups

FACTSHEET 12.3
A Few Eco-morphotypes at Generic Level
Neritic realm:
Planktonic: Larvae
Passive drifters: Long bodied serpenticone: Perisphinctes; some sphaerocones; some
Cretaceous heteromorphs
Vertical migrants: Heteromorphs like Turrilites (conspiral), Scaphites (partially uncoiled;a)
Mobile nektobenthos: Streamlined oxycones (Amaltheusb) and platycones (Hildoglochicerasb)
Sluggish nektobenthos: Highly ornate planulates (Acanthoceras; Stephanoceras?; Hoplites?—for all threeb)
Oceanic realm:
Planktonic larvae in shallow water
Fragile shelled heteromorphs in deeper water
Vertical migrants in still deeper water
(a) Prominent and numerous ribs: For strengthening shell/As hydrodynamic stabilizers (from pitching and rolling)/As
camouflage measure (by diffusing outlines)
(b) Streamlined, strongly septate forms: mobile pelagic

12.18 Indian Case Studies In Kachchh examples, colour markings are

Colour markings in fossil shells, though rare, are
reported from Middle Cambrian to Holocene.
Organisms secrete pigments that are incorporated
into shells as dietary and/or metabolic byproducts
and colouration patterns develop, as in molluscs, as
a result of differential secretory activity of pigment
producing cells located along the mantle edge. An
Indian example is reported by Bardhan, Jana and
Dutta (1993) from colour bands found in four
specimens of the ammonoid genus Calliphylloceras
from Jurassic Chari Formation of Kachchh.
Pigmentation is generally held to serve
functions like mimicry and camouflage, a
protective measure from larger predators like
fishes. More recently (e.g. Kobluk and Mapes
1989) it has been suggested that melanins,
common pigments in invertebrates, increase the
strength of shells and, thus, protect infaunal
organisms more efficiently against abrasion. In
dark habitats colouration perhaps is used to reduce
reflectance of the shell by absorbing light towards
the blue end of the wavelength making the shell
appear darker.
restricted exclusively to the areas of internal ridges.
These ridges themselves mark periodic pauses in
growth and added strength to the shell wall in
lifetime. Association of the two suggests the same
function of colour markings too. Occurrence of
similar colour patterns in four different specimens
of a single species point to low intra-population
variability and, thus, their being adaptively
significant.
Phylloceratids have compressed, oxyconic,
highly streamlined, smooth and involute shells and
are considered to be pelagic; they are held to be
deeper water dweller (below photic zone). In
Kachchh, they are present throughout Jurassic and
are found in almost all beds of Chari Formation
deposited on a warm, agitated shallow shelf. Their
pelagic habit and post-mortem drift might have
led to the poor facies control of their occurrence.
But the evidences, such as their well-preserved
shells with oyster epibionts, assemblages of
Calliphylloceras with small juvenile to large
individuals, association with typical shallow water
fauna, suggest a life distribution without transport.
Associated macrocephalitids are found to possess

a jaw apparatus preserved in body chamber
implying little or no post-mortem transport. In
addition, the end-Tithonian regional mass
extinction of Kachchh during regression at that
time affected all ammonites including
phylloceratids. Had they been deeper water forms,
it would have been otherwise. So, the authors
(Bardhan Jana and Dutta 1993) concluded that
colour patterns in Calliphylloceras were
functional and that the genus had a nektopelagic
habit in shallow water.
12.19 Evolution of Ammonoidea
The subclass Ammonoidea of the class
Cephalopoda ranges from Devonian to Cretaceous.
Its fossils have been extensively used in particular
in biostratigraphic and evolutionary studies. The
following sections will deal with a summary
account of the main aspects of ammonoid
evolution.
12.19.1 Biostratigraphic and
evolutionary significance
Ammonoids assume biostratigraphic and
evolutionary significance for the following main
reasons:
1. Ammonoids show widespread occurrence
(geographical) even at specific level.
2. Being nektonic, they occur in rocks of different
sedimentary environments of ancient seas—
from shallow to deep water, quiet to turbulent.
3. Their fossils show excellent preservation and
even when the calcareous shell is dissolved,
the rather robust internal mold (or cast) is not
only well-preserved, but also provides ample
important morphological characters from gross
shell form to delicate sutures. These help in
relatively easy diagnosis of genera and species
and other studies.
4. Most ammonoid genera and species show short
vertical or stratigraphic ranges that add to their
Chapter 12 Cephalopoda (Mollusca) 203
importance for their value as time-markers. In
fact, for prolific occurrence and rapid evolution,
ammonoids have been used to erect zone of
even less than a million year duration in
different parts of Mesozoic. They have also been
useful in regional and even global correlation.
5. Each ammonoid shell preserves all the
ontogenic stages from the larval (embryonic)
stage to senility, generally attained after
maturity (through nepionic, neanic and
ephebic stages) and providing features of
senescence or old age (gerontic stage). In
addition to their intrinsic importance, these
ontogenic stages, when brought out from the
study of shells sectioned generally along
symmetry plane, also supply important clues
on phylogenetic changes.
12.19.2 Ancestry of ammonoids
Previously, some coiled nautiloids were considered
as the ancestors of ammonoids. Subsequently, it
was suggested that ‘bactritids’ with straight
(orthoconic) shells, with marginal siphuncle, a
bulbous protoconch (the first chamber),
retrochoanitic (posteriorly directed) septal necks
and gently flexured sutures with lateral lobes, were
the ancestors to ammonoids. These forms are now
included in a separate subclass Bactritoidea,
derived from orthoconic nautiloids, viz.
sphaerorthoceratids, themselves descendants of
michelinoceratids. The earliest true ammonoids
(anarcestids) are recovered from Hunsruck Shale
of early Devonian of West Germany (also see
Factsheet A. 2.5).
12.19.3 Phylogeny
The oldest true ammonoids belong to Anarcestida
(order) of Devonian age. Of the descendant orders
Clymeniida (also of Devonian) is notable for its
dorsal siphuncle, Goniatitida (Devonian to
Permian, one genus in Triassic) for its apparently
conservative phenotypic traits, though giving
rise to later stocks. All these suborders are
204 Part Two: Major Invertebrate Groups
characterized by goniatitic suture, though
differences in ontogenic variations in sutures help
differentiate them. Of the later orders,
Prolecanitida shows ontogeny of suture that
appears to be the normal course of development of
all Mesozoic ammonoids. However, as mentioned
in section 12.16, end-Permian extinction severely
affected ammonoids. Only a few genera (e.g.
Xenaspis and Xenodiscus) are known to have
survived the Permian crisis. Ceratitida arose from
Prolecanitida in Permian (Xenodiscidae) with
typical ceratitic suture; the former, however,
evolved further subsequently. Ceratitida was
abundant and diversified in Triassic, apparently
radiating from the stem-genus Ophiceras, a
descendant of Xenodiscus. The order became
extinct by the end of the period. Phylloceratida
appeared from Ceratitida in Triassic; it continued
upto late Cretaceous, but was again a stock
conservative or persistent itself, though serving as
root-stock for all post-Triassic ammonoids.
Phylloceratida gave rise to Lytoceratida
possibly in early Jurassic, near the boundary of the
period with Triassic; the latter differentiated into a
number of genera and species, and became stable.
Ammonitida is the last major suborder, arising
probably from Phylloceratida in Triassic; highly
differentiated into about 37 families, this order is
now considered polyphyletic, i.e. derived from
different ancestral stocks of Phylloceratida and,
may be, Lytoceratida.
A separate fourth Mesozoic order Ancylo-
ceratida includes Cretaceous hetero-morphs with
four-lobed primary suture similar to that of
Ceratitida and opposed to five- or six-lobed primary
suture in Ammonitida and some Lytocratida,
respectively; Ancyloceratida is supposed to have
been derived from five-lobed Ammonitida. (Figure
12.5) (see Factsheets 12.2 and 12.4)
12.19.4 Phenotypic trends
Evolution of ammonoidea did never follow a
simple clear-cut general path. The different bio-
characters considered important for depicting
evolutionary, i.e., phenetic trends are:
(i) Protoconch (earliest chamber) and
proseptum (septum between protoconch and
first chamber).
(ii) Body chamber (large or small).
(iii) Size of shell.
(iv) Shape of the aperture (simple, constricted
or with lappets, that is simple or necked
spatulate projection of the edge of aperture
of body chamber or mouth border).
(v) The presence or absence of a keel along
venter and its nature—solid/hollow and
open/hollow and floored.
(vi) Ribbing and other ornaments.
(vii) Sutures.
12.19.4.1 Sutures: Of these, sutures are
considered the most important for the purpose,
progressive development of suture being a broad
trend in ammonoid evolution. It involves changes:
1. in the number of lobes in the primary suture
(that is, between first and second chambers);
and
2. in the denticulation of adult sutures.
The first case is elaborated in Factsheet 12.2.
It shows that the primary sutures are of the
following types: trilobate, quadrilobate, two
variations of quinquelobate, hexalobate and then
again a quadrilobate. The trend, thus, includes
increase in lobes of primary suture from three to
six and a reversal to four in Upper Cretaceous.
The different parts of ammonoid sutures,
particularly the lobes are listed in Factsheet 12.4,
along with the symbols used to mark and describe
them as well as their stage of appearance during
the ontogeny. (Also see Figure 12.5.)
The second trend of sutural changes is towards
frilling. It is a pervasive trend, both in overall
change from Palaeozoic through Triassic to
Jurassic-Cretaceous forms and within individual
groups.
 Chapter 12 Cephalopoda (Mollusca) 205

FACTSHEET 12.4
Parts of Ammonoid Suture, Their Symbols and Ontogenetic Stage of Appearance

Types of lobes Symbols Ontogenetic stage of appearance

Ventral/external E Appears first in ontogeny and phylogeny
Lateral L do
Dorsal/internal I do
Umbilical U Appears as finer lobe on dorsal lobe
Adventitous A Appears as finer lobe on ventral lobe
Median M Appears on a saddle on ventral lobe
Sutural S Saddle on umbilical lobe

On these two trends, the broad change of
sutures may be summarized in the successive
appearance, domination and complication found
in stages, goniatitic, ceratitic and ammonitic
(for definitions of different suture types see
Factsheet 12.2).
Appearance of simpler pseudoceratitic suture
in Cretaceous heteromorphs was considered as
degenerative reversal of progressive trend towards
the increase in complexity, an idea which has
since been rejected. Details are discussed in
section 12.14.
12.19.4.2 Size trend
1. Progressive size increase takes place both in
many specific lineages or, in general,
following Cope’s Rule. Thus, largest
Cretaceous (Parapuzosia sepperadiatus) is
bigger than largest late Jurassic (Titanites) or
largest Jurassic (Titanites) is bigger than
largest Triassic (Pinacoceras).
2. It is often held that rapid increase in size led to
rapid extinction because of extreme
specialization and scarcer food source telling
upon a number of individuals leading to
extinction.
3. Size progressively decreased in some
Cretaceous lineages.
12.19.4.3 Ornaments: From smoothness to
ornamentation of surface, increasingly more
complex with time, was a general trend, though
simple ornamented forms coexisted with complex-
ornamented ones.
12.19.5 Indian ammonoids
Indo-Pak subcontinent has a fairly rich ammonoid
record representing different stages of the history
of the group. More important constituents of the
record are given below. The earliest record comes
from Permian of Salt Range in Pakistan, the forms
being Cyclolobus, Xenodiscus, Xenaspis
(X.carbonaria).
Otoceras, Ophiceras and Meekoceras occur in
Triassic of Spiti and adjacent areas. They mark
three successive zones from the lowest Triassic:
Otoceras woodwardi, Ophiceras sakuntala,
Meekoceras varaha are the important species.
These and other ceratitine genera and species,
Ceratites, etc. characterize Triassic of different
areas.
In Kachchh of western India Macrocephalites
triangularis and M. macrocephalus occur in
Middle Jurassic Pachham/Jhurio Formation and
Chari/Jumara Formation, respectively. The former
species occurs also in Spiti and the latter in Salt
Range. Perisphinctes, Reineckia, Hildoglo-
chiceras, Torquatisphinctes, etc. occur in different
upper parts of Jurassic of Kachchh. A phylloceratid,
Calliphylloceras has been extensively studied from
the Chari Formation of Kachchh (Bardhan, Jana
and Datta 1993).
206 Part Two: Major Invertebrate Groups
From Cretaceous Hoplites, Turrilites, etc. are
reported from Cauvery Basin of South India.
Mention may also be made of Belemnites
sulcacutus and Belemnopsis gerardi, two typical
coleoids reported from Jurassic and Cretaceous of
Spiti and other Himalayan areas. In addition,
reports of nautiloid from Upper Cretaceous Bagh
Beds (Gangopadhyay and Halder 1996), including
that of time-marker Cymatonautilus (Halder and
Bardhan 1996) impart added significance to the
group.
12.19.6 General comments
Phylogeny of ammonoids reveal a number of
interesting features about modes and patterns of
evolution. They are as follows:
1. Phylogenetic history, in general, or at any
stage, is never of a uniform diverging
development from a modest initiation, to be
followed by a gradual decline to extinction.
2. On the contrary, it is marked by a few (at least
six of varying importance) major episodes of
adaptive radiation (Up. Devonian, Up.
Carboniferous-Lr. Permian, Triassic, Mid. to
Late Jurassic. Lr. Cretaceous and finally Up.
Cretaceous). During these episodes numerous
short-lived genera and species arose rather
suddenly from relatively slowly evolving
ancestral stocks and themselves evolved for
short periods to become extinct soon.
3. Each of these episodes is succeeded by a phase
of extinction, with or without an intervening
span of gradual decline in varieties of genera
and species, and in abundance (End of
Devonian, End of Permian with a decline in
Up. Permian, Tr./Jr. boundary, Early
Cretaceous and finally end of Cretaceous or
Maestrichtian with a decline earlier to it).
Notably, extinction phases are often marked
by regression of sea (Early Permian, Late
Triassic, Early Cretaceous, Late Cretaceous)
with the radiation taking place with renewed
transgressions.
4. There remained or appeared a stock or a few,
which were persistent, rather conservative. It
meant, they evolved slowly, but could survive
the phases of extinction and did give rise to
newer, numerous descendants in the succeeding
episode of adaptive radiation. (Anarcestida-
Goniatitida; Prolecanitida-Ceratitida; Phyllo-
ceratida-Lytoceratida and Ammonitida; in each
case the first is a conservative long-ranging
order giving rise to a short-lived order or more,
but itself continuing).
This phenomenon in which a conservative
ancestral stock from time to time gives rise to short-
lived groups, expanding and diverging rapidly and
replacing each other successively, is called
iterative evolution. More recent authors, however,
challenge the validity of this pattern on the charge
of over-simplification. (The phenomenon is also
called “palaeontological relay’.)
5. Ammonoid orders, e.g. Phylloceratida and
Lytoceratida, or down to lower levels, even
species, show parallel evolution. It means
different stocks, may or may not be with
ancestral-descendant relationships, evolved
parallelly with similar phenotypic trends.
However, actually the picture in most cases
was more complex, as trends and their rates
varied in different lineages seldom remaining
strictly parallel.
6. Evolutionary convergence leading to
homeomorphy is common in ammonoid
evolution. As expected, the number and type
of functionally viable changes that could take
place on ammonoid shells always remained
within finite, relatively narrow limits. The
changes are limited to a tubular external shell,
the phragmocone; to coiling that cannot
change in such a way as might impede its
floating or swimming habit; to ornaments
varying within limits of radial, spiral or
peripheral positions and ribs, tubercles or
spines as elements or lappets, etc. to sutures,
including prosuture, primary suture and adult

sutures, too, limited in their variations by
functional constraints. The resulting
evolutionary convergence or homeomorphy
may extend to the whole shell, or to specific
features such as the suture line. Homeomorphs
may be synchronous or heterochronous; it
may also be between taxonomically distant
groups and between stratigraphically separated
members of lineages (also see Factsheet 9.3).
7. Evolutionary divergence is well-documented
in ammonoid phylogeny. As mentioned above,
there may be as many as six phases of adaptive
radiation. In each of them, new genera and
species appeared and found adaptation in a
multitude of niches, which were left vacant
with a phase of extinction of earlier forms.
Thus, lytoceratids and ammonitids filled in the
niches evacuated by ceratitids with their
extinction. At lower levels, similar radiations
lead to diverse morphotypes, often convergent
between or within lineages because of limits
of possible changes.
8. (a) Ammonoid evolution serves well as
example of major modes of evolution. The
episodes of adaptive radiation or
macroevolution, mentioned above were
periods of high productivity of new viable
forms, filling in the ecological niches left
vacuum by earlier forms with their
extinction.
(b) Ammonoids were once considered to
provide good examples of palingenesis,
that is, ontogeny recapitulating phylo-
geny (Palingenesis : recapitulation of
preceding phylogeny in ontogeny of later
advanced forms). But, subsequently it
was found that reverse instances were
more common. Major innovations
occurred cenogenetically; new characters
appearing early in ontogeny (Ceno-
genesis: early appearence of a new
character in ontogeny). In palingenesis,
new characters will be expected to
appear late in ontogeny. Again, these new
Chapter 12 Cephalopoda (Mollusca) 207
features were either incorporated
immediately in the whole of the onto-
geny or progressively in succeeding
generations, proterogenetically (Protero-
genesis: incorporation of a new character
appearing early in the ontogeny in
ancestors, but made stable only in adults
of descendants) (see Factsheet 12.5).
(c) Macroevolution succeeded by cenoge-
netic and proterogenetic incorporation of
new characters and introduction of new
forms with these characters in varied
niches and in the phylogeny, was suc-
ceeded by microevolution (geronto-
morphism). During this phase, the new
forms passed through slower, less spec-
tacular stereotyped evolution, changing
gradually on divergent or parallel lines
through one or more of a series of stan-
dard changes.
This particular mode brought about:
(i) Specialization of evolving forms and
generally, their extinction.
(ii) Repetition of homeomorphy as
parallel or divergent evolution on
similar trends levelled down and
eliminated differences or
distinctions, or repeated some stock
characters.
(iii) What looked like orthogenetic trends,
but commonly not a straight line
evolution with predetermined trends
or directions; instead, each change
was rather an oppurtunistic response
to a change in local conditions. It was
bound by the cause and effect
relationship between adaptation of a
particular form to those conditions
and natural selection acting
thereupon. Selection, in turn, was a
product of scores of factors operating
inside the ecosystem, viz. available
space, predation, competition and
availability of food, etc.
208 Part Two: Major Invertebrate Groups

FACTSHEET 12.5
Palingenesis Versus Cenogenesis-Proterogenesis

PALINGENESIS: Ontogeny recapitulates phylogeny: X, Y, Z a lineage representing phylogeny O1, O2, O3 repre-
sents ontogeny of Z: Palingenesis suggests O1, O2, O3 repeats stages of phylogeny viz. X, Y, Z, respectively.
CENOGENESIS-PROTEROGENESIS: N1, N2, N3 represent ontogeny of Y and O1, O2, O3, represents ontogeny of
Z: a new character appears cenogenetically first in N1 stage of Y, but is not incorporated in mature stages, N2 and N3 of
Y; it is firmlly incorporated proterogenetically at the mature O3 stage of Z ; Y and Z represent members of a lineage, i.e.
phylogeny.

9. Ammonoid evolution may this be viewed as
an example of ‘organic evolution taking place
through organism environment interaction’. At
higher levels, adaptive radiation or macro-
evolution took place with availability of
favourable environment often left vacated with
the extinction of earlier forms and coinciding
with global or regional transgressions. Each
of them created newer ecological niches that
could be explored by newer forms. Succeeding
microevolution involved organism-
environment interaction in which newly
appeared forms adapted themselves to local
environments made available to them or in
which they appeared and were placed in. As
organisms as well as their environment
changed continuously, quickly evolving forms
suffered extinctions with the loss of
equilibrium between the two changes. To
interrupt the interaction between organisms
and its environment, changes in environment
triggered by major phases of regression might
have played important role, since they meant
squeezing and, thus, loss of marine niches.
In this general plan there were, however,
variations in the response of different forms.
The persistant, conservative stocks were
broadly and genarally adapted to presumably
a broader or commoner niche with more
evolutionary plasticity. They survived longer
with a bradytelic (slow) or horotelic (normal)
rates of evolution. With availability of newer
opportunities, they gave rise to multitudes of
newer descendants in a succeeding phase of
radiation. The changes were tachytelic (rapid),
but the resultant forms subsequently followed
slower rates of microevolution.
10. This whole scenario was operative on a group
of organisms with a particular biological
organization fitted to some particular habit and
habitat of living. Thus, ammonoids as
cephalopods, developed on the basis of
molluscan body plan adapted to active pelagic
habit. They had a characteristic growth rate of
body and shell in which shell increment on
one (ventral) side of the aperture was greater
than on the other (dorsal) resulting in coiling
of shells. Increments on the lateral sides were
generally similar to make the coiling bilaterally
symmetrical and planispiral, though in some
they were different, one lateral side growing
more than the other to produce helically or
conispirally coiled shells. Coiling could be
loose (evolute) or tight (involute) and in some
forms shells were uncoiled in parts. Be that as
it may, they could not go beyond such shapes
as to disturb the buoyant floating or swimming
habit. It implied that the shell and its whole
visceral mass in it must be held in equilibrium
position, necessarily in a vertical erect posture
to move efficiently through the fluid medium.
This precondition, in turn, demanded that the
centre of buoyancy and the centre of gravity
be coincident in the case of orthoconic or
straight shells; for coiled shells, the contrary
was true, as the greater distance between the
two centres over which the tension between
them acts, creates wider leverage and, thus,

greater mechanical advantage for stability. In
coiled involute shells, as also in the peculiar,
aberrant heteromorphs (some Triassic ceratitids,
but mostly Upper Cretaceous lytoceratids and
ancyloceratids) the distance is greater
suggesting their efficient swimming habit.
Coiled evolute shells have shorter distance and
they might have been slow-moving, perhaps
benthic animals or deeper water-dwelling
without much necessity of swimming.
Of other characters, size could not have been
too small for the relatively advanced
molluscan body, nor could it be too large for
its habit. Surface ornaments too, were
varying but limited by the necessities of
protection, maintenance of hydrodynamic
equilibrium, etc. Septa, rather sutures, were
possibly to solve hydrostatic problems, that
of maintaining the shell under the
tremendous hydrostatic pressure it would
have been subjected to. Finer frilling
increased the length of contact between the
septum and the shell so that the compressive
force borne per unit length of suture is
reduced. But this too, had limits and the
Cretaceous heteromorphs with four-lobed
primary suture of ‘pseudoceratitic’ type,
distinct from the five-lobed ammonitic suture
of their ancestors, might have avoided
making suture more complex to impart more
mechanical strength to the shell. Rather, they
made it less so and instead developed more
efficient systems of swimming to confront
the problem of higher hydrostatic pressure.
11. Ammonoid evolution, as understood today,
rejects a few long-held ideas:
(a) Palingenesis or Theory of Recapi-
tulation: At one stage of studies, am-
monoids were thought to be providing
good examples of palingenesis. However,
later studies have proved otherwise, where
cenogenesis and proterogenesis prevail.
The issue has been discussed in Factsheet
12.5 and in the discussion above.
Chapter 12 Cephalopoda (Mollusca) 209
(b) Orthogenesis, retrogression or degene-
ration: Orthogenesis or straight line
evolution implies a non-adaptational,
unidirectional progression towards an
inherent or predetermined end product.
Philosophically mechanical, it also
implies an inherent ‘vital’ or driving
force. As a corollary, any deviation was
termed as degeneration or retrogression
developed with racial senescence,
aberration or over specialization, etc.
Heteromorph ammonoids or pseudo-
ceratitic suture were cited as examples;
aberrant shell forms of heteromorphs
were previously interpreted as non-
functional, hence non-adaptational and,
thus, to have lost all evolutionary
momentum or potentiality.
But it was later found that:
(i) Heteromorphs were rare among
ceratitids and more common in one
order Ancyloceratida in Cretaceous.
(ii) Many Cretaceous uncoiled hetero-
morphs gave rise to normal coiled
forms and, thus, must have had this
ability imprinted in their genotypes.
(iii) Functional morphological consi-
derations reveal that such uncoiled
evolute shells were better adapted to
swimming than coiled evolute shells
were, and as stated earlier in this
discussion, their sutural simpli-
fication may be related to these
increased efficiency in function of
swimming. Hence, such hetero-
morphs, too, were definitely
adaptive, neither degenerate nor
retrogressive, rather progressive in
so far as the specific demands of
their conditions in some niches were
concerned.
The above account gives a brief generalized
picture of ammonoid evolution. The essential
aspects of the phenomenon relevant for a beginner
to know and understand it, are only mentioned.
 13
Echinoidea
(Echinodermata)
13.1 Introduction: Characteristics
of the Phylum
Echinodermata, one of the major phyla of inver-
tebrate animals, stands important in both biology
and palaeontology. It has a set of characteristics,
as follows:
1. Echinoderms are eukaryotic metazoan animals.
2. They are exclusively marine. They live
successfully and abundantly in warm, shallow
seas with their hard parts being made of
calcite.
3. The hard part of echinoderm animals is made
up of a large number of elements set in a
complex organization and arrangement.
Hence, it is referred as skeleton and not a shell.
Normally, however, test is the term used for
the purpose.
4. Echinoderm skeleton is internal, since it is
covered by a thin skin-like integument. On this
account, many biologists consider them closest
to the chordates among invertebrates.
However, the skeleton is functionally
external, as most of the softer parts are placed
210
within it and the skeleton acts both as abode
and as protection.
5. Echinoderm skeleton, rather its skin bears
spines on it; hence, the name: echine or spines
on dermis or skin.
6. Echinoderm skeleton or spines are made of a
large number of plates, each a calcite crystal.
With growing body, the skeleton increases in
volume in two ways. New plates are added as
also older plates grow larger with increments
added along the margin. Hence, growth in
echinoderms takes place by both accretion
and addition.
7. Echinoderm plates are clustered and combined
together into a few systems of plates, each
associated with some function or other. Of
these, ambulacra (sing. ambulacrum; referred
henceforth as ambs) are vitally linked with the
water vascular system of the body.
8. Echinoderm body or skeleton are characterized
by a pentameral organization consisting of
basic elements five in number. They are set in
either a radial or a bilateral symmetry. This
pentameral organization is, however, absent
in some early groups and in ‘carpoids’.
 Chapter 13 Echinoidea (Echinodermata) 211

(Subphylum Homalozoa; also called
calcichordates, i.e. like chordates but with
calcitic plates.) (See Factsheet 13.1)
9. The pentameral organization is based on a
water-vascular system placed inside the
skeleton. It consists of several tubular
elements, called canals, that store or act as
passageway for water utilized for varied
functions.
The system consists of a vertical tube, stone
canal that ends in a centrally placed circular
tube called ring canal. Five tubes called radial
canals are given off radially from this ring
canal. Numerous small tubes, called tubefeet
or podia (sing. podium) emerge from the radial
canal. All the parts of the system, canals or
tubefeet, are filled up with sea water and it
can expand or contract its parts. As and when
required, the parts may be expanded or
contracted, sending water into them or taking
it out of them. In echinoids and other groups,
each tubefoot has a sac or pouch (ampulla)
that acts as a storage, though there are no
ampullae in tubefeet in, say, crinozoans, where
they are controlled by the radial canal itself.
Tubefeet are flexible and peep out of the test
(i.e. skeleton) through pores in plates of
ambulacra that lie above the radial canals.
10. Echinoderms are gregarious; they live in large
numbers together; the groups are, however,
different from colonies in that the individuals
are not connected to each other during their
lifetime. They are also different from herds or
such other groups of higher animals as they
lack intercommunication among the
individuals. This gregarious habit imparts
some taphonomic peculiarities to fossil
echinoderm occurrences. In any bed or
horizon, the fossils tend to occur concentrated
at places, the rest of the bed or horizon being
virtually devoid of them.
11. Taphonomy assumes further importance also
on account of calcite-made skeletons of the
animals. It being a stable form of calcium
carbonate, calcite plates add to preservation
potentialities of echinoderm tests. But, at the

FACTSHEET 13.1
Pentameral Organization in Echinoderms

Where:
Characteristic in most groups except some early groups and ‘carpoids’(Subphylum Homalozoa; also called
calcichordates, i.e., like chordates but with calcitic plates).
Why:
Following possibilities are relevant:
1. Suture lines between plates are lines of weakness; even number of columns of plates places opposite
sutures in line with each other; odd number ensures suture against plates, thus reducing weakness; 3 is too
few a number, 5 ideal for maximum resistance. But the fact that collagen fibres hold plates together across
suture, goes against the argument.
2. Pore rows in multiples of 5, are ideal for strength of test and maximum number of tubefeet; pores impart a
‘postage stamp weakness’, an unnecessary larger number of tubefeet may thus weaken the test.
3. Pattern of 5 gives a nearly constant width in any direction with a small number of rays/ambs; the relation D
= n × (n – 3)/2 may also be relevant here, where ‘n’ is the number of sides/apices/vertices and D is the
number of diagonals from one vertex to another. The relationship entails:
for n = 3 4 5 6 7
D= 0 2 5 9 14
212 Part Two: Major Invertebrate Groups
same time, the test is made of numerous small
elements, the plates, joined along sutures that
act as lines of weakness. In addition, good
cleavage of calcite makes the plates and the
test susceptible to mechanical breakage. Thus,
there are instances of limestone beds made
largely of echinoderm remains, viz. disartic-
ulated plates and other parts, in which
complete well-preserved tests are relatively
rare or absent.
12. Echinoderms are benthic in habit, sessile or
vagrant. In fact, the phylum is often subdivided
on the basis of this variation in the living habit.
13.2 Subdivisions and Brief
History
The phylum was classified into two subphyla on
the basis of living habit as well as morphology:
one sessile benthic Pelmatozoa and the other
vagrant Eleutherozoa. Subsequently, this scheme
was revised to erect a few subphyla including
Echinozoa, Asterozoa and Homalozoa which
were vagrant and Crinozoa and Blastozoa sessile.
Though largely retained to date (Clarkson 1998),
in another recent opinion based on cladistic
analysis by Smith (1984b, 1988), the sessile (fixed)
and the vagrant (free-living) division are
considered phylogenetically valid and, thus,
natural. The opinion is, however, yet to gain
universal acceptance (Clarkson 1998).
Echinoderms appeared in Lower Cambrian
and became morphologically so varied that they
were grouped into a few classes under subphylum
Echinozoa. This was also because they were
different from later echinoderms of pentameral
body, thus demanding their grouping into separate
classes. The phylum diversified and reached a
height of radiation at a class level in Ordovician,
though by the number of genera it reached the acme
in Carboniferous only. Of Lower Cambrian genera,
Helicoplacus belonging to the monogeneric class
Helicoplacoidea, is considered the stem group
(Smith 1988) from which were derived two lines.
In one, there were ‘fixed’ ‘pelmatozoans’ through
Lepidocystis (class Lepidocystoidea) and in the
other, were ‘free-living’‘eleutherozoans’ through
Middle Cambrian Cambraster. Thus, though the
Cambrian classes and genera were short-lived and
often localized, they stood important in
understanding echinoderm phylogeny. Subsequent
to Lower Palaeozoic, a number of classes and
subclasses appeared and/or continued till date.
Of them, Echinoidea is the class that assumed
importance in palaeontology. This present chapter
will deal with this class. Factsheet 13.2 provides
some basic information about the other groups.
13.3 Introduction: Echinoidea
Echinoidea is a class of the subphylum Echinozoa
of the phylum Echinodermata. Obviously, it
possesses the above-mentioned characteristics of the
phylum. It is distinguished from other groups of
echinoderms on certain other vital characteristics.
Echinoid tests, made of numerous small
calcitic plates are, in general, hemispheroid or
discoid in shape. As in other echinoderms, there
are five ambs in an echinoid test, each lying above
a radial canal of the water-vascular system and
meridionally placed in the test. Each amb consists
of a series of plates joined together and perforated
or pore-bearing. The pores provide passage for
tubefeet that emerge from the radial canal.
Echinoid tests are further characterized by five
meridional bands of plates, the interambs. Ambs
and interambs occur alternately, the adjacent ones
being held together by tissues. Thus, ambs and
interambs make the major part, called corona, of
a rigid test (flexible in some).
Generally, ambs are narrower than interambs
and their plates are perforate in contrast to
imperforate interamb plates. Since ambs are more
functional providing passage for tubefeet, their
lesser width prevent tubefeet from becoming
unnecessary long. On the contrary, interambs
become wider to fill in the space between the two
narrow ambs. (Figure 13.1).
 Chapter 13 Echinoidea (Echinodermata) 213

FACTSHEET 13.2
Key to Phylum Echinodermata

Subphylum Class Earlier Position GeologicalAge Characters

Echinozoa Echinoidea Eleutherozoa Ordovician-Recent Vagrant, rigid hard

skeleton
Holothuroidea and fixed ambs
Edrioasteroidea Lower Cambrian
to Carboniferous
A few other extinct Palaeozoic classes belong to this subphylum
Asterozoa Stelleroidea Eleutherozoa
Asteroidea Ordovician-Recent Vagrant, flexible
in starfish
Ophiuroidea do Flexible skeleton
in brittle stars
Somasteroidea do
Crinozoa Crinoidea Pelmatozoa Middle Cambrian Sessile, stalked, with
to Recent flexible ambs as in
sealilies
Blastozoa Diploporita Pelmatozoa ?Cambrian Sessile, stalked and
Cystoidea to Devonian fixed ambs
Rhombifera do
Blastoidea Silurian-Permian
Eocrinoidea Lower Cambrian
to Silurian
Homalozoa Ordovician
also grouped as subphylum Calcichordata of phylum Chordata

The test seats on a flatter side, which latter also
has the mouth and is called oral side. The other
side of the test is more or less convex and is known
as aboral. Earlier the lower or oral side was
recognized as ventral and the upper or aboral as
dorsal. Presently, however, oral-aboral terms are
preferred.
Corona, made of ambs and interambs, is a
system of plates. In an echinoid test, there are three
other major and a few minor systems of plates. The
major ones are as follows:
1. Peristomial system or peristome: A
membranous system of plates covering mouth
situated on oral surface during lifetime.
2. Periproctal system or periproct: A similar
membranous system of plates covering anus
during lifetime; it is differently placed in
different genera, being oral, aboral or marginal
between the two in position.
3. Apical system or apical disc (also called
oculogenital system): Lying at or near the
centre of the aboral surface, this system
includes two kinds of plates, ocular plates
connected with the water-vascular system and
genital plates used for reproduction. These will
be treated further in later sections.
It has been mentioned that echinoderms grow
by both accretion and addition. In echinoid tests,
214 Part Two: Major Invertebrate Groups

An
An
1 1
5b

7
2
3 3
4 4
5a

Po
Po
(b)
(a)

Ab
2

6
8
9
4
Or
(c)

Fig. 13.1 Echinoid morphology: regular echinoid with simple ambs and hemispherical test.
(a) Aboral view: ambs, interambs, apical disc, periproct; (b) Oral view: ambs, interambs, peristome;
(c) Lateral profile: maximum height at centre.
Index: 1 Interambs, 2 Madreporite, 3 Ambs, 4 Ambitus,
5(a) Peristome, 5(b) Peristomial membrane, 6 Periproct,
7 Lantern in mouth, 8 Genital plate,
9 Ocular plate (at the end of each amb),
An (Anterior), Po (posterior), Ab (aboral), Or (Oral)

new plates are added to the ambs and interambs
at their ends with the apical disc, whereas
accretion takes place particularly near the margin
between oral and aboral surfaces (this margin is
also called ambitus; there is however, a less
common usage of the term meaning the region of
maximum width of any amb, which generally
coincides with the oral-aboral boundary in what
is known as simple ambs).
For all practical purposes, the morphology of
echinoid tests are more easily comprehended, when
judged on a format as follows:
1. Shape, symmetry, orientation and dimension
of test/tests.

2. Corona, i.e. ambs and interambs.
3. Peristomial and periproctal systems.
4. Apical or oculogenital system.
However, for a real appreciation of echinoid
morphology it is required to be judged on the
background of the adaptive changes in the group.
13.4 Adaptation and Symmetry
Echinoids are marine, benthic animals known
more as sluggish mover or vagrant, though a few
genera may be temporarily fixed. They may be
either epibenthic or endobenthic. The former live
on hard rocky substratum, whereas endobenthics
live either in holes or more commonly, within
sediments, in burrows they make. This difference
in the mode of living brings about morphological
differences too. The most evident of them is the
difference in symmetry. Epibenthic echinoids
have a radial symmetry that may also be the
primary and primitive symmetry for the class, as
can be found from echinoid ontogeny. But
endobenthics which appeared later in echinoid
phylogeny have, in general, a bilateral symmetry,
which is acquired during their ontogeny too. The
bilateral symmetry is, thus, an adaptation, which
does not remain only phenotypic. During the
course of evolution, this trend from radial to
bilateral symmetry developed in many lineages
to assume a permanent status. A brief account of
this major adaptive change in echinoids has
already been given in section 3.2.5. It will be
further referred to in proper context in the
following discussion.
13.5 Water-Vascular System
As an echinoderm, echinoids characteristically
possess a water-vascular system. Like most of the
echinoderms, they also present a pentameral
organization of their body, which includes five
radial canals of the water-vascular system and
corresponding five rays or ambs.
Chapter 13 Echinoidea (Echinodermata) 215
An echinoid body is filled with liquid and
gaseous material in such a way that it looks like a
fluid-filled baloon. It has two main passages for
circulation of fluids. The first is the tubular gut, a
digestive organ, which starts from the oral opening
on the lower oral surface and winds upwards to
end at the anal opening, situated commonly near
the centre of the upper, aboral surface. The second
is the water-vascular system (Figure 13.2).
The system has already been briefly introduced
in section 13.1. Recapitulation along with some
additions may be helpful. The system starts from a
sieve-like plate of the apical system, called
madreporite. It lies at one end of the stone canal,
the vertical tube which runs at the other end into
the centrally placed ring canal. From it, emerges
the five tubular radial canals that run meridionally
immediately below the test or its ambs. They finally
emerge at the aboral end through the plates, known
otherwise as ocular plates of the apical disc. Thus,
the latter, the apical disc, which was earlier known
as serving for vision (with the help of ocular plates)
and reproduction (genital plates emanating
reproductive materials), is now found to be more
closely linked in echinoid life-activities, namely,
those of the water-vascular system.
Numerous small tubes, called tubefeet or podia
emerge from the radial canal. The system, with all
its parts, canals or tubefeet, is filled up with sea
water and it can expand or contract its parts as and
when required, sending water into those parts or
taking it out of them. In echinoids and some other
groups, each tubefoot has a sac or pouch (ampulla)
that acts as a storage. Tubefeet are flexible and
peep out of the test (i.e. skeleton) through pores in
plates of ambulacra that lie above the radial canals.
Each tubefoot bifurcates into two parts, which latter
emerge through a pair of pores in an amb-plate
(or its component part, in the case of compound
amb-plates), only to again coalesce together outside
the plate. This is why pores for tubefeet are
generally present in a pair. Only in few new plates
introduced in the amb near the apical disc or few
old plates near the mouth, there may be a single
pore in an amb-plate or its component part.
216 Part Two: Major Invertebrate Groups

7
1 2
11
3
6 5
4 6
7 8 10
11 14 14
9
13 14
10 (c)

(a)
(b)

a 15
17 18

19
16
(d)
(e)
(f)

Fig. 13.2 Echinoid water-vascular system, spines, tubefeet, pore pair in amb-plates and fascioles.
(a) Water-vascular system and spines, (b) Tubefoot (sucker tubefoot of oral side), (c) Part of (b)
enlarged to show pore pair, (d)-(f) Fascioles
Number Index: 1 Madreporite, 2 Pedicellariae,
3 Ocular plate taking radial canal through, 4 Gut (digestive organ),
5 Gonad (reproductive organ,) 6 Stone canal,
7 Ampulla, 8, 9, 10, 11, 12 Radial canals (or their tips),
13 Perignathic girdle, 14 Pore-pair, a–(Anus)
15 Anal, 16 Sub-anal,
17 Endopetalous, 18 Peripetalous,
19 Lateral, AP (Ambplate), IAP (Interambplate)

The way each tubefoot bifurcates before
passing through the amb-plate and the parts
coalesce thereafter, ensures that the tubefoot does
not slip inside the test when its water is being
withdrawn into ampullae and the tubefoot itself is
shortened.Water enters the water-vascular system
through the madreporite, which sieves out the
particulate matter to prevent clogging. Via stone
canal, water passes on to the radial canals and
therefrom to the tubefeet. Ampullae there store the
water and send it into the tubefeet as required. The
prehensile tubefeet extended further, perform
different functions such as locomotion (for those
on the oral side), food-gathering, respiration and
protection, etc.
The water-vascular system is not preserved in
fossils. But the features that bear its marks are:
1. Multipored madreporite in the apical disc
presenting the entrance; it is further significant
for being invariably located in the right anterior
position; for radial forms, without any other
criterion, madreporite is thus the character that
helps in orienting the test.
2. Pores in amb-plates attesting to the presence
of tubefeet.

13.6 A Format for Morphology
As mentioned earlier, there are four major and a
few minor systems of plates in echinoid tests. The
format is best framed on their basis. Details may
be studied as follows:
1. Shape, symmetry, orientation and dimension
of test/tests; shape includes geometry of the
ambitus and the curvature (concave/convex/
flat) of oral and aboral surfaces; symmetry may
either be radial or bilateral; fixing orientation
involves marking anterior-posterior that
defines the length and oral-aboral
(dorsoventral) that defines the height.
2. Corona:
(a) Ambs: Similarities-dissimilarities;
structural characteristics, viz. simple,
subpetaloid, petaloid; appearance: raised,
depressed or flat in relation to interambs;
pore-zones and pores: number, shape,
position and conjugation; width of pore-
zone, etc.
(b) Interambs: Similarities-dissimilarities;
shape and size.
(c) Tubercles: Varieties and their nature;
fascioles and their types and positions.
3. Peristomial and periproctal systems, their
position, alignment and outline geometry; with
peristome, floscelle, lantern, food groove, etc.
Normally, these two systems are not preserved
in fossils; they are represented by corresponding
openings. Mouth and anus are really smaller
openings included within the larger openings
of peristome and periproct, respectively.
4. Apical/oculogenital disc/system: position,
shape; number of plates, their position;
madreporite.
13.7 Shape-Size-Symmetry, etc.
of Test
Echinoid tests show either radial or bilateral
symmetry. Though generally and typically
Chapter 13 Echinoidea (Echinodermata) 217
hemispherical or flatter discoid, they show quite a
varied type of shapes. (Figures 13.3, 13.4, 13.5).
It is controlled by several parameters. Of them, the
geometrical outline of the ambitus, i.e. the margin
between aboral and oral surfaces is an important
one. In ‘regular’ echinoids, the ambitus is circular;
but in ‘bilateral’ forms it varies. At ordinal levels,
in cassiduloids and holectypoids, ambitus is
circular, elliptical or oval, while clypeasteroids
have a pentagonal to circular type. In spatangoids,
ambitus is heart-shaped, with a characteristic
reentrant at the anterior end and an abrupt straight
or truncated margin at the posterior end. The
margin between oral and aboral surfaces is
generally smoothly rounded; in such cases, ambitus
is the greatest horizontal circumference.
While ambitus gives the horizontal component
of the shape of echinoid test, lateral profile giving
the curvature of oral and aboral surfaces determines
the shape in vertical section. In regular echinoids,
any vertical section serves for the profile, but in
irregular echinoids the lateral profile is drawn
along the symmetry plane. Aboral surface is always
convex, high, moderate or low, being largely
characteristic of different genera and species.
Besides, the position of maximum height on this
surface varies. In radial, i.e. ‘regular’ forms it is at
the centre of the aboral surface. But in bilaterally
symmetrical ‘irregular’ forms, it varies at ordinal
to lower taxonomic levels. In Clypeaster or other
genera, the surface is convex in the central part
and flattened as a rim along the margin, imparting
a hat-like look to the test. In spatangoids, the
convex aboral surface tends to have maximum
height at or towards the posterior margin. The test
is then sloping towards anterior. The oral surface,
on the other hand, may be convex, concave or flat,
generally characteristic at generic level.
Like symmetry, the shape of the tests is also
dependent on the mode of living of echinoids.
Radial symmetry, along with circular ambitus,
hemispherical or disc-shaped tests having
maximum height at the centre, characterizes
epibenthic echinoids, living on hard substrates and
facing similar ambience all around.
218 Part Two: Major Invertebrate Groups

m
a
a (iii)

(i) (ii) (a)

ant
a
a
m
(b)
post
a

a
m
(c)

a
m
(d) a

ant post

Fig. 13.3 Echinoid tests.

(i) ambitus in aboral view, (ii) ambitus in oral view, (iii) lateral profile
Characters ambitus profile ambs periproct
(a) circular hat-shaped petaloid inframarginal
(b) heart-shaped anteriorly sloping paired petaloid marginal
(c) pentagonal convexo-flat simple (not shown) supramarginal
(d) elliptical/oval biconvex subpetaloid inframarginal
m (Mouth/peristome), a (Anus/periproct), ant (Anterior), post (Posterior)
 Chapter 13 Echinoidea (Echinodermata) 219

(a) (b)

(c)

Fig. 13.4 Regular echinoids.

(a) Oral view of a recent test; (b) Aboral view of the same test; (c) Fossil of regular echinoid.
Note: Simple ambs and multiserial pores in (a) and (b); uniserial pores and flexuous ambs in (c).
220 Part Two: Major Invertebrate Groups

(a) (b)

(d) (e)

(c)

(f) (g)

Fig. 13.5 Irregular echinoids.

(a), (d), (g) Oral views; (b), (c), (e), (f) Aboral views
(a) Periproct at the bottom of the figure (c); Uniserial inner circular, outer slit-like pores in
(d) Typical tubercles (e) and (f) Petaloid paired ambs (g) Distinct carina.

Endobenthic organisms, in their turn, face
different environments in front (the dead end of
the burrow) and at back (sediment water interface
and the water mass to derive oxygen and nutrients
from, as well as free space to discharge wastes),
but similar environments on the sides. So, they
assume bilateral symmetry, defined by elliptical,
oval, pentagonal or heart-shaped ambitus of tests.
Burrowing is facilitated by an anteriorly sloping
test and so spatangoids, the typical burrowers, have
the maximum height at or towards, the posterior
end. Clypeaster tends to bury its test beneath a thin
veneer of sediments, with the apical disc just
peeping out. Centrally elevated hat-like test with a
thin rim around, suits this mode.
Orienting echinoid tests involve fixing anterior-
posterior. A set of criteria may be used as shown in
Factsheet 13.3. In radial forms, madreporite is the
only feature that helps in marking right anterior
position. Dimensions of echinoid tests are termed
length (maximum distance between anterior and
posterior ends), breadth (at right angles to length in
horizontal plane) and the height measured in lateral
profile at right angles to length.
13.8 Corona: Ambs and
Interambs
Corona is the main part of the test made of five
ambs and five interambs placed alternately and
running meridionally from the apical disc to the
peristome. Ambs are definitely more important than
interambs in life activities and for taxonomic
purposes. Interambs primarily fill up the space
between two ambs to complete the rigid test.
Ambs make one of the most important
characteristic that reflects symmetry of echinoid
tests. Thus, regular echinoids with radial symmetry,
have five similar ambs, of which any two adjacent
ones subtend an angle of 72° at the centre. Bilateral
symmetry disturbs this similarity of ambs. Five
ambs are grouped into two pairs and one unpaired.
Ambs being functionally important and anterior
Chapter 13 Echinoidea (Echinodermata) 221
being an uniquely significant direction for the
animal, the unique unpaired amb marks the
anterior. The rest are included in an anterolateral
pair and a posterolateral one. The shape, size,
alignment, position and other characters of these
ambs reflect the symmetry.
Variation in ambs, characteristic for genera and
species, mainly depend on two factors: number,
arrangement, etc. of amb-plates and characteristics
of their pores for tubefeet. As a part of echinoid
test, ambs are made of numerous calcitic plates
(Figure 13.6). Barring a few Palaeozoic groups,
most groups and their genera have their amb-plates
arranged in two columns. The zigzag perradial
suture between the two columns mark the line
along which the plates are imbricated (boundary
between ambs and interambs is called adradial
suture). It has already been stated that ambs run
meridionally above the radial canals of the water
vascular system and that each amb-plate (excepting
a few adapical and adoral plates) bear a pair of
pores to make passage for the tubefoot. Alternate
position of plates along the perradial suture appears
to meet the two ends. Junctions between adjacent
plates of one column are lines of weakness; their
alternate position ensures that such lines of one
column are shifted laterally to end against a plate
of the other column. It, thus, reduces weakness to
an extent. Secondly, since tubefeet are given off
on both sides of the radial canal, their alternate
position, as against opposite one, provides more
space for them to emerge.
Columns of amb-plates may extend upto 20
in number in the few Palaeozoic groups referred
above, which are also significantly different in
many other respects (even in the absence of
pentameral arrangement) from later echinoids.
Amb plates are called simple, when it is a
single unit of calcite crystal each. It may otherwise
be a compound of various types, consisting of two
or more component parts (smaller ones called
demiplates). Each plate of simple amb or each
component part of a compound amb carries a pair
of pores (Figure 13.6). These occur near the two
222 Part Two: Major Invertebrate Groups

FACTSHEET 13.3
Orienting Echinoid Tests

Feature Marks Found in

Ambitus: Heart-shaped Anterior Spatangoida
Lateral profile: Maximum height marginal Posterior Spatangoida
Ambs: Unpaired amb Anterior ‘Irregular’
Periporct: Exocyclic position Posterior ‘Irregular’
Peristome: Shifted from centre of oral surface Anterior ‘Irregular’(Spatangoida)
Apical disc: (a) Madreporite Right anterior In all echinoids
(b) Four gonopore, two on either side
of symmetry plane; fifth absent Posterior Spatangoida

(ii)
(i) (iii)
(a)

(i) (ii)
(iii)
(b)

Scrobicular
tubercles

(i) (ii)
(c)

Fig. 13.6 Echinoid test : plates, pores, tubercles.

(a) Amb-plates; (i) simple; (ii) diademoid compound (lowest member small); (iii) echinoid compound
(lowest member large), (b) Pore series; (i) uniserial Palaeechinus; (ii) biserial Lovenechinus;
(iii) multiserial Melonechinus; (c) Tubercles; (i) one primary tubercle surrounded by a ring of scrobicular
tubercles and other granules on one single plate (in plan); (ii) same in elevation.
Note: One compound plate has one primary tubercle: figures diagrammatic.

adradial sutures along a zone called pore-zone;
its width is often characteristic of genera; thus, in
Hemiaster or Micraster, the zone is wide, whereas
in Echiolampas or Cidaris it is narrow.
As said before, pores for tubefeet occur in
pairs. In a few newly introduced adapical plates
(near the apical disc), there may be a single pore
in a plate; also in a few senile plates near the
peristome, tubefeet may become functionless and
their pores plugged or lost. Similarly, in genera
with petaloid ambs, portions beyond the petals have
their pores plugged, whereas those within the petal
area are highly developed and specialized for
respiration. Normally, pores are circular, more so
in regular echinoids (e.g. Cidaris, Temnopleurus,
etc.). But in many irregular echinoids one pore of
the pair (the outer one, i.e. towards adradial suture)
or both may be elongated and slit-like;
Echinolampas, Clypeaster provide examples of the
‘inner circular, outer slit-like pores’, whereas
Hemiaster, Schizaster have both pores slit-like in
paired ambs. Pores may remain separate and so
non-conjugate or be conjugated by a groove
(e.g. in Echinolampas, etc.) or having been placed
within a common depression called peripodium
(e.g. in Micraster, Breynia).
In one pore-zone, the pair of pores may be
stacked in one series (i.e. actually two lines of
pores called uniserial arrangement). This is
particularly found in ambs with simple plates, as
in irregular echinoids. But in many regular
echinoids with compound amb-plates, the pairs
may be laterally shifted giving way to two series
of pairs in one pore-zone (i.e. actually four lines
of pores; biserial pores) or more (multiserial)
(Figures 13.4, 13.6).
Tubefeet find various uses in echinoids. To
know them in correct perspective, we need to deal
with certain important variations in ambs
themselves. Structurally ambs may be simple or
petaloid, with an intermediate subpetaloid type
between them. Simple ambs are found mainly in
radially symmetrical regular echinoids. In this type,
each amb, after starting from the margin of the
Chapter 13 Echinoidea (Echinodermata) 223
apical disc, gradually widens to reach the
maximum width at the ambitus; passing over to
the oral surface, it then gradually narrows down
towards peristome. More significantly, simple
ambs are characterized by the uniform nature of
their pores, barring minor variations near the
adapical and adoral ends, as indicated earlier.
Generally, pores in simple ambs are circular, non-
conjugate. Ambs which contain compound plates
are invariably simple.
On the other hand, particularly in spatangoid
and clypeasteroid irregular echinoids, a portion of
each amb on the aboral surface is characterized by
numerous, densely packed amb-plates with one or
two slit-like pores in each pair. In the remaining
portions of the amb, pores are completely plugged
off. The former portion, an enclosed area may be
depressed in relation to adjacent interamb, but is
always associated with specialized and
characteristic pore-pattern. It has an appearance
of a petal and, hence, such an amb is called a
petaloid amb.
A third type, called subpetaloid, is found in
some irregular echinoids, more particularly in
cassiduloid-holectypoid groups. Here the aboral
petaloid portion of an amb is not fully formed into a
totally enclosed area of specialized tubefeet; there
is generally one slit-like and one circular pores in
that part and this pore character changes gradually
on the aboral surface towards its margin, i.e. ambitus.
As we mentioned earlier, radial echinoids are
mainly epibenthics, while endobenthics have
bilateral symmetry. The latter have to live under
sediments and are, thus, provided with limited
oxygen supply. It calls for their adopting special
device to acquire adequate oxygen. Slit-like pores
for appressed respiratory tubefeet in ambulacra
serve the purpose. They develop on aboral portion
of ambs, particularly in the four anterolateral and
posterolateral petals. The unpaired amb, diving into
the burrow ahead of the paired ambs, is
functionally different from them and is often
inactive and so lacking respiratory tubefeet or their
slit-like pores. It may be added here that the flat
224 Part Two: Major Invertebrate Groups

surface of these tubefeet provides advantage for
gaseous exchange involved in respiration (also see
Factsheet 13.4). Since echinoid metabolic rate and
oxygen consumption increases with temperature,
warm water forms have longer petals, needing
more efficient gaseous exchange systems than
those from cold waters.
Radial forms do not require these specialized
tubefeet for respiration. But their epibenthic mode
of living demands more protective means and more
efficient current-producing movement of tubefeet.
This seems to have been achieved with the help of
increasing number of tubefeet for a given size
(diameter and height) of the test through
development of compound plates bearing
multiserial tubefeet or their pores.
13.9 Interamb, Tubercles and
Fascioles
Alternately lying with ambs, interambs are the
second components of corona. They are generally
larger than ambs in area and size of plates.
The latter are, however, imperforate, since they
have nothing like tubefeet to pass through them.
They are also less varied for the same reason. In
regular echinoids, the five interambs are similar
in shape and size; but in irregular echinoids they
are divided into two pairs and one posterior
unpaired.
The major variations in interamb morphology
pertain to the presence and characters of tubercles
(Figure 13.6). The latter are also present on ambs,
but there they are less varied and so unimportant.
Echinoid tests bear spines during lifetime of
the animal. These spines serving different functions
(Factsheet 13.5; Figure 13.3) are held to their bases
with the help of tissues, which decay after death,
leaving the spines disarticulated from their base
on the test.
Tubercles serve for the base of spines. Large
or small, a tubercle is like a mound with a dome-
shaped head that bears a perforation that makes
room for the tissue to emerge. The spine that sits
on a tubercle has a concave base with a
corresponding perforation for the tissue that holds
the two, spine and tubercle together. The couple
acts on a ball and socket arrangement as we find
with vertebrate bones. It ensures rotatory movement
of the spines. Different parts of tubercles and spines
are shown in Figure 13.6.

FACTSHEET 13.4
Gaseous Exchange and Respiratory Tubefeet

Transport of oxygen to and carbon dioxide from internal tissue poses problems to any globular animal because
of its small surface area to volume ratio. Further, echinoids lack respiratory pigments in coelomic fluid. Thus,
for efficient gaseous exchange there were a few morphological adaptations in echinoids, most of which (3, 4,
5 below) were more with the endobenthic forms facing a lesser supply of oxygen:
1. One-way circulation system between tubefeet-ampullae and radial canal, in Silurian from single-pored ambs
in Ordovician. It included:
(a) a valve at the junction between the ampullae and the radial canal, which prevented water from flowing
back into the radial canal; and
(b) pore-pair preventing tubefoot to slip back into the test.
2. Partition to separate oxygenated and deoxygenated currents within the tubefeet and ampullae.
3. Thinner walled, suckerless aboral tubefeet for respiration.
4. Since mid-Mesozoic, more prominently in Tertiary, there was a trend to increase in surface area relative to
volume through flattened and elongate respiratory tubefeet (ending in slit-like pores).
5. A respiratory shaft reaching the surface and ciliated spines of facsioles to create necessary currents in
spatangoids.

FACTSHEET 13.5
Multiple Use of Echinoid Spines
1. Protection of tests of:
(a) of tubefeet
(b) of oral cavity
2. Acquiring food with the help of sticky mucus
3. Preventing sediments from getting into mouth
4. Carrying food into mouth
5. Helping locomotion
6. Making burrow
In respect of size, position and function,
tubercles are of three main types, viz. primary, the
largest, secondary the intermediate and miliary the
smallest, often microscopic.
Primary tubercles have the following
characters:
1. Commonly found in regular echinoids.
2. Fewer in number, each interamb (or amb) plate
bearing one primary tubercle.
3. Add strength to the plates themselves.
4. Present in ambs, where plates are compound;
the tubercle then helps hold the component
parts together.
5. Occur in rows on the test.
6. Those on aboral surface hold spines used for
protection; spines on large tubercles on oral
surface hold the test above ground and help in
locomotion.
Secondary tubercles bear the following
attributes:
1. Smaller than primary tubercles, these are,
however, visible to naked eye.
2. Equally important in regular and irregular
echinoids.
3. Include two kinds. The first, called granules,
are numerous, distributed evenly all over the
test or over a large area; generally larger on
oral surface than their counterparts on aboral
surface; found typically in cassiduloids and
clypeasteroids.
Chapter 13 Echinoidea (Echinodermata) 225
4. The second type of secondary tubercles are
fewer in number, occur in a raised scrobicular
ring around a primary tubercle; so they are
referred as scrobicular tubercles; found in
regular forms with primary tubercles.
5. Perform similar functions as the primary
tubercles.
The third type, i.e. the miliary tubercles cannot
be located by naked eye or under a hand lens. They
have the following characteristics:
1. These are innumerable.
2. These occur in narrow depressed bands or
channels called fascioles, found mostly in
spatangoids (Figure 13.2).
3. Miliary tubercles in any fasciole moves
continuously during lifetime to create a flow
of water along the channels of fascioles.
4. Fascioles present in different parts of the test
are given different names; primarily they
supply fresh water for respiration and food
particles or for preventing particulate matter
(sand and mud particles) and foul water from
entering the vital areas or openings. Thus,
fascioles may be
(a) Peripetalous: Surrounding the petals;
(b) Endopetalous: Within the petals;
(c) Marginal: Around ambitus;
(d) Lateral: Occurring diagonally from the
peripetalous to the marginal fasciole; and
(e) Anal and Subanal: The former
crescentic around the anus and the latter
ring-like below the anus.
Variation in tubercles is also linked with the
basic adaptation of these animals. Epibenthic
echinoids living on the substrate are largely
unprotected; protection of test and locomotion are
the two main functions of tubercles in these forms.
On the other hand, protection becomes less
important for those endobenthic echinoids that are
partially covered by sediments. So, they have more
granules and less or primary tubercles. For
burrowers, the principal needs are supplying fresh
226 Part Two: Major Invertebrate Groups
water for respiration and food, etc. and removing
the wastes, ensuring that the two do not mix up.
There is also the need for preventing particulate
matters from clogging the test openings. Thus,
spatangoids, the typical burrowers characteri-
stically have miliary tubercles.
13.10 Peristomial System,
Aristotle’s Lantern
Peristomial system or peristome is among the three
major systems other than the corona. It is a system
of plates around mouth and with a membranous
cover during lifetime. It is always situated on oral
or lower surface, central in position in regular
echinoids and often eccentric, shifted towards
anterior in irregular echinoids. In fossils, it is often
not preserved; however, position, alignment and
geometrical outline of the peristomial opening,
including the mouth are often characteristic of
genera and species. Besides these general features,
peristome, ambs and interambs around it develop
certain specialized structures or systems related to
feeding and food-gathering processes of the animal.
Particularly in regular echinoids, there is a
structure called Aristotle’s Lantern, which is made
up of about 40 plates and is used for various
purposes. There is also a system of continuous or
discontinuous ring of internal processes around
peristomial opening, called the Perignathic girdle,
that acts for attachment of muscles to support and
control the lantern. The lantern has five component
units, each having a maximum of eight plates: a
pair of pyramid or hemi-pyramid; a pair of
epiphyses, a tooth, a rotula and a compass divided
into two. There are as many as 60 muscles to operate
the lantern and its different parts. They help throw
out the whole structure through the peristomial
opening and open it like a dredging machine to
grab/dredge materials with food from the substrate.
It then closes and retracts inside the test.
In cassiduloids, in particular, another system
around peristome helps in food-gathering. It looks
like a flower and, hence, is called ‘floscelle’. It
consists of five phyllodes and five bourrelets. The
former are broadened ambulacral areas that have
a dense concentration of tubefeet. Its small tubefeet
pick up small particles and larger number of them
can process larger amount of sediments in a more
efficient way. These together allow the animals to
live on less organically rich sediments. Tubefeet
are also suckered that prevent the organism to live
on fine sediments. Bourrelets in interambulacral
areas are provided with spines to help push food
materials into the mouth.
Large clypeasteroids developed food grooves
around the mouth. These were narrow channels in
ambulacral position leading into the mouth. The
grooves are provided with numerous cilia or hair-
like processes that produce current or mucous
strings which transport food material. Nearest
tubefeet simply place food particles near the food
groove. As a result, they are freed from creating
water current to carry the particles and can, thus,
capture more particles. With branching of food
grooves in some genera (e.g. Encope or Scutella),
the efficiency is further increased with more
tubefeet being involved in the process. This
technique appears to have developed in echinoids
living in sandy sediment acting as an efficient
means for sieving surface sediments.
Spatangoids also have their typical
developments around mouth. They, thus, develop
labrum, a more or less enlarged and modified lip-
like plate at the margin of the peristome in the
position of the posterior unpaired interamb. There
is also the plastron, a more or less inflated and
enlarged part of the same unpaired interamb, with
characteristic absence of tubercles. But the most
significant development in spatangoids in
connection with food-gathering strategy is the
deeply sunken unpaired amb on the aboral surface
(e.g. in Schizaster). Deep burrowing echinoids
naturally face problem of finding enough nutrients,
as the latter decreases downwards away from the
surface. These animals develop the sunken
unpaired amb provided with specialized spines for

protection and other purposes. They channelize
nutrient particulates within the amb to reach onto
the mouth on the lower surface and prevent large
particles from the burrow walls to clog or
contaminate the flow.
13.11 Periproct and Apical Disc
Like peristome, periproct of echinoids is a similar
membranous system of plates covering the anus
during lifetime. In regular echinoids, periproct lies
at the centre of aboral surface, diametrically
opposite to peristome and included within the
apical disc. This position is called endocyclic. On
the other hand, in ‘irregular’ echinoids periproct
comes out of the apical disc towards posterior,
giving rise to exocyclic position of periproct.
Outside the oculogenital ring, periproct may lie
on the aboral surface on the posterior unpaired
interamb (supramarginal in position as in
Stygmatopygus) or may be situated right on the
ambitus (marginal; as in Hemiaster at the top of
the posterior truncation) or may move onto the
lower oral surface (inframarginal; as in
Echinolampas or Clypeaster). The position,
alignment and outline geometry of periproct are
diagnostic of genera or species.
Apical system lies at or near the centre of the
aboral surface in regular and irregular echinoids,
respectively. In the former and some primitive
irregular forms, it includes the periproct within its
confine (endocyclic); in the latter periproct moves
out of the apical disc (exocyclic). This system
includes two kinds of plates, ocular plates
connected with the water-vascular system and
genital plates used for reproduction. Generally, in
regular forms, they are arranged in a single
continuous ring of ocular and genital plates
alternately placed, the former in amb positions and
the latter in interamb positions. This is insert/
monocyclic condition of the apical disc. But in
many regular and all irregular echinoids, the
condition is exert/dicyclic, where there are two
Chapter 13 Echinoidea (Echinodermata) 227
concentric rings the outer a discontinuous ring of
ocular plates and the inner a continuous ring of
larger genital plates.
All ocular and all but one genital plates have
a pore each. The right anteriorly placed genital
plate has many pores giving it a sieve-like
appearance. As mentioned earlier, this is the
madreporite, a vital part of the water-vascular
system. Besides, it helps in orienting radial tests
lacking any other relevant criterion. Pores in genital
plates are called gonopores, which are larger in
female tests than in male ones. In irregular
echinoids, adjacent genital plates are often fused
together (characteristically for genera or species)
and there may, thus, be four to even one single
genital plate.
Ocular plates were earlier believed to be linked
with the vision of the animals. But since, radial
canals emerge through the pores in ocular plates,
the latter are now interpreted as vitally linked to
the water-vascular system of the animal.
13.12 Mode of Living and the
Plate Systems
It will be clear from the discussions above that
morphology of echinoid tests, including its
different systems are intimately linked up with the
basic adaptative change in the class, i.e. adaptation
from epibenthic to endobenthic mode of living.
(Also see Factsheet 13.6; Figures 13.4, 13.5) The
picture becomes clearer when viewed in relation
to the feeding processes or habits of these
organisms.
Echinoids are deposit-feeders feeding on
organic material acquired from the substrate.
Perhaps they were primarily detritus-feeders, which
grabbed sediments from the surface and brought
out the food material from them. With evolution,
the process changed regular echinoids collected
food largely by ‘scraping’ with the help of their
lantern. On the other hand, endobenthic irregular
echinoids living on softer substrates, took help of
228 Part Two: Major Invertebrate Groups

FACTSHEET 13.6
Echinoid Mode of Living and Some Major Morphological Variations

Epibenthic ‘Regular’ Endobenthic ‘Irregular’

SYMMETRY Radial Bilateral

AMBITUS Circular Circular/elliptical/oval/pentagonal/heart-shaped
AMB
General Similar Dissimilar
Simple Simple/subpetaloid/petaloid
Amb-plate Simple/compound Simple/compound
Amb pore-pairs Both circular Both circular
One circular, other slit-like
Both slit-like
Rows Uniserial/biserial Uniserial/biserial
Multiserial
Pore-zone Narrow/broad Narrow/ broad
PERISTOME Central Central to subcentral (anterior)
PERIPROCT Within apical disc Outside apical disc
Central (aboral) Central/subcentral/posterior (aboral)
Posterior on the ambitus
Posterior on oral surface
APICAL DISC Endocyclic (periproct within the disc) Exocyclic (periproct outside the disc)
Generally insert/monocyclic Generally exsert/dicyclic (two concentric rings,
outer of ocular, inner of genital plates)

swallowing bulk material to choose organic amount
from that. Their excreta, thus, contained more
sediments and was larger in volume. Regular, and
some primitive irregular echinoids, epibenthic or
partially endobenthic, had or have endocyclic
periproct. But later endobenthic forms could not
accommodate with that kind of position for
periproct, on account of their difficulty mentioned
above, i.e. their larger and different kind of
excreted materials. Chances of pollution of water
for the vascular system entering through
madreporite increased in these sediment
swallowing echinoids. This was solved with the
exocyclic condition of periproct, which then moves
out of the apical disc and progressively shifts
towards the posterior.
There were changes also with respect to
peristome. Regular echinoids with functional
lantern needed larger and simpler peristomial
opening to provide for free passage for the latter.
In irregular echinoids peristome becomes smaller;
in those irregular forms still possessing lantern,
the latter is not thrown out of the test. It lies inside
and instead of scraping food material, it is used
mainly for crushing them or for mastication.
With evolution, food gathering process in
regular echinoids involved changes in the lantern
and perignathic girdle. In irregular forms, on the
other hand, there were development of floscelle,
food grooves and various features in spatangoids
as discussed in section 13.10.
The summary of wide morphological variation
in echinoids, dwelt upon, proves in outline, how it
is related to the adaptive changes in this group of
animals. The basic adaptive change from
epibenthic to endobenthic mode decided the course

of evolution of echinoids. But at lower taxonomic
levels, similar adaptations have recurred once and
often in different lineages, producing
morphological convergence. That is why, regular
and irregular groups are now considered
polyphyletic by many authors.
13.13 Brief Phylogeny
Echinoids appeared in Upper Ordovician;
Bothriocidarids that made their appearance in
Middle Ordovician are no longer considered as
echinoids. They are held as phylogenetically more
related to holothuroids.
Palaeozoic echinoids found an adaptive
radiation in Carboniferous. The orders of that time
were characterized by combinations of imbricated
plate arrangement (instead of sutured one), number
of ambulacral columns, tooth structure in lantern,
etc. very much different from those found in later
forms. Perhaps only one genus, Miocidaris,
survived the end-Permian extinction.
Palaeozoic orders are included in the subclass
Perischoechinoidea. Later orders include regular
Cidaroida (and Miocidaroida) of the same subclass
and those of the other subclass Euechinoidea, both
regular and irregular (eognathostomates,
neognathostomates and atelostomates) echinoids.
Irregular echinoids appeared in early Jurassic and
diversified later to become about 47 per cent of
living species.
Fossil record of the group is, however,
modified by the preservation potential. It depends
on (i) rigidity of tests and (ii) type of environment
inhabited. Imbricate plated tests have lesser chances
of preservation and the orders that had such tests,
extinct or still living, have poor fossil record. An
order of irregular echinoids (holasteroids) being
deep-sea forms also have poor fossil record.
Epibenthic regular echinoids living on substrates
without finer sediments, and subject to current and
scavenging actions also have lesser chance of
preservation. Irregular echinoids being deposit-
Chapter 13 Echinoidea (Echinodermata) 229
feeders or sediment swallowers living on loose
sediments in areas of active sedimentation are best
suited for preservation. They are also abundantly
represented in Tertiary column. In India too,
echinoids are significantly represented only in
Tertiary. A few examples are Temnopleurus in
Tertiary of Pakistan and Kachchh, Cidaris from
Cretaceous-Tertiary of South and Central India,
Pakistan and Kachchh; Clypeaster, Schizaster,
Echinolampas, Breynia from Tertiary of Sind–
Baluchistan and Kachchh. A few age-specific species
are also known; Stygmatopygus elatus of Upper
Cretaceous is known from Ariyalur Formation of
Cauvery basin and from Assam; Breynia carinata
of Lower Miocene from Sind and Kachchh.
13.14 Appendix: Echinoidea
13.14.1 Characters of a few genera and
suggesting functional
morphology on them
Breynia: Bilateral symmetry; heart-shaped
ambitus; anteriorly sloping lateral
profile with maximum height
towards posterior end and
posteriorly located marginal
periproct suggest infaunal
burrowing habit. Petaloid-paired
ambs and slit-like pores (or rather
pear-shaped outer one of the
uniserial conjugate pores placed in
peripodium in petals of paired ambs;
the inner one is circular) suggest the
presence of respiratory tubefeet for
infaunal living. Depressed unpaired
amb, presence of carina, labrum,
plastron and typically arranged
tubercles on oral surface point to
specialized food-gathering adap-
tation inside the burrow and multiple
function of tubercles for locomotion
including burrowing.
230 Part Two: Major Invertebrate Groups
Micraster: Bilateral symmetry; heart-shaped
ambitus; gently anteriorly sloping
and steeply posteriorly sloping
lateral profile and posteriorly located
marginal periproct suggest infaunal
burrowing habit. Petaloid ambs and
conjugate pores (albeit circular in all
ambs, placed in peripodium in petals
of paired ambs) suggest the presence
of respiratory tubefeet for infaunal
living. Strongly depressed paired
ambs and depressed unpaired amb
indicate deep burrowing habit. The
presence of labrum on oral surface
points to specialized food-gathering
adaptation inside the burrow. Other
parts like carina, plastron, etc. are not
well-developed.
Schizaster: Bilateral symmetry; heart-shaped
ambitus; strongly wedge-shaped test
with anteriorly sloping lateral
profile, maximum height towards
posterior end and posteriorly located
marginal periproct suggest infaunal
burrowing habit. Highly depressed
petaloid paired ambs, deeply incised
unpaired amb and slit-like pores in
petals of paired ambs (though pores
are circular in unpaired) also suggest
infaunal habit with respiratory
tubefeet for deep infaunal living.
(Specialized food-gathering (water
channelizing) processes such as
carina, fascioles, plastron, labrum
and specialized tubercles typical of
spatangoids including the genus may
not be always well-preserved.)
Clypeaster: Bilateral symmetry and posteriorly
located inframarginal periproct
indicate infaunal habit. Centrally
elevated hat-like shape points to
living below a thin veneer of
sediments; petaloid ambs and slit-
like outer pores (of the uniserial
pores with inner ones being circular)
attest to the presence of respiratory
tubefeet for infaunal living; petals
flush with interambs and similar in
all the five ambs suggest shallow
infaunal habit; food grooves and
even-sized granules on oral surface
speak of specialized food-gathering,
digging and locomotory processes
for infaunal habit.
Eurhodia: Bilateral symmetry defined by
differentiated though simple am-
bulacra and posteriorly narrowing
triangular ambitus, posteriorly
located inframarginal periproct
indicate the irregular status of the
genus and the infaunal habit. But
simple nature of ambs, echinoid-type
of compound plates with only
circular pores and high dome shape
of the test suggest weak/temporary
infaunal living.
Cidaris: Typical regular echinoid of epifaunal
habit; radial symmetry, high dome-
shaped test with circular ambitus,
undifferentiated simple ambs with
circular pores and periproct placed
at the centre of the aboral surface
diametrically opposite to the
peristome confirm epifaunal living.
 14
Trilobita (Arthropoda)

14.1 Introduction: Arthropoda living forms belong to three major subdivisions

Arthropoda is a very well-known group of
invertebrates, being the largest in number of genera
and species so far known. Formerly known as a
phylum, the group is now recognized as a
polyphyletic superphylum (Factsheet 14.1). Its
(formerly known as classes, now ranked as separate
phyla), viz. Uniramia (insects, etc.), Chelicerata
(spiders, scorpions and mites) and Crustacea (crabs,
lobsters and shrimps), whereas extinct forms include
trilobites and eurypterids (both of Palaeozoic age,
the latter including giant water scorpions).

FACTSHEET 14.1
Arthropoda and its Major Divisions

Taxon Characteristics Age Examples Earlier position

Arthropoda (S) Polyphyletic, segmented Cambrian Phylum

body, articulated appendages -Recent
Uniramia (P) Marine-terrestrial, undivided, do Centipedes, Insecta and
uniramous appendages insects other classes
Hexapoda (C), etc.
Crustacea (P) Marine, also terrestrial do Lobsters, crabs Class
Ostracoda (C) Marine to terrestrial, important (Subclass)
in micropalaeontology
Chelicerata (P) Marine to land; two do Scorpions, Consists of some
main segnments spiders, earlier classes
horseshoe,
crabs
Trilobita (C) Marine; only fossil, Cambrian Class
three lobes -Permian
S: Superphylum, P: Phylum, C: Class
231
232 Part Two: Major Invertebrate Groups
Arthropoda is not only numerically strong, it
shows a very wide range of adaptations. Yet it is
characterized by a set of general attributes such as:
(a) bilateral symmetry and segmentation of the
body and its exoskeleton (many also have an
internal skeleton, i.e. endoskeleton), the latter
providing firm anchorage for muscles and other soft
parts that help operate the appendages efficiently;
(b) jointed appendages (from which in Greek, the
name Arthropoda is derived), including jaw
structures which are highly varied in their number,
arrangement and morphology, so much so that they
may be successfully used in taxonomy; (c) hard
jaw structure or mandible to grind, crush or bite,
leading to a fairly advanced mode of feeding
through predation; (d) growth by moulting or
ecdysis, i.e. on the mode “growth of body-shedding
of existing hard part as it becomes inadequate-
growth of new hard part”; (e) highly developed
digestive, circulatory and sensory systems,
including eyes; (f) durable exoskeleton, often
mineralized leading to high preservation potential
to make the rich fossil record of the group. In this
regard, particular mention must be made of the
effect of ecdysis; it produces a series of remains
for one individual, each representing an ontogenetic
stage in which the skeleton was moulted out
(Factsheet 14.2). Thus, a single individual may be
represented by a number of fossils, adding to the
numerical lot but demanding close scrutiny as well.
It always remains a possibility that variation of
morphology through ontogeny reflected in fossils
of its different stages may be mistaken as variation
at species level, if not properly judged.
As mentioned, Arthropoda is now considered
a superphylum by many authors (Manton 1973,
1977, Clarkson 1998). They subscribe to the view
that Arthropoda combines heterogeneous elements
that evolved from different ancestors; Manton even
considers that the evolution of arthropod-like
organisms (arthropodization) occurred at least
thrice producing three or more distinct phyla. The
issue is, however, not settled for the final.
FACTSHEET 14.2
Trilobite Ontogeny
Under exceptional conditions, ontogenetic stages of
trilobites from the larval stages may be preserved.
They (Figure 14.4 A, B, C) include:
Protaspis: A cambered disc, ventrally open;
segmented central axial lobe to become glabella later;
tiny eyes on anterior margin, which migrate inwards
and bring the facial suture with them.
Meraspis: Pygidium appears as a free segment
following the development of a transverse furrow;
thoracic segments are really parts of the pygidium.
Holaspis: Adult number of thoracic segments
reached; after a few more moults, the animal becomes
a fully grown adult.
The present discussion will be confined to
Trilobita, a very well-known group of Palaeozoic,
important in Indian perspective too. Systematic
position or affinity of this extinct group is also
debated; generally it is considered a class, though
some authors raise it to the rank of independent
phylum. Factsheet 14.1 presents some more
information on different groups of arthropods.
14.2 Introduction: Trilobita
As stated above, trilobites are generally ranked into
a class within Arthropoda. They are the earliest of
arthropods known, appearing in Lower Cambrian,
immediately above the early Cambrian Tommotian,
non-trilobite fauna; they ranged upto Permian. In
late Precambrian, well below the stratigraphic level
at which trilobites are found, there are certain
paired chevron-type markings, a kind of trace
fossils that are interpreted as scratch marks of
appendages of some animals, trilobites or, more
probably, their soft-bodied precursors.
Trilobites were exclusively marine. During
Palaeozoic, they evolved fairly rapidly into 1500
genera and several thousand species (Clarkson
1998). They were particularly numerous,

morphologically and taxonomically diverse and
spread out in all marine environments, specially
shallow water, in Cambrian and more so in
Ordovician. In result, they have been successfully
used in biostratigraphy of rocks of these ages. Their
importance in Lower Palaeozoic is further
enhanced by the fact that the other rapidly evolving
group of invertebrates of that time, namely,
graptolites, could not survive the rigours of shallow
marine realms on account of their delicate skeletal
remains and, hence, are rarer in these deposits.
Though there were free-swimming (nektic)
trilobites, like the miomerids (explained later) and
some might have had meroplanktic stage, most
trilobites were benthic and provincial in
distribution. They have, thus, been used in
demarcating biogeographic provinces and even in
palaeogeographic reconstruction of ancient
continents (Burrett and Richardson 1980). Further,
there have been interesting studies on the
functional morphology and adaptive significance
of morphological variations adding new flavours
to the multifarious use of these extinct animals.
Trilobite exoskeleton, called carapace, is borne
on the dorsal side of the animal, extending slightly
onto the ventral. The latter side is, thus, largely
vulnerable. For this reason, there are fossils of
trilobites, in which the carapace is preserved in a
rolled up condition, obviously preserving the event
of its having been attacked. The skeleton grows by
ecdysis. Particular problems or advantages of this
kind of growth have been indicated above; it may
be added here that during the phase immediately
succeeding a moulting and preceding the growth
of a new moult, the animal is left without any
protective covering. It, thus, increases the chance
of loss by accident, had there been any during those
phases. This also has its effect on fossil record.
14.3 Three Lobes and Segments
The name Trilobita points to the presence of three
lobes in the skeleton or carapace. In fact, there are
Chapter 14 Trilobita (Arthropoda) 233
tripartites divisions both along and across the body
or skeleton of these animals. The three main
transverse divisions, the anterior most cephalon,
thorax behind it and the posterior most pygidium,
refer to three major segments; whereas the
longitudinal divisions are referred as lobes. They
include an axial lobe along the symmetry plane
and two pleural lobes, one on either side of the
axial lobe. Dorsal furrow is the suture that
demarcates the axial lobe. For cephalon, part of
the axial lobe there is called glabella and the two
pleural lobes are called cheeks (Figures 14.1, 14.2).
Though lobes are given weightage in naming
the class, the three transverse segments are more
important in body organization, taxonomy and
other aspects of this group.
All the three segments may have further
segmentation in them. Of these smaller segments,
those of thorax are particularly well-marked.
Certain trilobites like the agnostids and the
eodiscids have fewer segments (two and two to
three, respectively) in thorax; they are the
miomerid trilobites. But in most other trilobites
there are several thoracic segments, the number
and morphology being typical of genera and
species; this is known as polymerid condition. In
comparison, segments in pygidium are often fused
together, specially along the margin and in
cephalon, they are evident only in the axial that is
the glabellar part. In the cheeks, equivalent to the
pleural parts, there are only two divisions: fixed
cheeks or fixigena adjoining the glabella and free
cheeks or librigena on the outer sides. The thoracic
segments are attached to the adjacent ones in such
a way that as and when the carapace rolls up, the
movement is compensated by each segment
slipping out from underneath the one lying anterior
to it. Thus, in normal condition, there is a portion
of each segment, called articulating half-segment,
that extends beneath the preceding segment.
Figures 14.1 and 14.2 present different important
morphological features of trilobite carapace.
Glabellar and pygidial segments are relatively
fixed or immovable and as said, often fused together.
234 Part Two: Major Invertebrate Groups

(a) (b) (c)

(d)
4 3 1
2

(e) (f)

Fig. 14.1 Trilobite carapace; lobes, segments and rolling.

(a) to (c) Lobes and segments: (a) Axial furrow defines the axial and pleural lobes, (b) Cephalon,
thorax and pygidium superimposed on the three lobes, (c) Segmentation complete, (d) Lateral view of
carapace, (e) One thoracic segment in dorsal view; (1) Axial ring; (2) Pleuron; (3) Articulating half-
ring; (4) Articulating facet; (f) Rolled up carapace in which thoracic segments slip out with articulating
half-ring making up for the extra length due to stretching.

14.4 Cephalon
In addition to glabella and cheeks, another
important feature on cephalon-part of the carapace
is the facial suture (Factsheet 14.3). It is the margin
between the two parts of the cheek, that is, between
fixigena and librigena. Its variation is
taxonomically important, but first we should note
its significance on the carapace.
During moulting of carapace, it is the
librigena or free cheek that is first separated along
the facial suture. What remains on the carapace
is glabella and fixed cheek together, called
cranidium. This is shed apart in the second stage
of moulting. Eyes of a trilobite lie on the facial
suture; when the free cheek and cranidium are
shed apart successively, the eye stands on the bare,
unprotected body.
Figure 14.3 shows the position of facial suture
in front of glabella as well as near the posterior
margin of the cephalon. The latter character that is
the posterior portion of the facial suture is specially
important as one of the main criteria for
classification of the group.
Morphological variation in glabella is
represented in the number and nature of its
segments, whether they are complete or partial and
fused, in the shape of glabella, particularly in
respect to its width, its height above the dorsal
surface, etc.
Another important part of cephalon is the eye.
Arthropoda, as a group, is characterized by well-

2
5
3 11
4
6
7
8
(a)
Fig. 14.2 Trilobite carapace.
(a) Dorsal view, (b) Ventral view. Palpebral lobe: raised platform behind eye, Occipital lobe/segment:
last segment of glabella, Hypostome: raised plate attached behind doublure, Doublure: extension of
the carapace onto the ventral surface,
1 Anterior, 2 Glabella,
5 Palpebral lobe, 6 Occipital lobe,
9 Posterior, 10 Hypostome,
developed and organized eye of a compound lens
type. In fact, trilobites may be considered to have
the first highly developed eyes in the animal world,
well-organized in structure and capable of powerful
vision. The compound lens of trilobites is made of
a large number of smaller lenses. Each lens is a
uniaxial calcite crystal, formed perpendicular to
the optic axis. Such a lens could, thus, avoid the
effect of double refraction. Trilobite eyes are of
two types in their internal organization. In most
genera, more so of Cambrian or such older age,
eyes are made of numerous small lenses of circular
or hexagonal cross-section. They lie in contact with
each other. This type, known as holochroal, is
generally smaller in size. In some genera,
particularly of later age and specially belonging to
Phacopina, small lenses of circular cross-section
are not in contact with each other. It makes the
Chapter 14 Trilobita (Arthropoda) 235
1
10
(b)
3 Eye, 4 Facial suture,
7 Pleura, 8 Thoracic segment,
11 Doublure
eyes larger in size. The type is called schizochroal.
The two types, differing in their organization,
must have been different in their working, though
it is not precisely and fully known how the two
types worked and how they differed between
themselves. However, a broad correlation between
the variations in eyes and mode of living can be
arrived at.
For instance, some trilobites have such eyes
that were probably non-functional, as it appears
from their organization. Presumably they were
blind. Similarly, trilobites without any eyes were
blind too. They might have been nocturnal or
inhabitants of deeper dark water, or might have
lived in crevasses or holes. Blind trilobites,
however, appeared later in the phylogeny of
the group, as such types are not found among
older forms.
236 Part Two: Major Invertebrate Groups

FACTSHEET 14.3
Facial Suture in Trilobites

Type Part behind the eye Example

Proparian Meets cephalon margin Phacops

in front of genal angle Dalmanites
Opisthoparian Meets cephalon margin Isotelus
behind genal angle Ptychoparia
Paradoxides
Marginal Along the cephalon margin Cryptolithus
not found on dorsal side Harpes
Gonatoparian Meets the genal angle Trimerus
Calymene
Part in front of eye and glabella Example

Trimerus, Isotelus
Joined on dorsal surface
Dalmanites
Calymene, Phacops,
Joined on ventral surface
Ptychoparia

FACTSHEET 14.4
Trilobite Enrolment

Spheroidal enrolment in Acaste which makes a ball of the carapace.

Double enrolment in which pygidium and last few thoracic segments are tucked under the cephalon.
Discoidal enrolment where the first few thoracic segments bend, and the rest held as flat plate, as in harpids
and trinucleids.

There are two alternative explanations of 14.5 Thorax and Pygidium

exceptionally large eyes of trilobites. One is that,
they were deeper water forms that needed powerful
eyes to look through the darkness. The other
explanation entailed that such big eyes helped the
animal to see clearly over a large area around.
So, they must have been nektic in habit, rather than
being the usual benthic type. A few trilobites had
the eye set on stalk that raised it above the glabella.
They might have lived within sediments with the
eyes peeping out above the sediment surface.
Other characters of cephalon are shown in
Figures 14.1 to 14.4.
Thorax, the second major segment of trilobite
carapace from anterior is characterized by its clear
and distinct smaller segments. The number of these
segments is typical of genera and species. Each
thoracic segment has an axial ring arched up and
flanked on either side by a pleuron (pl. pleura).
Two other parts of thoracic segments are
articulating half-ring and facet. The former lies
along the anterior margin of the axial ring which
is marked by an articulating furrow. The
articulating half-ring is semicircular in shape and
 Chapter 14 Trilobita (Arthropoda) 237

(a)
(b)
(d)
(c)

(e) (f) (g) (h)

(j) (k) (l)

(i)

Fig. 14.3 Trilobite ontogeny and cephalon.

(a) Protaspis stage of ontogeny (not to scale), (b) Meraspis stage, (c) Holaspis stage, (d) Bulbous
glabella with eyes placed on stalks, (e) Marginal facial suture, (f) Gonatoparian facial suture,
(g) Proparian facial suture, (h) Opisthoparian facial suture, (i) Cephalon with glabella, eyes, cheeks
and facial suture, (j) Same with glabella and librigena (free cheek) stippled (k) Same with cranidium
(glabella and fixigena) stippled, (l) Trilobite (only cephalon shown) without eye.

extends beneath the segment or the axial ring in
front of it. Distal end of each pleuron has a sharply
turned anterior portion called the articulating facet.
When the animal rolls up for protection, the
articulating half-ring slid back and out to protect
the anterior portion of the axial ring, exposed by
rolling (also see Factsheet 14.4 for rolling).
Conversely, the articulating facet of each segment,
or rather its pleuron, were tucked underneath the
pleuron in front, for the same purpose of protection.
Axial, pleural and interpleural furrows marked the
lateral boundary of the axial ring, a furrow midway
on the pleuron and the boundary between two
successive pleura, respectively.
A second feature of thorax, important in
taxonomy, is the biramous appendage. Each
pleuron has an appendage attached to the ventral
side; there is no mark of that appendage on the
dorsal side. Thus, they can be found only in rare
well-preserved specimens such as those found in
the Burgess Shale of Middle Cambrian of British
Columbia, etc.
These appendages have two parts each, one
above the other. Perhaps the upper one functioned
as gill, and the lower one helped in locomotion.
Both thoracic segments and appendages become
smaller in size, as does the carapace itself as it
narrows down towards the pygidium that is towards
the posterior.
Morphologically, thoracic segments may have
blunt, smoothly rounded ends or may be spinose
with sharp ends (Factsheet 14.5). Like the number,
238 Part Two: Major Invertebrate Groups

1
3
2

4
5
(c)
(a) (d)

(b)

7 8
9 (f)
(g)

11
12 13
(i)
(h)
(e)

Fig. 14.4 Different kinds of trilobite spines.

(a) to (d) Pleural and genal spines; (a) Both long drawn (1); (b) Both short (2 and 4); (c) Both sharp
and moderately long (3); (d) Macropleural spines (5); (e) Cephalic spine (6) and genal spine (10),
both long; (f) Long pleural spine (7); and long telson (9), extension of a thoracic segment; (g) Marginal
spines (8); (h) Axial spine (12); (i) Moderate genal spines (11) and occipital spine (13).

this character of thoracic segment may help in
identifying genera or species.
FACTSHEET 14.5
Spines and Allied Structures
Genal spines: Extension of genal angle.
Macropleural spines: Very long extension of
pleural ends on thoracic segments (sixth in
Cybeloides).
Denticles: Modified spine-like structures along the
anterior and lateral margin of glabella.
Features of pygidium, the posterior most major
segment of trilobite carapace that may be used in
generic or species identification, are as follows:
(i) pygidium may be micropygus (smaller than
cephalon; typical of most genera), isopygus
(equal to cephalon as in agnostids) or macropygus
(larger than cephalon, rather rare, found in the order
Lichida); (ii) the presence and number of pygidial
segments and on their basis the nature of the margin,
whether serrated or smooth; (iii) the presence of
spines. Factsheet 14.6 and 14.7 provide a few
additional facts on trilobite ecology and trace fossils.

FACTSHEET 14.6
A Few Facts on Trilobite Ecology
l Trilobites present convergent forms repeatedly
through from iterative evolution exploiting similar
niches.
l A number of morphotypes, occur, as below
l ‘Phacomorphs’, etc. : tuberculate, large eyed,
convex, nearly isopygous; infaunal suspension
feeders.
l Large eyed, axially raised, reduced thoracic
pleurae; unsuitable for resting on floor, hence
pelagic, slow swimmers; occur independent of
facies.
l Elliptical carapace, no spines, eyes flush with
surface of cephalon, head long-snouted; fast
swimmers.
l Small, blind, isopygous, miomerid (only two
thoracic segments) and global distribution;
pelagic mode for agnostids.
l Spines, denticles, etc. are variously interpreted as
resting, crawling devices.
FACTSHEET 14.7
Trilobite Tracks and trails
l A number of form-genera and form-species of
trilobite tracks and trails are recognized. Most
well-known of these are as follows:
l Cruziana are bilobed chevron trails that are taken
as traces of crawling, ploughing, shovelling,
burrowing activities.
Originally known as of algal origin, later
interpreted as trilobite trails, but contested; no
body fossil found associated.
But occurs in beds that are interbedded with
trilobite bearing units; are numerous and diverse
in Cambrian and Ordovician and rare in Silurian-
Devonian fitting with the distribution pattern of
trilobites, etc.
l The only body fossil directly associated with
these trails and traces is Calymene associated
with Rusophycus which are bilobed ovoid
resting, burrowing or surface excavation traces.
l There are also other traces such as Protichnites,
Trachomatichnites, Diplichnites, etc.
Chapter 14 Trilobita (Arthropoda) 239
14.6 Chemical Composition of
Carapace
Chemical composition of the carapace material is
another significant aspect. Most arthropods are
made of chitin in their hard parts. It was earlier
believed that trilobite carapace was also made of
chitin. But later studies could not bring out any
trace of chitin in trilobite carapace. Rather it has
been found to be made up of fine needle-shaped
or microcrystalline and laminated calcite. These
calcite crystals were probably joined together by
some organic compound, but its identity is yet
unknown.
14.7 Palaeobiogeographic and
Stratigraphic Use of
Trilobites
Trilobites gain importance in palaeobiogeography
on account of their provincial distribution
particularly in Lower Palaeozoic. Provincialism
decreased since late Ordovician through Silurian,
whereby trilobites became cosmopolitan. Early
Devonian again brought in some provincialism
(South America, Falkland Islands, southernmost
Africa), but due to late Devonian extinction, it
became less distinct.
In Lower Cambrian, some authors (Clarkson
1998) recognize two realms olenellid and
redlichid; in Middle and Upper Cambrian, pelagic
miomerid agnostids present broad distribution,
while endemic benthic polymerids are provincial.
On a detailed study of distribution of trilobites on
the strength of 450 faunal lists based on 1371
genera, Burrett and Richardson (1980), however,
hold that biogeographic patterns of Cambrian
trilobites cannot be explained on an early
Palaeozoic Pangaea; it needs conceiving relative
movements of several continental blocks separated
by wide ocean basins. Five realms are recognized
restricted to one (or rarely two) of the major
tectonic clocks: American realm in low
240 Part Two: Major Invertebrate Groups
palaeolatitudes in the Southern Hemisphere,
Europe (including Morocco) in high palaeo-
latitudes, Siberia in temperate Northern
Hemisphere, Australia in low palaeolatitudes in
the Northern Hemisphere and China in low
palaeolatitudes in the Southern Hemisphere.
In Lower Ordovician trilobites were
provincial, with a few genera cosmopolitan
(miomerids), some might have been planktic. Four
provinces fit with early Ordovician continents
(McKerrow and Scotese 1990) with a
palaeolatitudinal control on them. These are,
namely: (i) calymenacean-dalmanitacean fauna
of cold water high-latitude shelves (present-day
France, Spain, Central Europe and Turkey); (ii)
dikelocephalinid fauna of low-latitude shelves of
South China and Australia, (iii) bathyurid fauna
of tropical platforms of Laurentia in North
America, Siberia and North China and (iv) asaphid
fauna of isolated Baltic-Russian platform of
intermediate latitude. In contrast to these shallow
water faunas, deeper water benthic trilobites
marginal to palaeocontinents appear less controlled
by latitudes or continents (for example, olenid
biofacies in early Ordovician).
These studies indicate that changes in
continental patterns may often be envisaged from
biogeographic changes. Conclusions are often
contested, but they provide examples of how
palaeobiogeography on fossils may be used in
palaeotectonic modelling.
Cooling towards the end of Ordovician led to
mass extinction, which in turn caused breakdown
of provinciality. Subsequently as continents came
closer, tropical-temperate faunas became widely
distributed.
Trilobites earn maximum stratigraphical value
in Cambrian-Ordovician time. They appear above
Tommotian (basal Cambrian) zone and present good
zone fossils with abundant, easily recognizable,
often vertically short ranged and geographically
widely distributed genera species. At the same time,
they were provincial and facies controlled. Thus,
they served for good intraprovincial correlations;
correlation beyond provinces could only be
attempted on pelagic forms.
Ordovician zones for offshore sequences are
generally erected on graptolites; stratigraphical
classification and correlation of nearshore shelly
facies are done on brachiopods and trilobites,
because graptolites stand less chance of
preservation in the more turbulent conditions there.
Problems mainly crop up, as mentioned above,
because of provinciality and facies control. In and
after Silurian trilobites, however, become less
important; ammonoids in Devonian, microfossils
in Carboniferous assume importance. Trilobites
find their some use in local correlation and
classification.
14.8 Indian Record
Indian trilobite record may be judged in the
background of the above discussion. Cambrian
rocks occur in three localities, viz. Spiti and
Kashmir in the Indian Tethyan Himalayas and Salt
range in the neighbouring Pakistan at the northern
or northwestern fringe of the Peninsular India.
Occurrence of Lower Cambrian sequence is a
debated issue. Middle Cambrian and to some extent
Upper Cambrian are more definitely established.
In Salt Range, Middle Cambrian is characterized
by brachiopod (e.g. Neobolus), but in Spiti and in
Kashmir trilobites are more important. In all the
cases the fauna is largely local with very few shared
species. Redlichia noetlingi, the most significant
species is, however, present in Salt Range and
Spiti, but not in Kashmir. Ptychoparia is
represented by different species in the three
regions. Agnostus and Microdiscus, the two
cosmopolitan genera are present in Kashmir and
Spiti (though as different species), but not in Salt
Range. The same is true also for brachiopods; for
instance Neobolus present in Salt Range is absent
in Spiti and Kashmir. On the other hand, Obolus
etc. are present in the Himalayas, but not in Salt
Range.

These differences in faunal characters
prompted authors to suggest that Salt Range, Spiti
and Kashmir were mutually isolated by geographic
barriers. But there are also views which hold that
the difference may be apparent. Their arguments
hinge on the following points :
1. In most cases the records are not complete.
2. Trilobites evolved very rapidly in Cambrian
and so slight difference in age would have
given rise to different species.
3. Environmental and ecological differences
without any biogeographic barriers may also
have caused difference in composition.
4. Himalayan fossils have suffered deformation
in varying extents. This often added to
difficulties in recognizing species or genera.
To conclude, these extrapeninsular Lower
Palaeozoic basins of Indo-Pak subcontinent may
have belonged to a bigger zoo-geographic province
Chapter 14 Trilobita (Arthropoda) 241
in a vast sea that stretched from Dead Sea in the
west through Iran, Himalayas, Southern China, Far
East Asia and Australia to even western parts of
Northern America. This is often named Indo-
Pacific Province.
Ordovician in extrapeninsular India is named
differently in different parts. In Salt Range, there
is a prolonged post-Cambrian hiatus upto Permo-
Carboniferous. In Spiti and Kashmir Himalaya,
there was a break followed by an Ordovician
transgression. Ordovician-Silurian succession,
thus, produced enclose newer trilobites such as
Calymene, Illaenus, Phacops, though they were
dominated by brachiopod of orthid and
strophomenid groups.
Devonian is also poorly represented in the
Himalaya and the Upper Palaeozoic shows varied
development of marine, continental, even subaerial
volcanic environments. Trilobites are poorly
represented in these upper parts.
 Part three
Miscellaneous
 15
15.1 Introduction: Definition
Microfossils occupy an important position in
palaeontological studies today. In the organic world
present today as also in the immensely vast fossil
record since Precambrian, microorganisms and their
fossils are obtained in such abundance and varieties
that studies on them have itself formed a major,
independent branch of palaeontology, called
micropalaeontology. At the same time, it must be
made clear that neither in methods nor in application
or otherwise, there is any fundamental difference
between the studies of macrofossils and
microfossils. It means that controls, forces and
processes that fossilize and preserve the remains or
traces of organisms or that bring about their
adaptation to environment and evolution, are all
basically the same in the case of both the larger and
the smaller fossils, the macrofossils and microfossils.
That, even in spite of these, micropalaeontology has
assumed its independent status, is mainly because
of the size factor of microfossils.
Microfossils may be defined as: Fossils of
small-sized complete body of organisms or similar
sized parts of larger organisms, that invariably
require microscopic studies under normal optical
microscope and/or electron microscope, for their
diagnosis and evaluation, are microfossils.
245
Microfossils
Naturally, because of their size, microfossils are
likely to be found in fair abundance in a small piece
of a fossiliferous rock. (They are reported even in
limestone fragment caught up within basalt or in
metamorphic rocks like green-schist, rarely blue-
schist). For the same reason of their size,
microfossils are generally preserved in a much better
condition than their larger counterparts in the same
hosts. Thus, it is easier to assess their morphological
variations and present a descriptive, even statistical,
quantitative account of them. Factsheet 15.1
provides a brief history of micropalaeontological
researches to reveal how during the last one and
half of the century, or more particularly, during the
last half of the century, the subject has made
phenomenal advancement. Of course, the rapid
growth of this branch of palaeontology is largely
coupled with the development of fossil fuel
industries, namely, with prospecting and exploration
of these fuels such as gas, oil, coal, methane.
Moreover, with the advent of studies under electron
microscope, or of different sophisticated and refined
geochemical methods and such other modern
techniques based on instrumentation-electronics,
micro-palaeontology has acquired the present, very
modern, vast, penetrative character of itself. By that,
it has not only accumulated a treasure of usable data,
it has also helped in understanding many facets of
the earth’s history in further details.
246 Part Three: Miscellaneous

FACTSHEET 15.1
Important Events in Studies of Microfossils and Scientists Contributing

l Strabo l Microfossils (Nummulites) in limestones of pyramids of Egypt noted;

58 BC-25AD significance yet to be realized.
l U. Aldrovandri ü
1522-1607 ï
l R.Hooke ï l Lens and microscope invented, could bring out identity and significance of
1635-1703 ý microfossils in 1665.
l A. van Leeuwenhoekï
1632-1723 ï
l Linneaus þ l A few species of foraminifera referred in the 10th edition of Systema Naturae
(Carl von Linne) (1758) under the generic name of Nautilus and Serpula.
1707-1778
l First half of l Almost all types of microfossils were known although their identity and
19th century systematic position were not always and fully known; foraminifera were
considered as tiny cephalopods, for instance.
l H.C. Sorby l Inception of studies on microfossils from thin sections of hard rocks.
1826-1908 (1849)
l C.G. Ehrenberg l In Microgeologie, significance of microfossils as rock-forming components
1795-1876 (1854) pointed out; father of ‘micropalaeontology’.
l A.Brongniart l Use of ‘Nummulites’ in biostratigraphy.
(1823)
l E.Forbes l Puerbeck Beds classified into zones on ostracodes.
(1850)
l W.Dames and l Turonian age of host rocks determined on foraminifera collected from
LG Bornemann drill-cores.
(1874)
l Second decade of l Microfossils included in university curricula as separate subject.
20th century l Research on microfossils in laboratories of petroleum companies.
l Independent research laboratory on foraminifera founded (by JA Cushman)
and journal published.
l Fourth-fifth decades l Rapid and extensive development of micropalaeontology in tune with that of
20th century hydrocarbon exploration.
l J.Cuvilier l Concept of microfacies.
(1945)
l N.N.Subbotina l Erection of zones on microfossils.
(and later H. Bolli)

15.2 Basic Varieties 2. Microfossil s.l. (sensu lato) includes those

There are two major types of microfossils. These
are namely,
1. Microfossil s.s. (sensu stricto), which includes
those organisms whose complete body is of
microscopic size.
microfossils, which are parts of a larger
organism, requiring studies under microscope.
Besides, fossils of still smaller organisms,
particularly less than 50 micron in size, are referred
as nanofossils; they require electron microscope
for their studies.
 Chapter 15 Microfossils 247

Factsheets 15.2 and 15.3 will help frame idea mode, autotrophic or heterotrophic, that may help
about the varieties of these types. A few points are group the concerned organisms with plants or
being discussed here. First, they show that all the animals, respectively.
five kingdoms of organisms have microfossil Fourthly, most of the microfossil groups are
representatives in them. However, the two marine and planktic, though they may have benthic
kingdoms of unicellular organisms, viz. Monera representatives in them too. Ancient most
and Protoctista, are entirely made of organisms, representatives of Monera, and
microorganisms and their fossils. Other kingdoms particularly the cyanophytes that formed
may include some microfossil s.s. in them. Most stromatolites were benthics themselves.
of the microfossil s.l. are obtained from the
kingdom Animalia.
15.3 Use of Microfossils:
Secondly, most of the microorganisms and
microfossil groups are still living. Systematics and Taxonomy
Thirdly, with microfossils often it may not be
easy to determine their plant or animal character. Broadly and generally, microfossils may be used
The only criterion, in those cases, is the feeding in the same way as the macrofossils. Some of
the particularities with the microfossils are
discussed below.
FACTSHEET 15.2
Without microfossils, a large portion of the
Microfossils sensu lato organic world would have remained unknown. The
Phylum Type of Variants existence and morphology and other characteristics
of different groups of Monera and Protoctista can
Porifera Spicules made of be studied only under microscope. Knowledge,
calcium carbonate Parts thus, made reveals the vastness of the organic
or silica of shell world, both in number and in variety. It also brings
Cnidaria Calcareous or out the antiquity of life and living beings and the
sclerite skeleton
nature of the earliest life and organisms. At one
Annelida Faeces, chitinous /juvenile
stage of human knowledge, it was believed that
scolecodont shell or
Mollusca Faeces, calcareous skeleton there was no life in Precambrian, as fossils were
prism, rhyncolite found from Cambrian onwards and those were of
(calcareous beak macroorganisms. Micropalaeontology threw new
of cephalopod) light on the problem of existence and the kind of
Crustacea Faeces life in Precambrian. It also helped develop the idea
Echinodermata Spines, pedicellarie on the early evolution of life.
plates In regard to systematics and taxonomy, a second
Hemichordata Carbonized important contribution of micropalaeontology is the
remain of
frequent and successful application of different
chitinous skeleton
Chordata Teeth, scales and statistical methods (such as bivariate, multivariate
otolith (internal analyses). Since they are obtained in large numbers,
aragonitic ‘ear’) Pieces of phenotypic variations of species or populations of
of fishes; teeth bones microfossils can be judged quantitatively to assess
of smaller and determine the similarities-dissimilarities
mammals e.g. correctly. Thus, numerical taxonomy draws many
rodents examples from microfossils.
248 Part Three: Miscellaneous

FACTSHEET 15.3
World of Microfossils

INDEX

l Systematics l Included organisms l Size (in microns)

l Geological age l Few characters l Habitat
l Feeding habit

MONERA

n Cyanophyta: Schizophyta n Schizomycophyta: Schizophyta

l Cyanobacteria, cholococcales, blue-green algae
l 1-25
l Precambrian-Recent
l Calcareous wall; single/colonial; spherical cell in
earlier genera, later threadlike; colonies threadlike/
filamentous, branching
l Marine-freshwater; tolerant of temperature, salinity,
poor oxygen, ultraviolet rays; mainly benthic.
l Autotroph; phycocyanin (blue-green) and
chlorophyll (green) help photosynthesis.
l Bacteria
l 0.25-2(width)/ 1.0-10(length)
l Precambrian-Recent
l Wall of organic compound; rod-like, spherical or
sharp conical in shape
l Marine, freshwater, land; live in both oxygenated
or oxygen-free condition; tolerant of salinity and
temeprature
l Mainly heterotroph; saprophytic; some
chemotroph; some anaerobic, autotroph

PROTOCTISTA

n Chrysophyta: Chrysophyta l Marine, planktic

l Silicoflagellate l Heterotroph
l 20-200 n Bacillariophyta: Chrysophyta
l Cretaceous-Recent l Diatom
l Siliceous; disc/hemispherical in shape; join l 5-2000(generally 20-200)
together to form hollow rod of opaline silica
l Jurassic/Cretaceous-Recent
l Marine; motile, nektic/planktic; photic; tolerates
l Siliceous; single/colonial; frustule centric spherical
salinity, varied in cold condition
or pennate oval
l Mixotroph; photosyn the size by golden-brown
l Marine to land; photic; centrics planktic, pennate
pigment
benthic
n Chrysophyta: Chrysophyta l Autotroph; olive/golden brown pigment
l Chrysomonad cyst
l 3-25 n Dinophyta: Pyrrophyta
l Mid. Precambrian/Cretaceous-Recent l Dinoflagellate (e.g. zooxanthellae)
l Nearly spherical l 20-200 (cyst), 5-2000 (motile core)
l May live beyond seas; benthic l Silurian/permian- Recent Cyst cells made of
l Mixotroph; photosyn the size by golden-brown organic compound and motile cell siliceous; cyst
pigment spherical/oval;
n Ebridian l Marine. photic; temperature tolerant; motile cells
l Palaeocene-Recent planktic, cysts benthic, cause of red tide in seas
l Siliceous, with flagella, motile l Mixo-/generally autotroph; carotinoid red pigment
(Cont...)
 Chapter 15 Microfossils 249

FACTSHEET 15.3 (Cont...)

World of Microfossils
PROTOCTISTA

n Haptophyta: Chrysophyta l Marine, rarely freshwater, motile

l Coccolithophorid; coccolith scales n Sarcodina (phylum):
l 5-20 and 3-15, respectively
l Triassic-Recent n Actinopoda (subphylum/class)
l Calcareous wall; spherical/oval; scales puriform/ l Radiolaria (subclass)
fusiform l Radiolaria (polycystina and pheodoria)
l Marine; photic; motile; aquatic; diverse in warm, l 100-2000 and < 250, respectively
tropical water l Cambrian-Recent and Eocene-Recent, respectively
l Autotroph l Siliceous, some with organic compound or
made of strontium sulphate; spherical, with
n Acritarcha pseudopodia
l Acritrachs l Marine, planktic, symbiotic with zooxanthellae,
l 10-150 forms ooze below CCCD
l End Precambrian-Recent l Heterotroph
l Diverse morphology; made of organic compound
n Acantharia (subclass)
l Marine to freshwater; planktic; tolerant of l Pleistocene-Recent
temperature l Made of chitin/silica, colonial; like radiolarians
l Autotroph l Freshwater, planktic
n Chitinozoa n Rhizopoda (subphylum/class)
l 50-300 l Foraminiferida (order)
l End Precambrian/ Ordovician-Permian l 0 micron-190 mm
l Hollow flask/bottle shaped; chitin made l Precambrian/Cambrian-Recent
n Ciliophora: Ciliata l Highly diverse; with pseudopodia; made of organic
l Tintinnids: calpionellids compound/calcareous/agglutinated
l Ordovician-Recent and Mesozoic, respectively l Marine; one subdivision freshwater/brackish;
l Made of organic compound/agglutinated/ planktic/benthic
calcareous l Heterotroph
PLANTA

n Tracheophyta
l Spore/pollen
l 5-50 (microspore); <400 (megaspore) l Marine
l Devonian-Recent l Autotroph; with chlorophyll or other pigments
l Wall of organic compound; diverse morphology
n Chlorophyta
l Reproductive organ of land plants
l Green algae
n Rhodophyta l Few microns to ten/ twenty metres
l Red algae l Calcareous framework; single
l A few microns to10/ 20 metres l Marine to freshwater
l Calcareous framework; single/colonial l Autotroph; with chlorophyll
(Cont...)
250 Part Three: Miscellaneous
FACTSHEET 15.3 (Cont...)
World of Microfossils
ANIMALIA
n Conodonta
l Conodonts
l 100-5000
l End Precambrian-end Triassic
l Tooth-like; made of calcium phosphate (apatite)
l Marine
l Systematics debated; held as chordate
15.4 Use: Biostratigraphy
Before micropalaeontology made its ground,
biostratigraphy had developed on the basis of larger
fossils. But often there were inadequate number of
specimens available and that too, in not-always-
perfect condition of preservation. This often affected
and limited the use of larger fossils as guide or zone
fossils for regional or wider correlation.
Micropalaeontology helped solve many
problems of biostratigraphy. Particularly after the
appearance of calcareous planktons, more so of such
planktic foraminifers in Upper Cretaceous, many of
the younger geological successions are replete with
them and they have been successfully correlated on
their basis among themselves as also with the
geological column itself. A number of biozones have
also been erected using them. Thus, the last
appearance of the genus Globotruncana has made
it possible to locate the KTB (Cretaceous-Tertiary
Boundary) in many succesions all over the world.
The biostratigraphic importance of
microfossils does not lie just in their small size
and abundance. Benthics of different larger
organisms are largely facies controlled. Hence,
though they may be useful in local or regional
stratigraphic classification, their applicability in
global correlation or classification becomes
limited. Many benthics may, however, have a
meroplanktic stage that may cause their
n Arthropoda(Superphylum)
l Ostracoda (class): Crustacea (Phylum)
l < 1000
l Cambrian-Recent
l Calcareous or organic compound; bivalved; diverse
morphology
l Marine to freshwater/land; benthic/nektic
l Heterotroph
cosmopolitan distribution. On the other hand, these
larvae lack hard parts and so have poorer chances
of preservation. In contrast, planktic micro-
organisms have their well-developed mineralized
parts and are, thus, much more useful for the
purpose. Many biostratigraphically useful larger
fossils, with shorter vertical range and wide
geographic, are nektic; ammonoids as a group is
an example. But since they are extinct, their
systematics, functional morphology, relation with
environment are often shrouded with uncertainty.
Planktic microfossils are, said before, relatively
younger and have their recent counterparts. It adds
to their advantage.
15.5 Use: Palaeoecology and
Microfacies
Palaeoecological use of microfossils is
fundamentally similar to that of larger fossils. The
limiting factors such as temperature, salinity of water,
clearness and depth, availability of nutrients and
food, hardness of substrate, relation with biotic
associates, such as producer-consumer ratio, which
control the distribution of larger fossils, especially
marine fossils, are equally operative on microfossils
too. But because of their abundance and existence
of their extant representatives, it is easier to draw
inferences in regard to these smaller fossils, rather
the organisms. Thus, it is directly observable which
 Chapter 15 Microfossils 251

8
7

1
2

1 3

6 3
5 2
4
4 9 5 6
7

10

(a) (b)

2
3
1

6 4
7 5

9
8

10 11

(c)

Fig. 15.1 Portion of the world of microfossils.

(a) Diatom {1–X350, 2–X150, 3-4–X130}; silicoflagellate {5–X215, 7–X270}; ebridia {6–X260};
dinoflagellate {9–X370}; acritarch {8–X195, 10–X290}; (b) Ostracod {1-2–X23}; coccolith
{3–X2900}; calpionellid {4–X180}; radiolaria {5–X65, 6–X55, 7–X30}; (c) Spores-pollens
{1–X450, 2-4–X200, 5–X260}; conodont {6–X20, 7–X16}; cyanophyta {8–X6.5}; chlorophyta
{9–X0.65}; rhodophyta {10–X98, 11–X13}

genera or species of microorganisms are living at
what depths or temperature or, for that matter, under
other conditions and based on these observations, it
is possible to infer relatively more precisely under
what conditions of shallow marine water, the
sediments with fossils of these groups, were
deposited. In that case, even the benthics may serve
the purpose quite well. This is particularly true, for
example, for groups such as foraminifera or
ostracoda.
252 Part Three: Miscellaneous

(a) (b) (c)

(d) (e) (f)

(g) (h) (i)

Fig. 15.2 A glimpse into the world of microfossils.

(a) Diatom, (b) Dinoflagellate, (c) Calcareous nanoplankton cell, (d) Dinoflagellate, (e) Diatom,
(f) Radiolaria, (g) Coccolithophorid, (h) Ciliate protozoa, (i) Photosynthetic silicoflagellate.

Microfacies analysis is an important tool for
palaeoecology or biostratigraphy. Details of
lithofacies and biofacies, including composition,
arrangement, form, chemical characteristics, etc.
determined through microscopic studies give
precise depositional facies, etc. of the lithounits,
even their smaller subdivisions. Obviously, such
studies are augmented considerably with the
microfossils present in the rocks, as they are more
readily available than their larger counterparts.
15.6 Use: Hydrocarbon
Prospecting and Exploration
It has been said earlier that the rise in application
of microfossils is intimately coupled with
prospecting and exploration of hydrocarbons, such
as oil, gas, coal or methane, etc. Factsheet 15.1
makes it further clear. The subcrops, viz. drill cores
and well cuttings, etc. made during subsurface
exercises, bring out subsurface lithostratigraphy as
well as fossils there. For obvious reasons,
microfossils are readily available in better state of
preservation. This makes them particularly useful
in the whole exercise.
The application of microfossils in prospecting
and exploration of hydrocarbons mainly involves
biostratigraphic and palaeoecologic studies. Thus,
if any time-indicative lithounit characterized by
a distinct microfossil assemblage or event
(FAD, LAD, etc.) is located in any drill associated
with a hydrocarbon deposit or trap, prospecting is
largely directed towards a search for the horizons
with the same fossils, implying the same age in
other drill. Microfossils, thus, help in correlating
the drills. Similarly, palaeoecological studies of
microfossils from different drills help build up
depositional-diagenetic environment and facies
maps, essential for any exploration attempt.
15.7 Microfossils and Lithogenesis
There is one more important aspect in regard to
the use of microfossils in prospecting and
Chapter 15 Microfossils 253
exploration of hydrocarbons. This relates to the
role of microorganisms in lithogenesis and
hydrocarbon generation.
Though some larger fossils, e.g. anthozoans
of coral reefs, play important role in building up
rock deposits of fairly large volume and extent,
deposits formed by microfossils are often larger
and more varied, having been formed in different
environments. But before discussing that, a few
points on the taphonomy of microfossils may be
relevant to site.
1. Most of the accumulations of microfossils on
basin floors are generally allochthonous.The
remains of benthic microorganisms are easily
shifted horizontally, more or less, by waves,
currents or turbidity currents. Besides
bioturbation may cause vertical mixing of
remains of different times. On the other hand,
dead remains of pelagic microorganisms
shower down from water on the basin floor.
Spores and pollens, that make important
continental microfossils, i.e. may also be
distributed widely by wind currents.
2. In spite of such removal from their living
places, skeletal remains of microorganisms,
because of their small size, are not readily
broken down by mechanical processes. They
may, however, be affected by biological or
chemical causes; they may, thus, be corroded,
dissolved or otherwise. Dissolution, oxidation,
necrolysis through bacterial decomposition are
the common effects. They may, otherwise, be
preyed upon by scavengers or may be lethally
covered by algal mats or fungi.
In regard to mineralized parts (see Factsheet
15.4), skeletons made of aragonite are dissolved
below the ACD (Aragonite Compensation Depth
at average depth of about 1000–2000 metre);
calcite below the CCD (Calcite Compensation
Depth at average depth of about 4000–5000 metre).
Solubility of silica, on the contrary, increases with
depth; hence, in shallow waters siliceous remains
are more easily dissolved.
254 Part Three: Miscellaneous

FACTSHEET 15.4
Composition and Stability of Shell/Skeleton of Microfossils

Aragonite Chlorophyte algae cnidaria, many Easily dissolved or changed into calcite
molluscs, some smaller foraminifera,
otolith
High-Mg Calcite Polychaete annelid, Converted into low-Mg calcite on
> 9 per cent mole Alcyonarian sclerite, loosing Mg
MgCO3 larger and some smaller
imperforate foraminifera,
echinoderm, sponge spicules,
cyanophytes, rhodophytes, etc.
Low-Mg calcite Perforate smaller foraminifera, Stable and unchangeable
< 9 per cent mole ostracoda, most bryozoa, brachiopod,
charophyte algae, coccolith
Siliceous Silicoflagellate, diatom frustule
radiolaria, siliceous sponge spicule

These two controlling factors of taphonomy
of microfossils (transport and erosion-dissolution)
determine, at the first hand, how much of the
otherwise prolific accumulation of remains of
microorganisms will ultimately be preserved in a
rock record. These include the biomass, the fleshy
remains as well as the mineralized skeletal parts.
The latter contribute to the formation and extent
of the rock deposits; the former constitutes the
main ingredient of the hydrocarbon accumulation
to form.
Rock deposits develop primarily in three ways:
1. Skeletal remains of microorganisms are carried
by waves and/or currents as clastic grains
(bioclasts) and deposited elsewhere. In this
case, these bioclastic deposits may develop
cross laminations, graded beds, size- or even
shape-sorting, controlled by the same factors
or environments, as that determine similar
features in clastic rocks.
2. Many microorganisms collect and accumulate
during their lifetime, elements such as Ca, Mg,
Si, P, Fe, etc. from around them. Post-mortem
accumulation of these elements may give rise
to saturated or concentrated deposits of them
in the host rock.
Then again, microscopic studies of carbon-
saturated coal-beds may reveal remains of
different parts of plant bodies at different stages
of alteration or transformation. These attest to
the fact that such carbon-rich beds have formed
through accumulation and transformation of the
organic material of plant bodies. The same is
true also with petroleum. It simply means that
the process is the first and primary step towards
hydrocarbon generation.
3. A third type consists of laminated or bedded
rock deposits formed biogenically. These are
the outcome of some or other kind of activity
of concerned microorganisms performed
during their lifetime. Thus, stromatolites or
oncolites formed by cyanophytes include, on
one hand, calcium carbonate secreted by those
organisms during their metabolism as also very
fine particles of the same mineral that
cyanophytes trap from the water above with
the help of their continuously agitating hair-
like cilia. Both these components accumulate
in layers on the cyanophyte mat; new mat
grows above the accumulation to add to the
growth of laminated calcareous deposit of
stromatolites or oncolites.

Microfossils of a rock record may lead to
prospecting for the above deposits. Thus, the
presence of bioclasts may point to a possibility of
a larger bioclastic accumulation around;
cyanophytes to the presence of stromatolites;
abundance of dinoflagellate cysts may speak of
phosphorous deposit nearby.
15.8 Microfossils and
Hydrocarbon Generation
It is now accepted (Bjorlykke 1984) that the
biomass of the accumulation of microorganisms is
the major contributor to the formation of
hydrocarbon deposits. In aqueous basins,
particularly in seas and oceans, the upper 30–50
metre of the water column, i.e. within upper one-
third of the photic zone, produces the maximum
amount of biomass. Phytoplanktons are the main
producers. Succeeding levels of trophic chain
include zooplanktons, herbivores, smaller
carnivores and larger carnivores; at each of these
levels the biomass is consumed by the higher level
of consumers. Some amount of biomass is retrieved
in the excreta and faecal pellets of animals as
undigested food materials. Biomass production may
be enhanced in oceans at the sites of upwelling
where oxygenated cold water of depth comes up
when heated for some reason or other. On land,
plants are the primary producers; they dominate in
humid, warm to temperate climates. They are
succeeded, as for aqueous organisms, by herbivores
and carnivores of different levels of trophic chain,
similarly consuming the available biomass.
Unconsumed, remaining biomass accumulates
on the basin floor. But here again, it may be affected
by different chemical-biochemical processes. Thus,
oxidation, bacterial decomposition may change or
destroy a portion of the biomass. In result, only a
small fraction of the total material finally
accumulates. If that deposit takes place in an anoxic
condition, lack of oxygen and absence of aerobic
bacteria, burrowing or scavenging animals, etc.
Chapter 15 Microfossils 255
may prevent further destruction of the biomass to
a large extent. If burial is slow from a slow rate of
sedimentation, that enhances chances of
destruction; a rapid sedimentation, on the other
hand, may bring in large volume of clastics to
reduce the proportion of biomass in it. Thus, an
anoxic basin with a moderate rate of sedimentation
is the best choice for biomass to form an
accumulation least affected and destroyed.
Kerogen is the polymer-type of component that
does not dissolve out in organic solvents. It is a
kind of admixture of organic compounds, whose
nature and composition depend on the original
components of the biomass (Factsheet 15.5). It is
from this kerogen that oil, gas or coal form under
the controls of temperature, pressure, duration of
burial; presence of different minerals or chemical
elements or compounds that act as catalysts or
inhibitors.
Concerted efforts of geologists, palaeont-
ologists-micropalaeontologists may search out the
different events or products at different stages of
the processes discussed briefly above. That helps
in suggesting locations of hydrocarbon deposits.
Thus, if palaeoecological, geochemical or such
other kinds of studies locate a plant-rich succession
of shallow seas or of land or a site of upwelling;
that suggests the possibility of thicker
accumulation of biomass around. If there is an
anoxic event or horizon in the succession and if
the rate of sedimentation is found to be suitable, a
next step towards hydrocarbon prospecting may
be envisaged. The type of microfossil assemblage
may give some idea about the type of hydrocarbon
that may be available. This is how micro-
palaeontology helps in knowing first-hand, the all-
important deposits of fossil fuels, their location,
nature, etc. But it must be categorically stated that
the above discussion gives a simple, elementary
idea about their generation and it is impossible to
locate these deposits of immense value precisely
and exploit them efficiently and economically,
without taking help of many more sophisticated,
costlier and complex methods of exploration.
256 Part Three: Miscellaneous

FACTSHEET 15.5
Biomass, Kerogen and Oil-Gas-Coal
A. Major organic compounds in different groups of organisms
Organisms Organic compound Organisms Organic compound

Land plant Lignin, Tannin Dinoflagellate, etc. Protein

Carbohydrate Phytoplankton Lipid
e.g., cellulose, glucose Radiolaria Lipid-dominated
Diatom Carbohydrate Forminifera Lipid-dominated
(zooplankton)
B. Kerogen

Kerogen Source Chracteristics of the Chemical property Results in

type organisms organic compound Primary ratio H:C-O:C

Type I Pollen Lipid-rich High-low Oil in

Planktic algae alliphatic-compound-rich land or sea
Animal remains
Type II Phytoplankton, Alliphatic and High-low Oil in sea
zooplankton, etc. naphthanic
+ land plant compound bearing
Type III Land plant Derived from Low-high Coal on land
(Humic) Lignin, Tannin,
cellulose, etc.
C. Petroleum-gas-coal

Depth of burial Products Temperature

1 30°C
Kerogen Subbituminous coal
2 60°C
3 90°C
Oil Bituminous coal
4 120°C
5 Gas Anthracite 150°C
 16
Microfossils:
Foraminifera
16.1 Introduction
Foraminifera, animals belonging to the order
Foraminiferida of the phylum Sarcodina have
earned a specially important position in the world
of microfossils and in studies on them. They occur
in abundance in marine carbonate or shale-
dominated rock successions of different ages,
particularly of Permo-Carboniferous and then of
Tertiary time. At different stages of their
evolution, they underwent rapid changes; also
with variations in their environment of living,
these animals exhibited considerably diverse
morphology. In result, the different genera and
species of this order present a rich fossil record.
They are useful in identifying stratigraphic units
of different ages and environments, and by virtue
of that, stand out as important means for
biostratigraphic and palaeoecologic-palaeo-
environmental studies. This has prompted their
wide application in palaeontology and, more so,
in oil exploration exercises. Besides, though
microfossils sensu stricto by definition,
foraminifers are much larger than most other
groups of that category. So, they are relatively
easily located in rock occurrences and can be
257
identified, albeit provisionally, at generic and
sometimes even at species level with the help of
common hand lens or even by naked eye.
Foraminiferal research has, thus, developed at a
much rapid rate and before those on other
microfossil groups could pick up any momentum.
The mention of foraminifers (rather
Nummulites) as rock builders of pyramids may be
found in Herodotus (5th century BC), Strabo
(58 BC-25 AD) and Pliny the Elder (1st century
AD). The first classification of foraminifera (1826:
though as included in cephalopods) and
biostratigraphy was proposed by Alcide d’Orbigny
(1802-1875); Felix Dujardin separated the group
in 1835 from cephalopods on the basis of their
having pseudopodia (fine free-moving hair-like or
filamentous bodies which are flowing extensions
of protoplasmic substance, as different from
flagella in some other groups) and called them
Rhizopodes. Studies on foraminifera were revived
and brought to their modern shape in 1950s and
1960s, with deep sea drilling projects (JOIDES
Program in 1965; DSDP in 1968).
Foraminifers are unicellular organisms; they
belong to Protoctista (formerly called Protozoa).
The protoplasm of a foraminifera cell is
258 Part Three: Miscellaneous
differentiated into endoplasm and ectoplasm and
gives rise to filamentous extensions, pseudopodia,
on the surface of the body. These pseudopodia are
used in food-gathering, locomotion, etc.
Characteristically, foraminifers build a test or
skeleton of inner ectoplasm. Each test may be made
of a single chamber (unilocular) or may be
partitioned into a number of chambers
(multilocular: much similar as to be found in
cephalopod shells). Wall of chambers as also the
partition between two chambers, called septum,
are perforate. Foramen is the larger perforation
among them, lying on septa and by way of which
the chambers are interconnected with each other.
Aperture, single or multiple perforation in the last
chamber, maintains connection with the outer
environment. The shape of the chamber/chambers,
the arrangement of chambers in multilocular tests,
the nature of foramen and aperture, surface
sculptures including nature of sutures, i.e. traces
of septa on the surface–all these characters show
wide variations that make test morphology of
foraminifera diverse and diagnostic at all
taxonomic levels.
Morphological variation in foraminifers has
another root. Even single species of these animals
may be found to have at least two different types
of tests. Difference in reproductive processes is
the cause of such variations. In one type, the animal
is born in asexual way, whereas in the other it takes
place through sexual processes. The first type is
called ‘A-form’ or megalospheric and the second
‘B-form’ or microspheric. In the first case, the test
is of small size, whereas the embryonic apparatus,
called megalosphere and made of the initial one
or a few chambers, is large. In the second case, it
is reverse in that the test is large, but the embryonic
apparatus (microsphere) is small and microscopic.
However, besides these differences, other
morphological characters are often so widely
different in megalospheric and micropheric forms
that it becomes difficult to judge them as belonging
to one and the same species without critical
examination.
16.2 Foraminiferal Ecology
Foraminifers present a wide range of adaptation.
Overwhelming majority of them are marine; a
sizeable portion of them are benthics. Planktics,
though numerically less, have assumed much
significance in stratigraphic correlation and age
determination. Among the benthics, some live on
seafloor and are vagrant; some others live within a
few millimetres in the sediment. A few other genera
live attached, with the help of their pseudopodia
or the calcareous tests, to the substrate, or shells
of other organisms or even floating objects, such
as seaweeds. In the last case, they are often referred
as pseudoplanktic.
Benthics, and to some extent planktics too, are
controlled by different aspects of their
environment. But before coming to that, another
relevant point needs to be discussed. Foraminifer
tests are generally made of either of the two
components. A few subdivisioions, viz.
Astrorhizida and Lituolida develop agglutinated
tests. Here the animal acquires particulate matter,
either grains of quartz, mica or other minerals or
biogenic remains like tests of smaller foraminifers,
sponge spicules, and forms its test with the help of
these cemented by organic material, or calcareous
or ferrugineous secretion of the body. But most
other foraminifer tests are calcareous (majorily
high-magnesian calcite, occasionally aragonite). In
this type, the fine fibrous crystals of the mineral
are aligned in such a way as to leave innumerable
perforations in the test material of the wall or
elsewhere. Thus, tests become perforate. They
often have a glassy look (hence, called hyaline)
and allow some amount of light to pass through
them in thin sections under microscope, thus
looking transluscent.
But in some cases, such as in miliolids, the
alignment of crystals does not leave any
perforations to make the tests imperforate. They
look like porcelain (hence, called porcellaneous)
and are opaque in thin sections seen under
microscope.

A third kind of calcareous test is micro-
granular, in which calcite occurs as small
equidimensional granular crystals. It is found in
some Permo-Carboniferous fusulinid foraminifers.
Agglutinated, microgranular or porcellaneous tests
are found in benthic foraminifers, whereas
calcareous hyaline tests occur in both benthic and
planktic forms.
Among factors controlling foraminiferal
ecology, salinity and depth are the most important.
Almost all foraminifers are stenohaline and cannot
live beyond the normal sea water. But some
porcellaneous miliolines (e.g. Alveolinella,) can
live in hypersaline lagoons and a few other genera
like agglutinated Egerella and hyaline Nonion in
brackish water lagoons or estuaries. On the other
hand, genera such as Elphidium, can adapt to
different salinity levels. Thus, foraminifers may be
used to bring out the salinity condition in which
some rock deposit might have formed.
Most of the benthic genera and species, as also
most individuals live in shallow marine water.
In general, porcellaneous forms are inhabitants of
lower depths, whereas hyaline forms may live at
any depth other than beyond the CCD level.
Agglutinated forms, too, can live at all depths, even
at 4000–5000 metre mark. Planktic foraminifers
normally live at 6–30 metre depth of water.
They are, however controlled more by temperature;
thus, different genera and species are found to
occur in latitude-parallel biogeographic provinces.
The temperature plays an important role also in
the distribution of benthics. Most of them,
including larger foraminifer genera favour warm
water (18°C–22°C).
These patterns of distribution of extant
foraminifers amply help in bringing out the
depositional palaeoenvironmental or palaeo-
geographic conditions of the host rock units.
16.3 Some Morphological Details
Vast morphological variations displayed by about
1400 genera and 30,000 species of foraminifera,
Chapter 16 Microfossils: Foraminifera 259
of which about 4500 species are still living, provide
materials for independent books (e.g., Cushman
1948, Loeblich and Tappan 1964 and Glaessner
1973 among others). Here only a brief summary is
provided for the beginners.
Most foraminifers of geological past and
present have tests of diameters between 0.1 mm
and 1.0 mm. But they may reach more than 100
mm value, as can be found in the family
Nummulitidae, particularly in some of its species
of Middle Eocene times. Foraminifers are called
larger or smaller principally on the basis of their
size. However, smaller foraminifers (of diameter
<5 mm), also called microforaminifers, are
identified mainly on their external morphology.
Larger foraminifers of diameter > 5 mm have more
complex morphology and require studies of both
external characters and internal ones, the latter to
be observed from thin sections and under
microscope (Figures 16.1, 16.2).
Morphological variation of foraminifers is
centred principally around the following
characters: composition and structure of the test
material; number of chambers, particularly, one or
many; shape and arrangement of chambers, and
nature of septa and suture; nature of aperture and
other perforations; a few other internal hard parts
and surface sculptures, etc.
Major types of foraminiferal test materials, by
composition and structure, have been mentioned
above. Of all the types, calcareous tests present
most of the variations.
It has also been mentioned that tests may be
unilocular (e.g. in Lagena) or multilocular.
Chambers may be arranged in one single, straight
or curved series. Nodosaria has a straight uniserial
test. Curved uniserial forms may be coiled
planispirally as in Elphidium (a smaller
foraminifer) or in Assilina (a larger form) with a
bilateral symmetry plane at right angles to the axis
of the coiling. They may, otherwise, be
trochospirally coiled, showing no symmetry, as
found in Globigerina (a smaller form) or
Dictyoconoides (a larger form). Chambers may,
however, be arranged multiserially, in two series
260 Part Three: Miscellaneous

(a) (b) (c)

(d)

(e)
(g) (f)

(k)
(i)

(h) (j)

Fig. 16.1 Larger foraminifera.

(a) Assilina (A form: eq); (b) Nummulites (A form: vr); (c) Alveolina (A form: eq); (d) Nummulites
(B form: eq); (e) A handful of foraminiferal sand from river; (f) Alveolina (A form: vr); (g) Scale for
Figures (h), (i), (j), and (k); (h), (i); (j) different species of Nummulites in hand specimen; (k)
Dictyoconoides (Figures (e), (h), (i), (j) and (k) of hand specimens and (a), (b), (c), (d), (f) of thin
sections; eq. equatorial section; vr vertical section; A-form megalospheric, B-form microspheric; in
figure (d) the bar represents 1 cm; in (e) the scale is in mm; in (a), (b), (c) and (f) figures are 16X, 19X,
4X and 3.75X respectively).
 Chapter 16 Microfossils: Foraminifera 261

(a) (b) (c) (d)

(e) (f) (g) (h)

(k) (l)
(i) (j)

(m)

Fig. 16.2 A few representative shapes, coilings and ornaments in foraminifera.

(a) and (g) Planispiral; (e) and (h) Unilocular; (f) Milioline; (l) A larger foraminera. (b, (c), (d), (i), (j),
(k) and (m) Trochospiral; Except (l), all the others represent smaller foraminifera.
262 Part Three: Miscellaneous
(biserial) in, say, Textularia or three (triserial) in
Egerella. In many examples, earlier and later parts
of tests have different arrangements. Thus, in
Egerella the test begins multiserially with more
than three series and later becomes triserial;
Clavulina has a test first triserial and then uniserial;
Siphogenerinoides begins as biserial and then
becomes uniserial, etc.
Variation in morphology is added by variation
in coiling. As mentioned, a few genera have tests
not at all coiled; for example Nodosaria, a straight
uniserial form, Textularia or Bolivina straight
biserial. Saracenaria test is initially planispiral,
then uniserial straight. Entirely coiled tests may
be planispiral or trochospiral. Two more important
types of coiling in foraminifers are annular and
streptospiral or milioline. The former is basically
a planispiral coiling in which chambers are so long
and curved as to cover much of one coil, i.e. whorl,
even half or full of the latter. Thus, successively
added chambers appear as concentric annulae.
Cyclolina, Orbitolites, Linderina, etc. are the
examples.
Streptospiral coiling is a complex type. Here
chambers are coiled in a plane about an axis; at
the same time the plane itself is rotated or coiled
about another axis, at right angles to the former.
As a result of this second coiling, the plane of
coiling is placed at angular distances of 120°C,
144°C, 180°C, etc. The chambers are added in
cycles of 3, 5 or 2, respectively (respective
examples are Triloculina, Quinqueloculina and
Biloculina). It should be added that many genera
such as Nummulites, Discocyclina, Orbitoides,
Lepidocyclina also present different complex types
of coiling. In the first named genus, chambers are
rotated on a second axis that lies at right angle to
the main axis of planispiral coiling. In the three
latter orbitoid genera, the main chambers grown
in annulae are placed in an equatorial plane
(which is also the plane of symmetry). But they do
not assume much height perpendicular to this
plane; instead there develops a layer of lateral
chambers on each side of this plane in which
smaller lateral chambers or chamberlets are stacked
in tiers to make a fairly large thickness or height
of the test. A kind of internal structure made of
bundles of fibrous calcite and called pillars, add
strength to the test developing in between the stacks
of lateral chambers.
The shape of chambers also varies in
foraminifers. It may be spherical (e.g. in Lagena,
Globigerina), tubular (in Triloculina, etc.) prism-
shaped (in Textularia, etc.) and so on; or, in thin
sections, it may be circular (in Orbitolites etc.),
rectangular (in Discocyclina, etc.), hexagonal or
polygonal (in Lepidocyclina, etc.), ogival, rhombic,
etc. (in Lepidocyclina, Miogypsina, etc.).
In multilocular tests, chambers are separated
by septa. Aperture is the opening in the last
chamber. Earlier chambers have foramen
(pl. foramina), standing for their aperture at the
respective stages of growth. Chambers are
connected between them as also with the outer
world through foramina and aperture, respectively.
There are a few other structures like stolon, which
also help maintain interchamber connections.
Shape and position of aperture and its
associated structures vary in genera and species
and are often diagnostic for them. Aperture may
be basal (at the base of septum or chamber),
terminal (at the end of the test), sutural (on the
suture), peripheral (along the margin of the test),
etc. In trochospiral forms, it may be central,
umbilical or spiral. In its simplest form, the aperture
may be a circular opening, or it may be slit-like,
crescentic, etc. In many genera, there are several
openings standing for multiple aperture, they may
be arranged in a linear fashion or irregularly
(cribrate). The aperture may have radially disposed
or dendritic extensions or parts. In some genera,
the aperture may be partially covered by calcareous
tooth-like structures.
The junction of a septum with wall is called
the suture; it may be raised like a ridge or grooved,
straight or curved, of uniform width or varying
length. In planispiral forms, sutures are similar to
the two sides of the symmetry plane. But in
 Chapter 16 Microfossils: Foraminifera 263

trochospiral genera, they are generally different on groups (e.g. Orbitoidae or the planktics,). Most of
spiral (i.e. apical) and umbilical sides. the families of this age were, however, extinct by
The internal space of chambers may remain the end of Oligocene or Miocene. This has left only
void or may be filled in by tube or tooth plate, etc. a few larger benthic genera and species to live in
In complex tests of larger foraminifers, there are the present time, though smaller benthics and
pillars, septulae, etc. inside chambers. planktics are abound.
The test surface in foraminifers may be smooth As evident, phylogeny of foraminifers
or ornamented by simple or complex sculptures. witnessed adaptive radiation, diversification and
The latter include ribs, hollow or solid spines, rapid evolution at several stages. In result, there
papillae or granules, etc. emerged a number of genera and species with very
short stratigraphic ranges. Of them, benthics are
particularly useful in local or regional geology,
16.4 Brief History
whereas planktics are often very effective in
intercontinental stratigraphic correlation. Factsheet
Barring a few probable unilocular agglutinated
16.1 gives the names of a few such genera.
tests, foraminifers appeared in Ordovician. Silurian
saw the advent of calcareous tests. Septa in tests
FACTSHEET 16.1
started to grow in Devonian; that introduced
multilocular tests. Trochospiral tests were found Few Biostratigraphically Significant Genera
first in Carboniferous. Towards the end of
Larger benthics useful in local or regional
Palaeozoic, two families, Fusulinidae and
problems in particular
Endothyridae became abundant. Modern history
of the group might have been initiated in Triassic Permian Neoschwagerina
to pick up momentum in Jurassic only. True hyaline Late Cretaceous Orbitoides
Palaeocene Miscellanea
tests, miliolid genera and planktic mode of living,
Palaeocene-Eocene Discocyclina,
all this might have started from this point or later. Alveolina
Cretaceous saw diversification and radiation of Eocene Assilina
foraminifers. Both benthics and planktics developed Eocene-Oligocene Different species of
fast in this period particularly in its later parts. A Nummulites
number of genera became extinct at the Cretaceous- Miocene Miogypsina
Tertiary boundary (KTB); they included larger Planktics for global correlation
benthic Orbitoides and planktic Globotruncana. The
last appearance of the latter is a successfully used End Cretaceous Globotruncana
datum (LAD) to mark the KTB. The extinction was, Eocene Hantkenina
Different Tertiary horizons Species of
however, accompanied by the appearance of newer
Globorotalia,
groups like Nummulitidae or rapid radiation and Globigerina, etc.
diversification of several families of some older
 17
Miscellaneous
Fossil Groups
17.1 Introduction
The groups discussed in Chapters 7 to 14, make
the most of the macro-invertebrate fossil records
of India (and of elsewhere too, in the main); they
are also the better studied and better known groups.
A few other groups of invertebrates having both
macro- and micro-representatives merit some
attention, though they leave poorer records (or none
at all) in India, or are not adequately studied and
known. Some of these groups also have doubtful
systematic affinity. In this chapter, brief discussions
on at least some of them are proposed, on account
of their this or that importance. Groups covered
will include Porifera including Archaeocyatha
and Stromatoporoidea, Bryozoa, Pteropoda,
Chelicerata and Crustacea of Arthropoda,
groups of sessile echinoderms and graptolites.
In addition, brief treatments on trace fossils and
stromatolites are also included in this chapter.
17.2 Porifera and Bryozoa
Porifera (including Archaeocyatha and
Stromatoporoidea, following Clarkson 1998) and
264
Bryozoa are two animal phyla, with biological
organization of their bodies, simpler than that of
cnidarians in the case of Porifera and immediately
above cnidarians with Bryozoa. Archaeocyatha and
Stromatoporoidea are two extinct groups and their
systematic position is still not free of uncertainties.
Archaeocyatha ranges from Lower to Upper
Cambrian only and may, thus, have some
biostratigraphic significance, while Stromato-
poroidea appeared in Cambrian, became abundant
in Silurian and Devonian. For a long time, its upper
age limit was considered at Cretaceous; presently,
however, it is extended to Oligocene. The four
groups, however, have left some contributions
in making of organic reefs of different times.
This underlines their possible palaeoecological
importance. It is also the reason why they are
considered together in the present section.
All the four groups of animals are sessile
benthic, filter-feeders. The two extinct groups,
Archaeocyatha and Stromatoporoidea, as well as
Porifera, in general, and Bryozoa were and are
marine, though the latter two groups have a few
freshwater genera. Stromatoporoidea and Bryozoa
are strictly colonial and form thin encrusting or
creeping layers of their calcareous skeletal remains,

spread out successively on the substrate and, thus,
building up laminated limestone deposits.
Archaeocyatha and Porifera (sponge) are generally
single or solitary, though there are colonial genera
too. However, both provided important frame-
building components of ancient reef deposits.
Sponges are also known for their role in producing
biogenic siliceous deposits.
Among the four groups, sponges have the
simplest bauplan or biological organization of their
bodies. Their body has different kinds of cells, but
they are not organized into tissues; nor does the
body have any nervous system. In its simplest form,
a sponge body is long, vase-shaped; but there may
be three successively more complex types
(Figure 17.1.II A). In the simplest body
(ascon stage), there is a porous wall made of a
kind of cells called ‘collar’. In the next stage of
complexity (sicon stage), a space between the wall
and the enteron, or the main body cavity, is
occupied by a few ascon-made chambers. In most
sponges, at a leucon stage, between the main
central cavity, called paragaster, and the wall,
there is a layer of several sicon-type of chambers.
Paragaster has mouth and osculum at its upper
end. The layer of collar cells that form the cover
of sponge chambers are also found in some
protozoan animals, suggesting some affinity
between them.
Most of the sponges have skeleton made of
spongin, an organic compound, and/or calcareous
or siliceous spicules. Spicules stand the best
chances of fossilization and when they are packed
dense and connected, the whole skeleton is
preserved in fossils (Figure 17.1 IIB).
The phylum extends from Cambrian to Recent
(Holocene). Chaetetida, the order, is the best-
known fossil group and is placed under the class
Demospongia. They have siliceous spicules. On
the other hand, Calcispongia has calcareous
spicules.
Stromatoporoids (Figure 17.1 IIH) are
grouped with sponges on the strength of a recent
discovery of an extant sclerospongidian group
Chapter 17 Miscellaneous Fossil Groups 265
(they are demosponges, but with aragonite-made
skeleton in additon to siliceous spicules). Skeletons
of the latter have a few features like astrorhizae
which bear similarities with similar features in
stromatoporoids. The latter formed one of the main
components of Lower Palaeozoic reefs; they
produced thinly laminated calcareous deposits.
Stromatoporoid fossils have a few radiating
grooves at the upper end of their skeleton; these
are called astrorhizae. In addition, their thin
sections show a fine mesh-like skeleton
(cenosteum) formed of horizontal laminae and
vertical pillars. Externally, some stromatoporoids
form small mound-shaped structures.
Archaeocyatha is an ancient animal group, in
which calcareous skeleton looks like a long cup.
In fact, it has two cones, the smaller placed inside
the larger one. Intervallum, the hollow space
between the two cones is partitioned by radial and
vertical septa, whereas the walls of both the cones
are porous, pores being smaller on the outer cone-
wall. The cavity of the inner cone is equivalent to
the paragaster of sponges. There may also be
horizontal tabulae or small dissepiments inside the
inner cone.
Archaeocyathid animals are generally of 10-
25 millimetre in diameter and 50 mm in height.
There are some giant forms of 150 mm in height.
The best preservation of archaeocyathid fossils
is found in Lower Cambrian of Russia, in a
continuous succession. They have been
successfully used in biostratigraphy there.
The phylum Bryozoa (known also as
Ectoprocta, particularly to US scientists) is known
from at least 3500 living species and about 15,000
fossil species. Yet the phylum could not gain much
popularity among palaeontologists because of the
small, fragile skeletal framework of the animals
affecting preservation potentialities to a great
extent and hence adding to difficulties in their
diagnosis and systematics. Notwithstanding that,
the importance of bryozoan fossils in palaeo-
ecology is widely accepted, as these sessile benthic
colonial organisms preferring hard substrates in
266 Part Three: Miscellaneous
quiet water are known for their sensitivity towards
temperature and salinity. Individual bryozoan
organism, zooid, is basically tubular and provided
with tentacles, looking much alike the cnidarian
polyps. The body is coelomate with a distinct gut
and a mouth at one end and an anus at the other.
The animals do not have any separate respiratory,
excretory and circulatory system. Each zooid
secretes around itself, a tubular or box-shaped
skeleton of calcium carbonate called zooecium.
The colony (Figure 17.1 II G) looks like a beehive
in which individual zooecium have porous wall
that maintains interconnection among the
organisms. The phylum extends from Ordovician
to Recent. Of the three main classes, one is
freshwater dwelling and does not possess
calcareous skeleton. Another class of bryozoans
lives in marine, brackish or freshwater; a few
genera of this class may have calcareous skeleton.
The third class is strictly marine, has calcareous
skeleton and includes, for example, mesh-like
genus Fenestella, reported from Upper Palaeozoic
horizons of extra-Peninsular and Peninsular India.
17.3 Pteropoda
Presently Pteropoda signifies an order within the
subclass Opisthobranchiata of the class Gastropoda
in the phylum Mollusca. They refer to a kind of
marine organisms that range from Eocene to date.
The organism is small, generally 2-10 mm in length
(< 30 mm) and bears a straight conical or partially
coiled thin skeleton, absent though in some genera.
Pteropods are pelagic-planktic or nektic. There are
about 30 species in the present tropical belt; some
genera can stand cold water. They live at different
depths, but not beyond the ACD (Aragonite
Compensation Depth), where its aragonitic
skeleton is dissolved.
Geological importance of pteropods lies first
in their producing biogenic deposits of pteropod
ooze in deep seas (above ACD). As zooplanktons
they show worldwide distribution, though being
more abundant in shallow basins and along the
continental margins with high temperature and
salinity. Secondly, as presently recognized,
pteropods range from Eocene upwards. Hence,
Hyolithes of Cambrian, or Conularia of Upper
Palaeozoic, which were earlier grouped with
Pteropoda are now derecognized. Affinity of
Hyolithes is still uncertain; conulariids are now
considered as scyphozoans under Cnidaria.
Indian examples of pteropods include
epipelagic and mesopelagic forms recovered from
topmost samples of gravity cores from continental
shelf of the Indian Ocean of North Kerala.
Pteropod assemblages distinctly differ between the
inner shelf and outer shelf-slope types. A
comparison of bathymetric distribution of
pteropods with the planktonic foraminiferal species
Globigerina bulloides (upwelling index) suggests
that the abundance pattern of certain pteropod
species (Limacina trochiformis, L. bulimoides,
etc.) is also influenced by the intensity of upwelling
(Singh and Rajarama, 1997).
17.4 Sessile Echinoderms
Crinoidea (Cambrian-Recent) of the subphylum
Crinozoa and Blastoidea (Silurian-Permian) of the
subphylum Blastozoa are important representatives
of sessile echinoderms. Though significantly different
in morphology, the two groups bear some similarities
between themselves. In either case, each individual
sits (or used to sit) on a narrow stalk, like a bud. As
echinoderm, these animals have water-vascular
system and a skeleton made of numbers of calcareous
plates. Generally, they are and were gregarious, i.e.
live or lived in groups. As a result, on the death of the
organisms, entire or broken skeletons or disarticulated
plates accumulate to form a fairly rich calcareous
deposit, not uncommon in certain parts of their fossil
records. Crinoid limestones of Mesozoic are good
examples, found also in extra-Peninsular India
Pentremites, a blastoid genus is a guide or index
fossil of Carboniferous, found from Burma.
Cystoidea, earlier recognized as a separate group,
is now considered as members of Blastoidea.
 Chapter 17 Miscellaneous Fossil Groups 267

8 7

6
5

3 4

2 3
2 C
1 1
B (a)
A

A (i) D
B C
(ii)
(iii)

H (i) (iii)

(ii) (iv)
G
E
F
(b)
Fig. 17.1 Miscellaneous fossils.
(a) Graptolites:
A Graptolites; 1 Theca; 2 Virgella; 3 Aperture;
B Graptolite stipes; 1 Pendent; 2 Deflexed; 3 Declined; 4 Horizontal; (relative to sicula);
5 Reclined; 6 Reflexed; 7 Reclined; 8 Scandent;
(c) Variations in graptolites.
(b) Others:
A Porifera: elements: (i) Ascon; (ii) Sycon, (iii) Leucon;
B Sponge colony, C, D Sponge spicules, E Archaeocyatha, F Bryozoan animal, G Bryozoan colony,
H Stromatoporoid: (i) Colony; (ii) Colony; (iii) Longitudinal section of colony;
(iv) Transverse section of colony.

17.5 Chelicerata an extinct group Eurypterida, water scorpions of

Classified presently as a phylum of the polyphyletic
superphylum Arthropoda, Chelicerata (Cambrian-
Recent) includes spiders, mites and scorpions, the
horseshoe crab Limulus and related organisms and
Palaeozoic time. Diverse and heterogeneous, the
phylum includes animals characterized by an
anterior prosoma (cephalon and thorax of trilobites
and other arthropods together) of 6 segments, a
posterior opisthosoma ( = pygidium) with <12
268 Part Three: Miscellaneous
segments, a pair of jointed pincers (chelicerae) in
the prosoma. The two main classes are
Merostomata (Cambrian-Recent), representing
water chelicerates, along with the eurypterids,
and Arachnida (Silurian-Recent) embracing
dominantly terrestrial animals like spiders, mites
and scorpions. Scorpions are known from Silurian,
and spiders and their allies from Devonian. Their
fossil record is definitely poor, but recent finds of
several exceptionally well-preserved records,
including those in ambers (see Factsheets Apx.
1.12, 1.13, 1.14) have added great evolutionary and
palaeoecological significance to the chelicerates.
Eurypterids, a rare but spectacular extinct
group, ranges from Ordovician to Permian,
occurring in marine, brackish, hypersaline and
freshwater rocks. They seem to form environment-
controlled associations for each of the above water
conditions. Detailed functional studies (Selden
1981, 1984 from Clarkson 1998) suggest that most
eurypterids were active benthos, able to swim; they
were predators in feeding habit.
Eurypterids include the largest arthropods ever
known, about 2 metre in length.
17.6 Crustacea
Another phylum of Arthropoda, viz. the Crustacea
that ranges from Cambrian to Recent, is also diverse
and abundant. The animals are mainly marine, with
also freshwater and few terrestrial groups. Well-
known crustaceans include shrimps and prawns
(swimming types), as well as crabs, lobsters and
freshwater crayfish (benthic types), all belonging
to the order Decapoda and barnacles to Cirripedia.
Ostracoda, another important group of
crustaceans classified variously and regarded as a
class under the phylum Crustacea, make a group
of microfossils, which is the second most abundant
among the recent microfaunal organisms, next to
foraminifera. They are specially abundant in
freshwater or brackish conditions to produce rock-
forming deposits.
Ostracods have hinged bivalved carapace,
quite similar in appearance and function to shells
of bivalvia (Figure 17.2). But ostracod carapaces
do not show growth lines, as they grow by moulting
and not accretion. Besides, the carapace bears pores
that provide passage for setae or sensory bristles.
The surface may, however, be ornamented
variously; generally, freshwater forms have thinner
and smooth-surfaced carapace, while benthic
marine types are thick-shelled and have coarse
surface ornaments.
Mention may be made of another group of
crustaceans, namely, the estheriids. They are the
fossil conchostracans that belong to the class
Branchiopoda of the phylum Crustacea. Estheriids
range from Middle Devonian to Recent and have
sufficient environmental significance. They are
essentially fresh to brackish water forms, intolerant
of salinity rise, but are more commonly found in
ephemeral pools of seasonal types. Thus, estheriids
cannot cross the seas and take land route for their
dispersal. This has prompted their use in correlation
or environmental studies of continental rocks,
devoid or poor in other suitable biota.
An example comes from reports and study of
estheriids from Gondwana formations of India,
particularly in an attempt to delve into the problem
of fixing Permian-Triassic boundary in them and
into their palaeoenvironmental and palaeo-
geographic implications. Several typical Triassic
forms or assemblages are found in the red beds of
Pali Formation, Kamthi Formation of Ib river and
elsewhere, Mangli red shale and green shales of
Panchet Formation of the Damodar valley basins
(Ghosh 1993).
17.7 Graptolites
17.7.1 Introductory remarks
Animals belonging to the class Graptolithina of
the phylum Hemichordata are commonly known
 Chapter 17 Miscellaneous Fossil Groups 269

(b) (c)
(a)

(e)
(d)

(f) (g)

Fig. 17.2 Arthropods.

(a) Trilobite, (b) and (c) Crab Fossils (d) Barnacles, (e), (f) and (g) Ostracods.

as graptolites. Two major orders of the class
Graptolithina, viz. Graptoloidea (Lower
Ordovician-Lower Devonian) and Dendroidea
(Middle Cambrian-Carboniferous) are known from
Palaeozoic only; none of them leaves any record
in India, though there are good occurrences from
the neighbouring Burma.
Graptolites are colonial marine invertebrates.
Their affinity was debated for long. At one stage
they were considered allied to corals, though the
270 Part Three: Miscellaneous
latter have a much simpler biological organization.
Largely on Koszlowski’s findings in 1948, where
he noted structural similarities between graptolites
and a recent colonial pterobranch Hemichordata,
Rhabdopleura, graptolites are now believed to have
affinity with vertebrates and are placed in the
phylum Hemichordata. They, however, present
interesting findings from recent studies that may
justify their being treated in this chapter.
Most commonly, graptolite fossils appear in
shale, often black shale, as tiny saw blades with
one or both edges serrated or toothed. They may
be straight or curved, sometimes spiral, single-
branched or bifurcating or many-branched. Finer
details are, however, lost in these common
preservations; only from rare well-preserved
specimens can their skeleton be reconstructed.
Graptolite skeleton consists of a series of
hollow interlinked cups or tubes, made of a thin
material, called periderm. The first cup or conical
tube is called sicula; its aperture, an opening, points
downwards. The apex at the other end extends into
a long, hollow tube or rod, called nema. The sicula,
for most of its parts, as also the other parts of the
graptolite skeleton, is made of half-rings or
complete rings, marking perhaps daily increments,
called fusellae, which join on one side of sicula
along the zigzag sutures. On the other side of
sicula, there is a stout rod, virgella, projecting
beyond its aperture (Figure 17.1 IA).
Sicula gives off a number of cup-like thecae.
Together they form the colony. Organisms lived in
the thecae and the sicula. Aperture of each theca
points upwards, while at the apical end there is a
foramen in the first theca and a similar opening in
all later thecae that join up into a common canal
close to the nema. The foramen and the common
canal provide connections between adjacent
thecae, whereby food material caught by one
individual is ingested and shared by the whole
colony. The organism, the zooid was probably
lophophorate, which used a lophophore for food-
gathering.
Graptolite colony is called a rhabdosome. It
may have a single series or branch of thecae or
more than one branch. Each branch is called a stipe.
Branches may be arranged in different ways, the
type designated on the basis of the position of their
nemas. Thus, in Diplograptus, the two stipes are
placed back to back with the nema in between. The
arrangement is called scandent and biserial. The
other types, viz. reclined, reflexed, horizontal,
deflexed, declined and finally pendent are shown
in Figure 17.1 IB.
The order Dendroidea has an internal tubular
system, called stolon system, which is an
interconnecting means between thecae. The order
Graptoloidea, however, lacks stolon system.
Shape of thecae may vary in graptoloids,
being characteristic of genera (Figure 17.1 IC).
Branching of stipes are also genera-specific. Light
microscope and transmission electron
micrography studies on graptolite skeletal
ultrastructure as well as chemical studies of
graptolites have thrown new lights on the structure
and composition of periderm material. It was once
believed that periderm was made of chitin or any
related organic compound. Recent work has,
however, failed to find out any trace of chitin. The
exact composition is not yet known.
17.7.2 Stratigraphical use
Graptolites were widely and successfully used as
stratigraphical indicators, particularly for
subdivision and correlation of Ordovician-Silurian
rocks. This is chiefly because:
1. they were planktonic and widely dispersed,
2. most species were eurythermic (could live in
different temperatures) and, thus, not confined
to latitudinal belts,
3. the majority of graptoloids (not dendroids)
were epipelagic and, thus, might have been
preserved both in deep water and shallow-
water facies, and
4. the stratigraphic ranges of many species were
short.

There are, however, problems, such as:
1. Graptolites are usually preserved as
compressions in which details are lost and so
there may be problems in precise identification.
2. The time range of some graptoloid species is
rather long.
3. Graptolites being delicate in structure are
better preserved in fine-grained sediments. It
makes them normally confined to black or
grey shale facies. They are relatively rare in
coarser grained rocks, and, thus, in areas of
different facies, graptolites may not be
preserved in the unfavourable facies though
they should be expected at that time and
place. Mixed or alternating sequences of
graptolitic and shelly (trilibote-brachiopod)
faunas are considered the best for the purpose
of directly correlating the two. Such
sequences are relatively more common in
areas of shoreline oscillations.
4. Common association of graptolitic facies with
black shale with carbon content and syngenetic
pyrite in the latter also suggests that these
planktic organisms were preserved in
conditions where no benthic organisms,
scavengers or burrowers could live and destroy
graptolite remains. But they are also found in
other lithofacies, indicating that the
preservational factors might have played the
crucial role.
A high-resolution stratigraphy using as many
closely spaced events (first and last appearances
of species in time), rather than a few selected
‘correlation fossils’ has made it possible to
precisely correlate Tremadoc (Cambrian)–
Llandeilo (Lowest Silurian) sequences on a global
basis with the help of 130 graptolite species.
At the same time, it is appreciated that many
graptolites are widespread, many others are really
geographically restricted. This has caused
palaeontologists to define faunal units larger than
zones, but of more or less worldwide application.
These successive graptolite faunas covering the
Chapter 17 Miscellaneous Fossil Groups 271
whole stratigraphic range of the group are
anisograptid fauna, dichograptid fauna,
diplograptid fauna and monograptid fauna.
17.7.3 How did graptolites live?
Of the two major groups of Graptolithina,
dendroids were benthic, while graptoloids that rose
from dendroids to become a more dominant group,
were planktonic. Benthic dendroids are found
mainly in silts, sands and limestones of shallow-
water inshore origin, their remains not having any
floatational structures.
Majority graptoloids are considered as
probably free-floating (holoplanktonic),
microphagous feeders (feeding on microscopic
organisms). There are two opposing views as to
how planktonic graptolites actually lived. Some
authors suggest that graptoloids passively floated,
as drifters, in the ocean perhaps at different levels.
The alternative view is that graptoloids, after a
larval benthic stage, became planktonic as young
adults and were actively automobile.
Had they been drifters, they would need some
mechanisms to prevent them from sinking. Gas
bubbles within the nema, or extrathecal tissue
containing tiny gas pockets, called vanes, which
are lateral extensions to the nema and may be
flattened bladders, expanded webs in some
dichograptids that increased the surface area.
Spirally coiled graptoloids (Cyrtograptus, etc.) that
rotated round and round and upward to buoy up or
the final change to the monograptid pattern
producing long, slender, less dense colonies, are
possible adaptations for floating.
No graptolites have ever been found actually
attached to algae, as was envisaged earlier. There
are synrhabdosomes, which are radially arranged
associations of rhabdosomes, with their nemas
directed towards the centre. These may be genuine
life-associations. Bubble-like floatation structure
at their center is also not substantiated.
Hypothesis of automobility contrasts with that
of passive drifting. Feeding currents produced by
272 Part Three: Miscellaneous
zooids are thought to have been powerful enough
to propel the colony. Vanes, webs and other such
structures could have been functional in mobile
graptoloids equally as in drifters. Reduction in stipe
number, increasing symmetry and change in
inclination of stipes may also be evolutionary
response to this mode of life. However, that many
graptoloids could regenerate after breakage,
growing in the opposite direction, may go against
automobility. It may not have been likely for the
same organism to generate the same type of
currents helping same type of movement, even if
growing in opposite directions.
Computer analysis and simulation of the
branching structure of multiramous (many
branched) graptolites help in functional analysis
of graptolite forms. Computer simulations have
shown that various branching patterns just like
those in ‘real’ multiramous graptolites can be
generated by permutations of very simple rules. In
most dichograptids branching is dichotomous, i.e.
each growing stipe splits into two equal daughter
stipes at the same time, and as the colony grows
there will come a further zone of dichotomous
branching. The simplest computer-generated
models were based on just such standard
dichotomies. Instructions were given (1) for
branches always to split dichotomously at nodal
points, but (2) to alter the angle of dichotomy in
some forms and/or (3) to delay the initiation of
dichotomy. ‘Graptoloid’ shapes, thus, generated
were very similar to the quadriradiate
(Staurograptus), triradiate (Bryograptus) and
biradiate (Clonograptus) genera of Lower
Ordovician. A computer model with instructions
to generate curving stipes produced a close match
with Cyrtograptus.
To ascertain the significance of the patterns, it
may be argued that, if we envisage the zooids of
adjacent stipes cover the feeding area in between,
then a multiramous rhabdosome may be an effective
‘harvesting machine’. It could either fed by rising
or sinking vertically through the water, while the
zooids strained off the nutrients, or by remaining
stationary while currents passed through it.
This has been further ascertained in more
recent times with the help of experimental model
of graptolites. Life size models of aluminium tubes
covered with ‘clingfilm’ to simulate the density of
a living graptolite, were allowed to fall, in
horizontal orientation, freely through a column of
still water. Most of the multiramous forms rotated
slowly as they sank. In the process they spiralled
downwards. It increased the path covered without
overlap between the paths of descending zooids.
In large multiramous forms, such a spiral fall
sweeps out a broad harvesting path, and in thin
scandent forms it exploits a narrow path. The
experiment adds a new dimension to our
understanding, though how and when the fall
stopped and the rhabdosomes rose again, is not
yet known.
17.8 Trace Fossils
Though no organic group in particular, trace
fossils are included in this chapter, simply for
convenience sake.
By definition (see Chapter 1), ‘fossil’ en-
compasses traces of activities of an ancient
organism made during its lifetime. However,
understanding of such traces have changed
profoundly during the last 50 years or so. As a
result, studies on traces made by recent organisms
as well as of trace fossils have now developed to
form ichnology, a separate branch of biology-cum-
palaeontology (Palaeoichnology refers to the
branch on trace fossils and neoichnology to that
on recent organisms; trace fossils themselves are
alternatively called ichnofossils or ‘lebensspuren’
; their genera and species are termed ichnogenera
and ichnospecies, which are really form genera and
form species, since they are erected on
morphological features, without often being
known, which organism is responsible for which
ichnofossil).
In actuality, trace fossils are sedimentary
structures made by organisms. However, to
distinguish traces or trace fossils from many other
 Chapter 17 Miscellaneous Fossil Groups 273

biogenic structures, some authors prefer to add that reptile teeth marks on ammonoid shells are cases
the former must be related more or less directly to in point. In another scheme, a more descriptive
the morphology of the organism that made it or at approach is maintained using morphological and
least must convey some idea about it (Frey and preservational characteristics for classification
Pemberton 1985). Accordingly, biostratification (see Factsheet 17.2).
structures like stromatolites, etc. or other traces that As sedimentary structures, trace fossils have
lack diagnostic anatomical features are excluded
from traces.
Trace fossils find multiple use in FACTSHEET 17.1
palaeontology. They may serve as an evidence of
Behavioural Types of Traces
existence of organisms whose remains might not
have been yet located, in general or in any Dominichnia dwelling traces
particular section or succession. They may also be Cubichnia resting traces
evidences of some vanished fauna that could not Fugichnia escape traces
be preserved. For example, worms are often Repichnia crawling traces
important members of benthic communities. But Fodichnia feeding traces
they can only leave traces such as burrows and Pascichnia grazing/ surface
rarely microscopic jaws. Though not generally, in feeding traces.
some cases, traces may tell of what kind of
organism could have made it. Footprints or tail FACTSHEET 17.2
marks of dinosaurs are quite useful in exposing
the organisms responsible. Trace fossils may occur Some Common Ichnogenera:
Their Morphology and Preservation
in deposits, like coarse clastics, in which body
fossils are normally not to be expected due to pre- Cruziana (trilobite trails) ü
burial high energy milieu or post-fossilization Nereites (worm trails) ý
þ
effects like diagenesis. Rather traces are often
l Tracks/trails at sediment/water interface
augmented or reinforced by diagenesis; burrows
made in loose sediments are cemented to retain its Asteriacites (resting mark of starfish)
l Radial horizontal markings
form and structure. Trace fossils are preserved in
situ that is, at the same spot where the activity was Skolithos (vertical worm tubes) ü
Chondrites (branching galleries by worm) ý
performed. A bivalve shell with gastropod borings þ
may be removed by current or otherwise but l Tunnels and shafts within sediment

without any shifting of the borings themselves. Rhizocorallum (horizontal U-tubes);

A scheme of classification of trace fossils is ü
Zoophycos (inclined spirals),
based on the behaviour of the organism that made ý
Diplocraterion (vertical U-tubes) þ
the traces.Thus, there may be dwelling, resting,
escape, crawling, feeding and grazing (surface l Traces with a spreite (a web-like structure
feeding) traces (see Factsheet 17.1). As they tell usually with a series of concentric markings)
about feeding mechanism, locomotion or some Rusophycos (trilobite resting traces) ü
such physiological or biological activities, they Pelecypodichnus (bivalve burrows) ý
þ
help understand the autecology of organisms. Trace l Pouch-shaped markings
fossils are also helpful in throwing light on
Palaeodictyon (origin uncertain)
community structures, i.e. on synecology. l Net-like structure
Predatory gastropod borings on bivalve shells or
274 Part Three: Miscellaneous
sedimentological use too. Thus, they are useful in
top-bottom determination, as most traces are
depressions on the sediment surface, pressed or
excavated by the animal. They may also be used to
find out directions towards which the animals
moved. Generally, trace fossils are restricted in facies
range and are, thus, helpful in palaeoenvironmental
reconstructions. From distribution of trace fossils,
aided by sedimentological evidences, it may be
possible to link certain assemblage of traces with
depth of water and sediment types. On the first hand,
trace fossils indicate well-aerated water and the
absence of anoxic condition, in which the organisms
lived. Besides, a few ichnoassemblages may be
correlated with some conditions of deposition. They
are as follows. In shallow water, there is a higher
percentage of burrowing suspension feeders i.e.
filterers. Such an environment requires physical
protection from turbulence and light, where the latter
exposes the organism to predators. Besides, the
higher rate of sedimentation in shallow water may
cause sudden suffocation to death, which makes
deep burrowing less advantageous in such a
condition. Dwelling and resting traces assume more
importance in shallow water. Cruziana facies and
arthropod tracks are also typical there.
Deeper and quiter water support abundant
deposit feeders or swallowers, as well as feeding
and crawling traces, sediment miners and
Zoophycos facies. Lightless deep water, on the
other hand, find deposit feeders and grazers, with
feeding, particularly grazing traces and Nereites
facies being more common. Protection is less
important in this environment. At the same time,
rich supply of surface dwelling bacteria and other
planktons produce food-rich surface layers. This
prompts complex spiralling and meandering
patterns of systematic grazing.
Trace fossils may also suggest the rate of
sedimentation from the relative abundance of
certain burrows (like Diplocraterion) and from the
presence of borings in ‘hardgrounds’. The latter
indicate induration, with or without encrustation,
that had taken place before boring was made,
suggesting a temporary break in deposition before
induration.
However, trace fossils have their limitations.
The same kind of traces, if they served for same
kind of feeding or locomotory mechanisms, may
be made by different organisms and, as mentioned,
it is often difficult to detect the trace-making
organism. Cruziana is generally believed to be
trilobite trails. But it is known also from post-
Palaeozoic rocks, even Eocene deposits of northern
Spain. This restricts the use of trace fossils for
taxonomic purposes. It is also the reason that
behavioural classification falls short of
universal use.The morphological-preservational
classification is, on the other hand, based upon
artificial, rather arbitrary criterion. Generally, trace
fossils are also not useful for biostratigraphic
purposes, except for local stratigraphy.
Particular mention should be made of the
importance of trace fossils in early Phanerozoic.
They are important in understanding the explosion
of life at the Precambrian-Cambrian boundary and
the evolution of metazoan life. Precambrian traces
show a general evolutionary pattern from
geometrically simple to more complex and
variously shaped traces representing initial
evolutionary diversification of metazoans, with the
development of gut-bearing bodies. However, no
undoubted trace fossils are obtained below the
youngest Precambrian glacial sediments.
17.9 Early Life and Stromatolites
Prokaryote (cells without nuclei)- eukaryote (cells
with nuclei) separation is one of the greatest
divisions among organisms. They differ not just in
vital characteristics (see Factsheet 17.3); but the
basic environments in which they thrive also differ
fundamentally. Prokaryotes vary widely in their
tolerance of or requirement for free oxygen. Thus,
they were the earliest forms of life to evolve, during
a period of fluctuating oxygen. Eukaryotes, on the
other hand, require oxygen for metabolism and for
 Chapter 17 Miscellaneous Fossil Groups 275

FACTSHEET 17.3 blue-green algae, i.e. cyanophytes (Walter 1976;

emphasis added by the present author). They are a
Prokaryote-Eukaryote Contrasted kind of algal or microbial mat, in which organisms
Organisms Bacteria, Protoctists, fungi, metabolize, grow and reproduce on an actively
cyano- plants and animals accreting surface or zone. They are, however, really
bacteria products of complex microbial communities,
Cell size 1-10 m 10-100 m generally made of several to many species of
Metabolism Anaerobic/ Aerobic microorganisms interacting with a large number
aerobic of environmental factors. These organisms, neither
Motility Nonmotile/ Usually motile, now, nor in the past, are not necessarily
with flagella with flagella/cilia cyanophytes only (Walter 1976). The study of the
Cell walls Of sugars Of cellulose/chitin, evoluion of stromatolite-building organisms may,
and peptides not in animals thus, be analogous to understanding the evolution
Reproduction By binary By mitosis of forest-forming or reef-forming organisms. At
fission /meiosis different times and places, different independent
Cellular Unicellular Unicellular combinations of interacting organisms might have
organization Multicellular adapted to stromatolite-building habit. Oncolites
are moving stromatolites, i.e. developed on shifting
or moving objects like pebbles being rolled by
synthesis of various substances; so they must have currents. Thrombolites are non-laminated, clotted
appeared and evolved only when an atmosphere or mottled varieties.
with sufficient free oxygen was established. A few words need be spent on ‘cryptalgal
However, as evident in other instances of evolution, deposits’ with ‘layered fabrics’. Such deposits are
even after eukaryotes diversified and established of three types of which stromatolite is one, oncolite
themselves as successful forms of life in an is another. Thus, stromatolites are defined as
oxygenated atmosphere, prokaryotes could and did domal, club-shaped, columnar, etc. structures
find their niches and continued successfully there. characterized by a generally non-planar (curved)
Prokaryotes were and are represented by lamination ranging from a few micron to a few
bacteria and cyanophytes (or cyanobacteria, as they millimeters, possessing well-defined three-
are also called). Without having any hard part, they dimensional boundaries and which grow attached
are likely to be preserved rarely. But the long to the substrate. Oncolites are unattached
stretch of Precambrian time presents important subspherical, ovoid, or flattened structures
records of prokaryotic life through what are known otherwise similar to stromatolites. A third type,
as stromatolites. They are not fossils, yet they are cryptalgal laminites are laterally continuous
extremely significant and useful in understanding stratiform deposits characterized by nearly
the history of the earth and its life, of a period when continuous planar lamination of somewhat greater
life had just begun. thickness than stromatolites.
Stromatolites can form in any free-standing
17.9.1 Stromatolites: what, where and body of water, which fulfils the following
how conditions:
Stromatolites are organosedimentary structures 1. The environment is suitable for the growth of
produced by sediment trapping, binding, and/or appropriate microorganisms.
precipitation resulting from metabolic activity and 2. The rate of growth of constructing micro-
growth of organisms, primarily or principally organism build-ups exceeds their rate of being
276 Part Three: Miscellaneous

FACTSHEET 17.4
Types of Algal Mat and Environment

supratidal film mat

blister mat rarely flooded
upper pustular mat
intertidal
tufted mat desiccation
(increasing upwards)
lower
intertidal smooth mat
gelatinous mat
rarely exposed
subtidal colloform mat
open coastline tidal pond
sedimentation rate (increasing to the left)
NB: italicised mats form columnar growth; others cryptalgal laminites (modified from Hoffman 1976)

destroyed by other organisms, like scavengers, 1. Stromatolite bioherm or biostrome

borers and burrowers or of being eroded out
or mechanically or chemically otherwise
affected.
3. The rate of sedimentation is enough to preserve
the internal fabric and structure, but not so
much as to prevent colonization of micro-
organisms.
Morphogenetically defined, stromatolites are
small to large organosedimentary structures whose
growth is recorded by a succession of laminae.
These laminae represent intervals of accumulation
of fine sediment on surfaces of a microbial mat,
populated by a community of microorganisms,
particularly cyanophytes.
The gross morphology of stromatolites reflects
the environment in which they occur, representing
the products of interaction of physical and chemical
conditions of sedimentation and organic films and
plexuses. They are best developed in subtidal,
intertidal and supratidal zones of marine-basin
margins.
In ancient stromatolites, aside from
constituent mineral grains, there may be one or
four distinct structural entities of different scales.
They are—
(stromatolith);
2. Individual stromatolite form (coenoplase);
3. The fundamental lamination; and
4. The cellular structure of microorganisms,
rarely present.
Each of these four has its own features,
significant at its own scale of development, yet
influencing and dependent on others.
Precambrian stromatolites generally do not
contain structurally preserved organic remains or
consistent cellular microstructure. All that is found,
are morphologically distinct build-ups with internal
laminations. However, their similar morphology
and geological setting as that of Palaeozoic and
modern stromatolites, leave no doubt about their
biogenic origin.
Under conditions, where the appropriate
microorganisms are available, the macro-
environment places limits to the overall extent,
shape, size and make-up of stromatolite
development. Maximum relief which a columnar
intertidal stromatolite may attain above the floor
is governed by its position in the intertidal zone
and the tidal amplitude. It also requires early and
progressive lithification to prevent breakdown
 Chapter 17 Miscellaneous Fossil Groups 277

FACTSHEET 17.5
Indian Precambrian Stromatolites

Anabaria : Few species: Jammu and Kashmir and Andhra Pradesh (AP) (Cuddapah Spg)
Baicalia : Several species: Vindhyan Spg of Madhya Pradesh (MP); PC of Rajasthan, equivalents of
Jammu and Kashmir, Uttar Pradesh (UP), Himachal Pradesh (HP)
Collenia : Number of species: PC of Karntaka, Rajasthan, AP ; Vidhyan Spg.equivalents in MP, UP
Colonella : Several species: Cuddapah of AP, Vindhyans of MP and equivalents in UP, HP,
Jammu and Kashmir
Conophyton: Several species: Precambrians of AP, UP, HP and Jammu and Kashmir; Vindhyans
Cryptozoon : Few species: Karnataka, AP, MP
Jurasania : UP, HP, Rajasthan, AP
Stratifera : Few species: UP, MP
Boxonia : Collumnaefacta: Crossia: Epiphyton: Garwoodia:
Gaya : Gymnosolen: Nzeria: Kusinella: Masloviella
Newlandia : Nucleela: Platella: Plicatina: Plumia
Sajania : Spongiophyton: Tungussia: etc.

from currents and waves. In intertidal and
supratidal settings, prolonged periods of exposure
to air favours formation of fresh encrusting
microbial mats.
Biological activity in stromatolites are mani-
fested at the lamination level (microenvironment).
It involves cell growth, attendant production and
concentration of organic matter into a multitude
of small structures. This organic living mass
precipitate, agglutinate or trap mineral matter.
Eventually they undergo bacterial decay, leaving
only the harder mineral materials to remain.
Periodic, yet successive accumulation of such
mineral matter ensures preservation of stromatolite
structures. Location, extent and make-up of the
algal association are adaptations in balance with
the environment. The controlling factors are
sunlight (cyanophytes are photosynthetic),
temperature, CO2, water volume and chemistry,
turbulence, ionic concentration and presence or
absence of competitors, scavengers, etc.
For a stromatolite to grow, a sediment supply
of proper fine grain size must also be available.
Sediments are localized by traps of sticky or
velvety organic films.
The presence of stromatolites in otherwise
unfossiliferous Precambrian successions confirm
the existence of life at that time. Besides,
particularly in Proterozoic, stromatolites of
different times show a morphological variation,
leading to recognition of different form genera that
could be used in stratigraphical classification and
correlation of these successions. As evident from
the above discussions, stromatolites may also be
used in the reconstruction of palaeoenvironment.
17.9.2 Indian examples
Stromatolites have been reported from different
stratigraphic units of various localities of India.
Only a few examples are mentioned here as
representatives.
Thus, stromatolitic horizons of dolomitic
limestone are located in Iron Ore Group in Orissa,
Barbil area, associated with jasper and black chert
as well as Fe ore deposits. Stromatolite occurs as
laterally connected, internally regular columns
with evidences of microbiota in it (Oscilloriacea
family).
Stromatolites have further been used in
correlation of Middle to Late Proterozoic (socalled
278 Part Three: Miscellaneous

Purana) successions of different Indian basins. patterns of stromatolites and attendant

Following four stromatolite assemblage zones are
established from Lesser Himalaya.
1. Kussiella-Colonella assemblage zone.
2. Conophyton assemblage one,
3. Baicalia assemblage zone
4. Jurasania assemblage zone.
They are observed in the Vindhyan
Supergroup. The Cuddapah Supergroup presents
a complex zonation; while Chhattisgarh basin and
the Kaladgi Group exhibit some of the zones.
Regional variation is attributed to difference in
sedimentational pattern controlled by local
tectonics (Raha 199X)
In a different kind of work, Sarkar and Bose
(1992) show variable morphology and growth
hydrodynamic changes in a continuous set of
different environments (see Factsheet 17.6).
Stratigraphically younger example is provided
by De (1999). Here discrete bodies of
stromatolites, identified as biogenic shallow
marine cryptalgal bioherms, morphologically
distinct from fresh water and brackish water
lagoonal algal stromatolites, occur in Talchir
Limestones of the Gondwana Supergroup in the
Talchir Basin, Orissa. They stand for an additional
evidence of marine transgression during Middle
Permian into an otherwise continental succession
of the Gondwanas. Associated ichnological and
sedi-mentary features point towards a moderately
high, fluctuating hydrodynamic conditions of
deposition typical of tidal flat environment.

FACTSHEET 17.6
Stromatolite Morphology and Environment

A case study from late Proterozoic marine carbonate formation in the

Godavari rift valley, India. (Sarkar and Bose 1992)

Wave-agitated reefs: Stromatolite columns are larger, widening and branching upward with a preferred
directional growth; intercolumnar areas wide and filled up with broken stromatolite fragments, etc; stromatolite
bodies linear, ridge-like with broad bases and narrow tops.
Relatively less agitated tide-affected nearshore subtidal plain: Columns smaller, less branching and widening
with a preferred, but reversing directional growth; intercolumnar areas proportionately wide and contain broken
stromatolite fragments; internal laminae both continuous and discontinuous between the columns owing largely
to tidal velocity assymetry; stromatolite bodies tabular.
Calm and quiet slope margin zone: Columns small, close-nested, little intercolumnar areas; columns vertical,
uniform in thickness and unbranching; bodies cylindrical albeit with rather irregular lateral boundaries.
 18
Vertebrata of Chordata
18.1 Introduction
Vertebrata (also called Craniata) is a subphyhlum
in the phylum Chordata and Chordata is just a
single phylum among the several which make the
organic world. Yet in material and methodology,
studies on vertebrates acquire the status of virtually
an independent branch, rather a full-fledged subject
in palaeontology. Vertebrates are animals with
backbones and, thus, include fishes, amphibians,
reptiles, birds and mammals. They are
characterized by many different living groups, as
well as many other extinct ones. Most of the extinct
forms can be correlated with some or other extant
kins. So, it is fairly easy to make out the functional
significance or other importances of different
fossils or their body parts, on their comparison with
living counterparts. Besides, by and large,
vertebrate fossils are bigger in size than most
‘invertebrate’ fossils. So, it is again easier to carry
out different kinds of observations and studies with
vertebrate fossils. All this, has given birth to vast
amount of data and knowledge on vertebrates. It
is meaningless to attempt to cover the whole range
in the span of one single chapter of a book.
Textbooks like Romer (1966), Colbert et. al. (2002)
279
or Benton (2005), etc. provide enough materials
for a more comprehensive knowledge.
There are problems with vertebrate studies too.
In many instances, fossils are not adequate or
distinct enough. New fossil finds are always there,
for instance, those with reference to the earliest
members of the group (see section 18.2) or new
fossils of hominids, etc. Not rarely, they are
changing the existing views drastically (also see
Factsheet 18.1). New methods are adopted, often
to the same effect. How molecular studies have
changed the ideas on the origin of placental
mammals and human have already been discussed
in sections 5.5.2 and 5.8.4, respectively.
Sometimes, the changes are not beyond the limits
of debate or controversy. Yet we may need to know
them, as it is. Further, a proper understanding of
vertebrate fossils requires a fairly in-depth
knowledge of physiology, zoology and anatomy
of vertebrate animal and body, including the
skeleton; itself, that makes a separate branch of
biology. Any attempt to cover it widely and
thoroughly in a textbook of composite structure,
may distract the general readers or beginners to a
large extent. We can at best leave the matter
recalling the invaluable phrase “We can learn
280 Part Three: Miscellaneous
FACTSHEET 18.1
Conodonts: New Finds End Dispute
Conodonts, long held problematic, are now
recognized as a group of early vertebrates, a group
of jawless fishes (Benton 2005) with notochord,
dorsal nerve cord, myomeres, the tail fin, cranium in
front of the notochord, etc. A complete conodont
animal has been reported from Lower Carboniferous
of Edinburgh (Briggs et al. 1983; the genus is
named Clydagnathus). Conodonts range from late
Cambrian to end of Triassic and are often abundant
in and characteristic of the marine rock-units, to be
successfully used to mark some biostratigraphic
zones.
everything, but for any stage of learning something
will remain unknown”. There will still remain other
important textbooks, referred above, to quench our
thirst, if we mean to.
So, here in this chapter, there will be no attempt
at detailing with anatomy or morphology or
classification. Rather it will contain only an
introductory outline of the subject. The group will
be discussed on the background of a few interesting
junctures of its history, with a view to imparting the
reader an overview knowledge, as far as
comprehensive it could be within that span. The
outline will be drawn on the frame of recent revisions
in the subject, more so in the systematics, to keep
the reader abreast with the state of the art. The
discussion will be oriented to make it meaningful
for particularly Indian learners, as and when
possible, with references to issues or occurrences
relevant to Indian geology as far as practicable.
18.2 Debate Starts from “Where
to Start”
In spite of the vast span of knowledge on the group,
the origin of vertebrates is a debated topic. For
palaeontologists, the earliest fossils are considered
very valuable in deciphering the origin of any
group. However, it must be admitted that many, if
not most, of the questions on the origin of different
groups, are now being resolved not on fossils, but
on embryology, molecular biology, genetics and
developmental biology particularly the study of
genomes that help relate specific anatomical
structures or morphological features to genes and
such other branches.
With vertebrates, we find a revealing example.
As mentioned, vertebrates are chordates, though
all chordates are not vertebrates. Chordates show
a number of common characters like notochord
(a flexible tough rod along the length of the body
near its back), dorsal nerve cord, myomeres
(V-shaped muscle blocks), etc. On their basis, it is
now held that the closest living relatives of
vertebrates are the seasquirts and amphioxus; the
former belong to the subphylum Urochordata or
Tunicata and amphioxus to the subphylum
Cephalochordata of the same phylum. The
question on the origin of vertebrates is in fact
related to the question on the origin of chordates.
Intensive search and research along with
increasing interest on exceptionally well-preserved
records of fossils have extended the age of chordates
considerably downwards. At one stage the genus
Pikaia from the Middle Cambrian Burgess Shale,
was considered the oldest chordate, indicating that
chordates appeared during the Cambrian explosion.
Small jawless agnathan fish and other basal
(primitive) deuterostome (explained later) fossils
from Lower Cambrian of southern China
(Chengjiang) pushes the age further downwards
(see Factsheet Apx 1.2). However, these fossil finds,
throwing much light on the problem, do not tell the
last words on the origin of chordates.
18.3 A Necessary Digression to
Clinch the Issue of
Vertebrate Origin
Some amount of background information,
apparently a digression, may help the discussion
(see Figure 18.1). Molecular studies since the
1990s as well as morphological evidences have
 Chapter 18 Vertebrata of Chordata 281

L Plants

Animals
E
B
O Fungi

Plant
chloroplasts

Protists

Mitochondria

Bacteria

Eukaryotes

Archaea

Fig. 18.1 ‘Universal tree of life’ (Benton 2005) showing the current opinion on the relationships of different
major groups of organisms.
Major clade within Animalia: (B) Bilateria (bilaterally symmetrical organisms) includes three clades
on morphological and molecular evidences, (L) Lophotrochozoa (brachiopods, phoronids, annelids,
molluscs, etc.), (E) Ecdysozoa (arthropods, nematodes, priapulids, etc.), (D) deuterostomia
(echinoderms, hemichordates, chordates).
282 Part Three: Miscellaneous
shown that the animals (Metazoa) have a major
group, Bilateria, with bilaterally symmetrical
organisms that can be divided into three clades
(each clade is a monophyletic group originating
from one common ancestor and including all
descendants of that ancestor). These are, namely:
Lophotrochozoa (brachiopods, molluscs,
annelids, phoronids and some minor groups),
Ecdysozoa (arthropods, nematodes, etc.) and
Deuterostomia (echinoderms, hemichordates and
chordates).
A deuterostome is recognized from
embryology. During ontogeny, each animal starts
as a single cell. It then multiplies into two, four,
eight, and so on. Eventually, a hollow sphere of
cells is formed. It is the blastula stage. At the next
stage, a part of the surface of the sphere pushes
inwards, ultimately producing a sac-like body with
an opening towards the end from where the surface
started pushing in. The stage is the gastrula stage
and the opening is the blastopore. The hollow
space of the sac serves as the gut.
Most invertebrates are protostomes, in which
the blastopore ultimately becomes the mouth. The
anus develops as a secondary opening. In other
animals, including the chordates, the primary
opening of blastopore becomes the anus and the
mouth develops as a secondary opening; these
animals are called deuterostomes. As mentioned,
chordates along with hemichordates and
echinoderms are considered to belong to the
monophyletic group, Deuterostomia.
Chordates, in their turn, have four major types
in their early records, though there are questions
about their identities or systematics. They are
urochordates, cephalochordates, vetulicolians
(yunnanozoans) and carpoids or calcichordates.
These are fairly well-represented in the Chengjiang
occurrence, though that does not solve the problem
of the origin of vertebrates.
As held at present, the solution comes from
the study of genome. Recent studies on amphioxus
have cast much light on the origin of vertebrate
characters, particularly the head and brain. On the
basis of amino acid sequences and related studies,
it is now found that both amphioxus and vertebrates
share the same genes that are responsible for the
development of brain and head, as also of skeleton
in vertebrates. Thus, though amphioxus has a very
simple brain and very simple sense organs, same
genes suggest that ‘phylogenetic precursors’
(Benton 2005) of vertebrate brains, eyes, skeleton,
etc. are there in amphioxus. In addition, it also has
embryonic cells with gene sequences that are
homologous with the neural crest of vertebrates.
The latter is unique to vertebrates and is responsible
for the development of other vertebrate characters.
Thus, the gene homologies suggesting
morphological homologies indicate evolutionary
relationships between amphioxus or cephalo-
chordates (also called Acraniata) and vertebrates.
At the same time, these evidences negate the earlier
suggestion that vertebrates were derived from
tunicates or urochordates.
18.4 Vertebrates Through Ages
It has already been mentioned that the Chengjiang
occurrence has the earliest fish fossil, marking the
appearance of vertebrates in Lower Cambrian and
that vertebrates include the aquatic animals, i.e.
the fishes, the amphibians which can live both on
land and in water, the reptilians, the birds and the
mammals. Their development through this time of
Cambrian to Recent, was never uniform, the acme
of the subphylum appearing to have reached in the
latest epochs. Here is a brief account of their
distribution through ages (Figure 18.2).
The fishes, a non-formal term, include a
number of groups, of which the classes
Placodermi (giant armour-plated fishes) and
Acanthodii are extinct, having lived essentially
and reaching maximum development in middle
Palaeozoic, between Silurian and Carboniferous.
The jawless fishes without paired fins, class
Agantha, make a paraphyletic group (a group that
includes some, and not all, descendants of a
 Chapter 18 Vertebrata of Chordata 283

1 2 3 4 5 6 7 8 9

Cz

J
*
T
*
P

Cb

D
*
S
O

Ca

PC

Fig. 18.2 Distribution of chordata through geological ages.

Index:
1 Agnatha, 2 Placodemi, 3 Acanthodians, 4 Cartilagenous fishes,
5 Bony fishes, 6 Amphibians, 7 Reptiles, 8 Birds,
9 Mammals, Cz (Cenozoic), K (Cretaceous), J (Jurassic), T (Triassic), P (Permian),
Cb (Carboniferous), D (Devonian), S (Silurian), O (Ordovician), Ca (Cambrian),
PC (Precambrian),
* Major mass extinctions : end-Devonian, end-Premian, end-Triassic, end-Cretaceous.

common ancestor) that ranges from Cambrian to
Recent; it reached its acme or maximum
development during the upper parts of Palaeozoic.
Chondrichthyes, another class, are cartilaginous
fishes, of which the present-day descendants are
the sharks and rays. They appeared in middle
Palaeozoic; doubtful instances come from
Ordovician and definite ones from Devonian, the
group radiating since Carboniferous to reach
maximum development in the Recent times. The
remaining class of the fishes, Osteichthyes, or the
bony fishes, are divided into two monophyletic
groups, viz. the subclasses Actinopterygii and
Sarcoptreygii, both appearing in latest Silurian,
radiating through Devonian and reaching acme in
the Recent times. The former are ‘ray finned’ and
284 Part Three: Miscellaneous
the latter ‘lobe finned’ ones. The lungfishes (Order
Dipnoi) and the coelacanths (Order Actinistia) are
the two living groups of sarcopterygians. Infraclass
Crossopterygii assumes importance for being
ancestral to the land-dwelling Tetrapoda that
includes all vertebrates other than the fishes. Fishes
were most varied in Devonian which is, thus, often
referred as the ‘age of fishes’.
In the modern cladistic view, Sarcopterygii is
a clade that includes all tetrapods. The latter has
in it, Amphibia and Reptilia both paraphyletic
groups that appeared in Devonian and
Carboniferous, respectively. Both of them have
recent representatives, though their maximum
development took place in Permo-Carboniferous
and Mesozoic, for amphibians and reptilians,
respectively. Tetrapoda also includes Aves and
Mammalia, the birds and mammals. The
celebrated Archaeopteryx, of late Jurassic is the
earliest bird. Since then, the group has undergone
a number of radiations particularly in the Cenozoic.
Mammals appeared in the late Triassic, though at
that stage, distinction between mammals and non-
mammals was not very apparent and the exact point
at which the synapsid reptiles (cynodonts) gave
risa to mammals is established rather by arbitrary
decision (Benton 2005). Mammals, too, reached
their maximum in Cenozoic radiation.
18.5 Basic and Major Features of
Vertebrate Body and
Skeleton
The basic vertebrate body plan inlcudes chrodate
characters such as notochord, dorsal hollow nerve
cord, myomeres, postanal tail, pharyngeal ‘gill’
slits. The characteristics of vertebrates include
among others: a true head, well-defined sensory
organs,viz. ear, nose and eye, a true brain with three
parts, otic (i.e. auditory or hearing), olfactory
(smelling) and optic (visionary or seeing).
The skeleton of vertebrates, on the other hand,
is constructed basically in the following plan. The
long axis of the body with a backbone or vertebral
column running parallel to it lies horizontally in
most vertebrates. Any deviation from this position
is a specialized adaptation. The backbone covers
both the thoracic and the tail regions.
At the anterior end of the backbone there is
attached the head region. It includes the skull or
cranium which houses the brain and the sense
organs. In most vertebrates other than the jawless
fishes, there are two jaws with well-developed
teeth along each of them. In fishes, amphibians,
reptiles and birds, the teeth serve mainly for
gripping and cutting (rather tearing); in mammals
they are differentiated morphologically
(e.g. incisors, canines, premolars and molars) to
serve different specialized functions, viz. cutting,
tearing, chewing, etc.
Immediately posterior to the head region is the
thoracic part, whose skeletal components include
the ribcage that houses the gut and different organs
within it.
Among other important features of vertebrate
skeleton are the appendages, fins or limbs, used
for locomotion and balancing. Barring the more
primitive forms like the jawless fishes, all the
other vertebrates have such appendages in two
pairs; they are attached to the body by bony
girdles, pectoral or shoulder and pelvic or waist.
Jawless fishes have non-paired median fins, which
continue in other fishes in addition to the paired
ones (see Figure 5.6).
While the vital soft parts such as notochord
or brain distinguish the vertebrate animals, the
skeleton is particularly pertaining to palaeon-
tologists. The basic plan of the vertebrate skeleton,
given above shows that it serves a few vital
functions. First, it provides a stable framework for
holding the body and housing the organs. Next,
the skeleton provides improved means for feeding.
Thirdly, the appendages serve for locomotion,
helping food-gathering, self-protection, etc. The
head region with sensory organs and brain in it
serves like a controlling panel.

Despite all niceties and complexities, the
history of this group may be visualized to have
been pivoted fundamentally on the efficient use of
this skeleton and body. It meant developing more
efficient means of feeding, of moving and
balancing the body and skeleton, of creating space
and framework for the organs which were
themselves developing and changing, and finally
more efficient means of sensing the environment
and controlling the self-functions through the
activities of more and more improving brains.
On this background knowledge, we may now
look into a few major events in the history of the
vertebrates that ultimately produced this vast and
varied group of animals.
18.6 Early Innovations
The jawless fishes that appeared in Lower
Cambrian continue till today. However, two
innovations, namely, development of jaws and
development of paired fins, opened up multitudes
of adaptations in regard to feeding mechanism, type
of food, locomotion for food-gathering and
protection from predators. This caused the middle
Palaeozoic radiation that produced placoderms and
acanthodiians and various other groups of fishes.
The origin of paired appendages was a major
evolutionary innovation for vertebrates themselves,
marking the first step towards fin-(and later limb)
driven locomotion. The earliest vertebrate fossils
lack paired fins but have well-developed median
fins, suggesting that the mechanisms of fin
development were assembled first in the midline.
Recent studies on shark and lampreys (a kind of
jawless fish), suggest that median fin development
involves the same genetic programmes that operate
in paired appendages; the former, i.e. median fins
arising predominantly from somitic mesoderm,
whereas paired appendages developing from lateral
plate mesoderm. Thus, despite their different
embryonic origins, paired and median fins utilize
a common suite of developmental mechanisms
(Freitas, Zhang and Cohn 2006).
Chapter 18 Vertebrata of Chordata 285
18.7 Advent of Tetrapods
18.7.1 Tetrapods and Amphibians
The next significant event in vertebrate history took
place when first tetrapods appeared on the earth.
In contrast to earlier vertebrates, that is the fishes,
they lived on land. New finds add many new details
on this transition from fish to tetrapod.
As mentioned, tetrapods include amphibians,
reptiles, birds and mammals. These days, they are
regarded not merely informally. Tetrapoda is given
a systematic status of superclass, with the above-
mentioned four groups as classes within it.
After the origin, the basal tetrapods, generally
referred as ‘amphibians’ (living forms frogs,
salamanders and newts) radiated extensively
during Carboniferous and early Permian. The name
suggests their living both in water and on land.
The class Amphibia, thus defined, that is, including
both the basal tetrapods of Palaeozoic and the
others of later times, is really paraphyletic, since it
does not include many other descendant groups,
namely, reptiles, birds and mammals, which come
from the same ancestor as that of the ‘amphibians’,
particularly the basal tetrapods. The present trend
is to restrict the term Amphibia to a monophyletic
group of tetrapods that inlcudes the living forms
which originated in Traissic from one common
ancestor.
18.7.2 Problems of living on land:
Support
Before we assess the changes involved in the
transition from the fishes to tetrapods, that is from
water-dwelling to land-dwelling, we may
enumerate the problems and advantages of living
on land.
In water, a fish is buoyed up by it; its body
weight, then, is virtually zero. On land, the body,
the skeleton and the organs are under the
downward pull of gravity while the limbs tend to
raise it upwards.
286 Part Three: Miscellaneous
The backbone of a fish is adapted for lateral
stretching and bending to help swim freely. A
tetrapod is acted upon by gravity and must have
the vertebrae and muscles around the backbone to
prevent the body from sagging down.
18.7.3 Problems: Locomotion
Fishes move in a smooth gliding motion; tetrapods
move in jerky steps. Paired fins (a pelvic or hind
pair and and a pectoral or front pair) of
sarcopterygian fishes help walking, rather
wriggling on land, but not in the way tetrapods
walk.
Eusthenopteron, a tristichopterid, had
humerus, radius and ulna in their pectoral fins (as
also other smaller bones of tetrapod limbs) and
femur, tibia and fibula in their pelvic fins (again
with others). But it could not have walked properly
on land on its fins. It was because, orientation of
Eusthenopteron limb bones was not helpful.
Tetrapods needed changes in length and structure
of existing bones, the appearance of few bones and
well-defined elbow and wrist joints.
In fishes pectoral girdle was a part of the skull;
in earliest tetrapods it was separated from the skull.
Pelvic girdle was enlarged and firmly attached to
the vertebral column to the add force to the hindlimb,
as tetrapod walking is controlled by hindlimbs.
18.7.4 Feeding and respiration
Tetrapod jaw acted in a much simpler way than in
most fishes. The early members probably fed on
small fishes and newly evolved invertebrates,
namely, millipedes, spiders, cockroaches, dragon-
flies, etc.
Living lungfishes have functional lungs and
fossil osteolepiforms and most other early bony
fishes probably had them. Early tetrapods were
perhaps marginally developed over them. In all
likelihood, with large mouth and straight, short
ribs, they breathed mainly by buccal pumping
(through mouth) as do the recent frogs and not by
pumping of lungs by ribs and costal muscles.
18.7.5 Sensory systems and water
balance
Sensory systems were also changed in early
tetrapods. The lateral line system of fishes was
retained only by aquatic tetrapods. Eyesight must
have improved with larger eyes; the sense of smell
must have improved too. But evidences from fossils
are not there to prove this. Bones of early tetrapods
do not suggest their strong hearing sense, as those
of recent amphibians and reptiles do.
Dessication or drying up in air must have been
a problem. Either the early forms preferred
viscinity of water to make up for the losses, or they
might have evolved semipermeable skin to cut
down water loss.
18.7.6 Reproduction
Even highly terrestrial apmhibians lay their eggs
in water where the young hatch out into free
swimming tadpoles, the larvae. Metamorphosis
changes them into adult form. Fossil ‘tadpoles’
obviously rare, has recently been found in adequate
numbers from Carboniferous and Permian of
France and North America. They suggest a similar
mode.
18.7.7 Fossil evidences
Since the 1990s, newer finds have added much to
the knowledge about early, i.e. Devonian tetrapods.
For obvious reasons, these land-dwelling
organisms have left trace fossils, their footprints,
as well as body fossils of bones, though the latter
are isolated and fragile. The examples are as
follows:
1. The oldest remains are some ill-defined
footprints from Australia and isolated bones
and footprints from different localities of
Europe, the Old Red Sandstone continent.
2. Some Late Devonian taxa from Scotland,
Australia and the Baltics may be close to the
transitional stages from sarcopterygian fishes
to basal tetrapods.

3. Unequivocal tetrapods of latest Devonian
come from Russia, Latvia, China, North
America; they have different names.
4. The most commonly known genera of
Devonian are Acanthostega and Ichthyostega
from Greenland. These were full-fledged
tetrapods in bone characters, though they
retained some fish characters. They are, thus,
considered as primitive early amphibians,
though the relationships between these
tetrapods and their closest fish-relatives are
controversial.
These last named forms might have been still
aquatic. They retained a tail fin, a lateral line system
of sensation, internal gills, flexible vertebral
column, probable movement by sweeps of their
tails, pectoral (shoulder) and pelvic girdles oriented
in such a manner and the limb bones of such shape
that suggest more swimming than walking; broad,
flat hand and foot used more as a paddle— all these
suggesting a dominantly water-dwelling habit.
They could have waddled about on land. Large
skull and heavy ribcage would have made them
rest on belly, intermittently.
The fossils are found in fluviatile sediments
of meandering rivers with lycopod- and fern-forest
around. Thus, they might have been still largely
aquatic, even with functional gills in adults. Fully
terrestrial tetrapods appeared in Carboniferous.
18.7.8 Carboniferous scenario
During Carboniferous, most of the continents were
joined, with no Atlantic Ocean and much of Europe
and North America around the then equator.
There were forests in the tropical regions with
large trees and lush undergrowth. The former
included giant Club mosses, 40-metre tall lycopods
(e.g. Lepidodendron), < 15-metre tall horsetails
such as Calamites, ferns and seed ferns
(pteridosperms). Their remains produced rich
Carboniferous coals of Europe and North America.
Trees provided new habitats for flying insects
(including dragonflies as large as pigeons);
Chapter 18 Vertebrata of Chordata 287
decayed plants or humus and undergrowth
sustained ground insects, spiders, scorpions and
millepods (some as big as 1.8 metre long). This
varied habitat led the tetrapods to diversify; some
remained aquatic feeding on fishes, while most
others explored and exploited the land habitats.
New finds from Scotland (early Carboniferous) and
Europe and North America (late Carboniferous)
have added valuable information on this radiation
(also see Factsheets Apx 1.6 and Apx 1.7).
18.7.9 Summary narrative on origin of
tetrapods
It is the suborder Rhipidistia of the order
Crossopterygii of the subclass Sarcopterygii, class
Osteichthyes, the bony fishes, that is now regarded
as the ancestor of the tetrapods. The same order
also includes Coelacanthiformes (coelacanths) that
is now represented by the genus Latimeria,
obtained from the sea between Madagascar and
Africa as also near Indonesia.
The early crossopterygian fishes of Devonian
have the characters that we may expect in ancestors
of land-dwelling animals. Osteolepis, a Devonian
crossopterygian genus, had a strong notochord,
prominent bony elements of vertebral column,
completely bony skull and jaws with well-defined
bony pattern. These, particularly the last one, along
with other evidences, indicate the position of
crossopterygians on the direct line of descent
between fishes and amphibians.
Sharp, pointed teeth, well-adapted to grasp
prey and indicating carnivorous habit of early
crossopterygians, also had a complex labyrinthine
enamel pattern as in early amphibians.
Nasal passages in crossopterygians went
directly from external nares or outer nostrils
through internal nares into the mouth or pharynx.
This was also as in higher land-living vertebrates.
Particularly interesting was the internal
structures of the paired fins of early crossop-
terygians. The single proximal (near the body)
bone in the paired fins may be equated with the
288 Part Three: Miscellaneous
upper limb-bone of tetrapods, viz. humerus in the
front leg and femur in the hind leg. The next two
bones may also be homologous with the radius
and ulna of the front leg and tibia and fibula of the
hind leg of land-dwelling vertebrates.
Osteolepis is an early crossopterygian genus.
It is included in the crossopterygian lineage which
is represented by the Suborder Rhipidistia. These
were freshwater fishes of Devonian to Permian age.
The late Devonian genus Eusthenopteron were
advanced over osteolepid ancestors and nearer to
early amphibians. In addition to the characters of
paired fins stated above, this genus had a skull
pattern, advanced nature of vertebrae, strong
notochord, dorsal spines on it, etc. that were very
close to early amphibian characters. ‘It was indeed
but a short step from Eusthenopteron to land-
dwelling vertebrates’.
18.7.10 Hypotheses on transition
On the anvil of appreciation of the problems of
living on land, as mentioned above, the available
fossil evidences and detailed observations of the
characters of fossils, it seems likely that air-
breathing, normally believed to be the critical
factor behind the transition from fishes to tetrapods,
might not have been the key hurdle. Rather, it could
have been the weight and structural support
conducive to the new modes of locomotion
accompanied by new ways of feeding, of sensing
prey and predators, of water balance and
reproduction that might have played the critical role.
This demanded a reassessment of the
hypothesis on the transition. The prevailing
hypothesis by Romer (1966) envisages that in
Devonian, the period of Old Red Sandstone in
Europe, being rather a period of seasonal drought,
streams might have dried up to be divided into a
series of smaller ponds and puddles. In such an
environment, some of the late Devonian
crossopterygians may have been forced to seek new
freshwater pools or streams in which they could
continue to live. Any adaptation that allowed them
to wriggle or move along the dry stream bed, would
have increased their chances of returning to a pond
or puddle that still had water in it. And this
adaptation could ultimately have led to the change
from an aquatic to a terrestrial mode of life.
However, evidence of aridity was considered
not enough to lend support to this hypothesis.
Besides, it could explain moderate terrestrial
adaptations, but was not enough to explain much-
modified tetrapod limbs.
A simpler alternative hypothesis from Benton
(2005) entails that vertebrates that could develop
adaptations for living on land proved to be more
successful, as they flourished on the rich and
untapped supply of food that opened up with the
advent and diversification of land plants and
terrestrial invertebrates living thereupon, in late
Silurian and Devonian.
18.8 Early Amniotes
Some small lizard-sized terrestrial tetrapods, living
in drier parts of forests on insects, laid eggs that
did not hatch in water. They were the first amniotes
and included the ancestors of all subsequent major
tetrapod groups (reptiles, birds and mammals).
These early amniotes were also reptiles. The
traditional class Reptilia, like Amphibia, is also
paraphyletic, as it does not include birds and
mammals, though they all have the same ancestor.
The oldest amniotes are obtained from middle
Carboniferous (310–300 million years ago) of
Nova Scotia. These and the other early amniotes
were extinct by the huge end-Permian mass
extinction. Among them were the ancestors of the
dinosaurs and the mammals that dominated later
periods.
Early amniotes fall into three main groups:
synapsids, diapsids and anapsids. Of these, the last
two, belonging to the clade Sauropsida, are
themselves broadly sister groups. Synapsida is an
outgroup (Benton 2005). Therapsida is a major
descendant of Synapsida. A synapsid of early
Permian of Texas, USA (Tetraceratops) may be
the oldest known therapsid, intermediate between

sphenacodontid pelycosaurs and later therapsids;
those from late Permian of Russia, including
Biarmosuchus are other examples of early
therapsids. They were characterized by enlarged
temporal fenestra, reduction of palatal teeth, etc.
the derived characters of therapsids, in comparison
with the ‘pelycosaurs’.
18.9 Importance of Triassic in
Tetrapod Evolution
Triassic marked the transition from older
(Palaeozoic) fauna to modern types. End-Permian
extinction reduced tetrapod fauna in both diversity
and spread. Synapsids witnessed new radiation with
newer groups, including aquatic groups (fish-
eating) like ichthyosars, etc. Diapsids in place of
synapsids became dominant. In late Triassic
dinosaurs, as well as other major groups like
crocodilians, pterosaurs, turtles and mammals arose.
All these took place on a Triassic earth, with
still a single continent, which, however, started to
break up towards the end of the period. However,
in early Triassic, the occurrence of dicynodont
Lystrosaurus and some others in widely apart
locations like South Africa, Antarctica, India, South
America, China and Russia point to a global
supercontinent.
Triassic climates were warm, though there was
a shift from warm and moist to hot and dry during
the period. This climate change affected floral
scene; early Triassic seed fern Dicroidium,
dominating southern continents, was replaced by
conifer-dominated flora of northern type. This
affected the herbivores, which gave way to the first
herbivorous dinosaurs.
18.10 Dinosaurs
18.10.1 Introducing the terrible lizards
Dinosaurs make one of the most well-known group
of fossils. As far back as in 1676, when Professor
Chapter 18 Vertebrata of Chordata 289
Plot of Oxford University described and illustrated
a bone-fossil larger than any known to man, he
had no idea as to which animal the bone could have
belonged to. But unwittingly he had opened a new
chapter in man’s knowledge. About one and a half
centuries later, in 1822 Mrs Mary Ann Mantell,
strolling leisurely in soothing spring air of a British
countryside called Tilgate Forest, picked up from
the roadside rubbles a piece of rock with a small
sharp glossy object peeping out (Colbert 1968).
Meanwhile her husband, Dr Gideon Mantell
(1790-1852), an ardent fossil hunter too, was
attending his patient indoors. The object came out
to be a tooth fossil. And this simple, uncere-
monious event led to the discovery of more fossils
from the same locality, to an arduous study by Dr
Mantell on this basis and finally to the discovery
of a second example (named Iguanodon in 1825)
of a group of reptilian animals that no longer lived
on this earth. Almost simultaneously, Dean
William Buckland (1784-1856), a theologist by
profession and a man with keen interest in
science, had described in 1824 fossils of
Megalosaurus, the first known example of the
same group of reptiles of the past. The fossils
found by Professor Plot belonged to this group.
Later, in 1842, the group was named Dinosaur,
the ‘terrible lizards’.
Eventually, the group was found to have
appeared in very early parts of late Triassic and to
have been extinct by the end of Cretaceous, at the
KTB. During this span, it developed remarkable
variation in morphology, as well as adaptations
(see Figure 18.3). From gigantic Diplodocus to
terrifying Tyrannosaurus rex, through the way
stations of the pterodactyl (now grouped with
Pterosauria and not Dinosauria) or Triceratops.
They had reigned unchallenged for about 200
million years on land, in the sea and in the air;
they were superbly adapted to their environment,
they never ceased to grow larger and larger; yet all
at once they vanished from the face of the Earth
some 65 million years ago. Why? (paraphrased
from Allegre in Courtillot 1999)
290 Part Three: Miscellaneous

0.5 m

(a)

(b)

0.5 m

(c)

(d)
Fig. 18.3 Dinosaurs: a few ecomorphotypes.
(a) Bony plates on back/quadruped, (b) Long neck giants/quadruped, (c) Horned quadruped,
(d) Giant biped.

18.10.2 Origin of dinosaurs
In middle to late Triassic, a major radiation of
archosaurs led to the initiation of dinosaurian
radiation. (Figures 18.4, 18.5) The oldest dinosaurs
come from very early parts of late Triassic. A sister
group Marasuchus is latest mid-Triassic, indicating
that the dinosaurs must have originated by that
time, from their common ancestor (about 230
million years ago).
Early Triassic evidences of skeletons and
footprints of dinosaurs, so far reported, have not
stood scrutiny in later studies. (Wild 1973;
Thulborn 1990; King and Benton 1996; Benton
2005). The oldest true dinosaur fossils are now
known from Ischigualasto Formation of Argentina;
they are from early parts of the Late Triassic
(Eoraptor and Herrerasaurus). They were rather
small (former 1 metre long , the latter 3 metre long)
animals with dinosaurian synapomorphies
(shared derived characters). Coelophysis from
North America are also held as Late Triassic.
Dinosaurs were once regarded as polyphyletic;
cladistics have indicated their monophyletic nature.
18.10.3 End triassic dinosaurian
radiation sparks off a
macroevolutionary debate
There was a major dinosaurian radiation in the
latest parts of Triassic; a macroevolutionary debate
prevails over the cause of this radiation, whether
it was a successful competition with the
synapsids, the basal archosaurs and the
rhynchosaurs, or an opportunistic radiation
following an extinction event. The second
alternative is now favoured on the strength of the
following evidences so far available, though it
always remains a possibility that newer data may
call for fresh changes in opinion (Benton 2005):
1. There were two rapid expansions and no
gradual change, one at the end of Carnic
(225 million years ago) and the other at the
end of Rhaetic (200 million years ago)
Chapter 18 Vertebrata of Chordata 291
(Divisions of Late Triassic: Carnic or Carnian;
Noric or Norian and Rhaetic or Rhaetian, in
the younging order). The first exterminated all
dominant herbivorous non-dinosaur groups;
herbivorous dinosaurs evolved rapidly in
Noric. After the Rhaetic extinction, dinosaurs
again radiated in Jurassic.
2. The first dinosaurs did not take over at once.
Of all the three major dinosaurian lineages,
theropods and sauropodomorphs did not
radiate immediately after their appearance
and the third, the ornithischians not before
Jurassic even.
3. Evolution of ‘erect’ gait, held as a mark of
superiority of dinosaurs among tetrapods was
there in many other Late Triassic archosaurs,
but they did not evolve further. So, it did not
prove dinosaurs a superior competitor.
4. The end of Carnic was marked by some events
both on land and in sea, such as (i) the
appearance of new types of corals,
scleractinians in seas; (ii) the extinction of
Dicroidium flora in the southern hemisphere
and rise of worldwide conifer flora; (iii) the
change of climate from pluvial (heavy rainfall)
to aridity over much of the earth. These might
have brought about the extinction of
herbivorous dinosaurs.
5. Simple competition as a means of replacement
of one major group by another is now
considered over-simplification.
More information on dinosaurs are given in
Factsheet 18.2.
18.10.4 Unusual fossils
Studies on dinosaurs have always been an
engrossing exercise. Through years, various kinds
of evidences and materials have been unearthed
and are still coming up to add newer and newer
lights to this topic. There have been reversals too.
Conclusions are sometimes debated, even
evidences challenged on authenticity. Even then,
it is felt that a brief mention of some interesting
292 Part Three: Miscellaneous

6 8

K D
A

Jr B
A

(b) (a)

Tr

Fig. 18.4 Dinosaurs: Phylogeny (simplified, showing a few major groups).

(a) Ornithischians, (b) Saurischians
Index: K (Cretaceous), Jr (Jurassic), Tr (Triassic)
A Ornithopods, B Sauropodomorphs, C Ceratopsians, D Theropods,
1 Coelophysis, 2 Stegosaurus, 3 Apatosaurus, 4 Camptosaurus, 5 Triceratops, 6 Ankylosaurus,
7 Tyrannosaurus, 8 Trachodon.
 Chapter 18 Vertebrata of Chordata 293

3 6 9 10

8
7

1
(a)

* * * E * *F * * I J ** L * * O * Q R
M
P
H
D G
C N

B
K

A (b)

Fig. 18.5 Dinosaurs: Cladograms suggesting phylogenetic relationships as accepted now (simplified
from Benton 2005).
Index: (a)1 Archosauria, 2 Avesuchia, 3 Crocodylomorpha, 4 Avemetatarsalia, 5 Ornithodira,
6 Pterosauria, 7 Dinosauromorpha, 8 Dinosauriformes, 9 Marasuchus, 10 Dinosauria.
(b) A Dinosauria, B Saurischia, C Theropoda, D Coelurosauria, E Tyrannosauridae, F Aves,
G Sauropodomorpha, H Sauropoda, I Brachiosauridae, J Titanosauridae, K Ornithischia, L Stegosauria,
M Ankylosauria, N Cerapoda, O Ceratopsia, P Ornithtopoda, Q Iguanodon, R Hodrosanridae.
294 Part Three: Miscellaneous

FACTSHEET 18.2
Dinosaur Information: Summary of Details

1. Earlier thought polyphyletic, artificial group; cladistics suggest it as monophyletic.

2. Appeared in Late Triassic about 230 million years ago; became extinct at the end of Cretaceous,
65 million years ago.
3. Earliest types were small to moderate-sized bipedal carnivores; by the end of Triassic there appeared
large quadrupedal herbivores; during Jurassic and Cretaceous there were large and small carnivores,
massive herbivores, small fast-moving specialized plant-eaters, forms with different types of armours,
bony plates, horns, etc.
4. Dinosaurs lived with their sister group pterosaurs, the flying reptiles, and the birds, derived from
carnivorous dinosaurs, in the sky, turtles, crocodilians, lizards, snakes and mammals on land,
ichthyosaurs, plesiosaurs and in Late Cretaceous, giant mosasaurs, in the seas that fed on fishes and
squids (coleoid cephalopods) there.
5. New discoveries from China (in 1996) confirm that birds are derived from maniraptoran theropods
and that feathers evolved in the earliest coelurosaurs (Benton 2005). But these were non-flying
animals,where the function of feather is not yet clear. It was only in Archaeopteryx, the first bird, that
the feather helped in flying.
6. Dinosaur relationships may be shown in the cladogram (Figure 18.6).
7. Dinosaurs are divided into two groups; the Saurischia, (reptilian/lizard hip) and the Ornithischia (bird
hip), on the basis of their radically different pelvic girdles. In fact, hindlimbs are now considered
important in characterizing Dinosauria as also the clades within Dinosauria.
Saurischian structure is more primitive, with pubis pointing forwards and ischium back. It is found
in all basal archosaurs. Ornithischians have pubis running backwards parallel with the ischium; there
is an additional prepubic process in front.
Characters of hindlimbs are related to the acquisition of upright posture.The ankle and foot are also
distinctive. Dinosaurs have digitigrade feet where the animal stands on its toes, and not on the flat of the
whole foot (plantigrade). Dinosaurs really uses three middle toes, second, third and fourth. Saurischians
include carnivorous theriopods as well as large herbivorous sauropods.
Ornithischian characters of pelvic girdle, skull and skeleton are derived. It arose in Late Triassic
(Carnic), rare until Jurassic; they were all herbivorous and divided into two groups: Cerapoda (bipedal
ornithopods, boneheaded and horned forms) and Thyreophora including ankylosaurs and stegosaurs.
8. The raging debate on whether dinosaurs were warm-blooded or not is yet undecided. Evidences in favour
of cold/warm-blooded nature are put forward, only to raise questions, even questioning methodology.
Below is provided a small list of them, only to keep the debate live.
Endotherms or warm-blooded animals control body temperature internally. Ectotherms or cold-blooded
animals rely on external sources of heat.
l Erect gait and ability for speed (like birds/mammals) suggest endothermy; but link between
endothermy and erect gait is not clearly demonstrated and speed estimates range from 6 to 60 km/h;
high speed may have been typical of only small bipedal forms.
l Haemodynamics (or blood circulation): For long-necked sauropods to pump blood to face and brain,
four-chambered heart like that of birds and mammals would have been necessary; but crocodilians too
have a four-chambered heart.

(Cont...)
 Chapter 18 Vertebrata of Chordata 295

FACTSHEET 18.2 (Cont...)

Dinosaur Information: Summary of Details

l Bone histology of dinosaurs reveals highly vascular bones and Haversian system of pattern like mam-
mals; but Haversian system can occur in modern reptiles, and many small mammals and birds have no
Haversian system.
l Fibrolamellar bone found in fast-growing mammals and some birds is also found in many dinosaurs; but
that may indicate only fast growth and no endothermy.
l Lamellar-zone bone with growth rings (lines of arrested growth during limited food supply or adverse
climate) found in modern reptiles, crocodilians and many dinosaur groups. A mixed thermoregulatory
pattern with both fast and episodic growth is envisaged.
l Birds and mammals grow fast, whereas reptiles grow slowly. Dinosaurs seem to have been fast growers
implying endothermy.
l New finds of feathers in some theropods might have been for insulation implying endothermy.
Small differences between core and peripheral body temperatures measured from O18/O16 ratio of
core bones (ribs, dorsal vertebrae) and peripheral bones (tail, limbs) were used to point out con-
stancy of body temperature, but here again interpretation is questionable.
l The absence of thin bone in the nasal cavity to absorb water from the air coming in, in modern
reptiles and dinosaurs go against endothermy.
l Herbivores (ecto-/endothermic) can support ~5 per cent of their biomass of endothermic predators,
about 30-50 per cent of ectothermic carnivores; if the predator ratio is greater it tells upon the
herbivore population; predator-prey ratio in early Permian is calculated at 50-60 per cent, late
Permian 10 per cent and 2-3 per cent in late Triassic, Jurassic, Cretaceous; this is held to suggest
endothermy of Mesozoic predators, the dinosaurs.
l Present opinion favours:
Small dinosaurs may have been endotherms with fibrolamellar bone or feather.
Large dinosaurs were probably inertial homeotherms or gigantotherms that had constant body tem-
perature (within 1°C or 2°C) by virtue of being large with very slow rates of internal temperature
changes during ambient temperature fluctuations.

case histories may add lively tones to the routine
classroom materials. Once more it must be added
that this is more a story-telling to hold the readers
back to the issue for some more time. Who knows,
one or two of them may pick up clues and initiate
more revealing research thereupon!
Dinosaur eggs from China: Dinosaur egg fossils
are, in most cases, limited to the egg shell. The
embryos themselves are rare, though only such
fossils in relatively advanced stages of
development, can help us associate dinosaur types
with fossilized eggs and egg shells.
Recently discovered remains of a female
dinosaur with two complete eggs preserved inside
the mother’s body, recovered from the Jiangxi
province of China elucidate the ways in which
these animals gave birth (Sato et al., 2005). The
pelvic and leg fragments are from a dinosaur, which
probably measured between 10 and 13 feet,
belonging to a group known as the ovira-
ptorosaurians. The larger egg is nearly 18
centimetres long and between 6 and 8 centimetres
wide. Modern female birds can only produce one
egg at a time because they have only one ovary
and oviduct, whereas in these dinosaurs similar
296 Part Three: Miscellaneous
sized eggs suggest that the creature’s two oviducts
simultaneously produced an egg each. The laying
of the eggs, however, appears to have been
distinctly birdlike. Dinosaur nests were earlier
reported from the same time period that contained
up to 15 eggs. That, together with the positioning
of the eggs, suggests that the dinosaurs would have
repeated the egg-laying process a number of times
to attain a full clutch just as modern birds do.
Reptiles, in contrast, lay their entire clutch at once.
The results provide further evidence that
dinosaurs share some aspects of reproductive
behaviour with birds, strengthening the belief that
modern birds are their descendants. In addition,
the find helps explain why the eggs in other
oviraptorosaurian nests appeared to be arranged
in pairs.
More egg types and embryos from late Cretaceous
of China: Shell fragments and their thin section
analysis suggest four main parataxonomic families
of eggs found in late Cretaceous of Henan and
Nanxiong Basins of China. They differ in shape,
size, structural morphotypes and pore systems, etc.
These are: (1) Faveoloolithidae (large, thickest,
without any continuous layer at the surface of the
shell; they are also the oldest and may represent a
hot and humid climate); (2) Dendroolithidae (with
a continuous layer at the surface; related possibly
to the taxonomic group Ornithopoda); (3) Spheroo-
lithidae (smaller, thin shelled); and (4) Elongatoo-
lithidae (elongated; includes the largest type found
to date; nests of Elongatus eggs indicate the eggs
were laid two at a time, the pairs being generally
oriented at 40 degree angles from the next nearest
pair forming a radial spiraling nest with each
successive layer buried with a layer of sand or mud)
(Zhao 1975, 1994).
Recent discoveries of embryos in one type of
Spheroolithid eggs indicate they are from a
Therizinosaur called Segnosaur; embryonic bones
are preserved on account of calcite replacement.
Therizinosaurs (Scythe reptile) were named for
their large bony claws. An aberrant group of
theropod dinosaurs which, based on tooth structure,
appear to be vegetarian, yet sport a most formidable
hand claw. One of the claws from the hand found
in Asia measures 28 inches and this does not
include the horny part of the claw which would
have made it even longer! Just what these claws
were used for is a mystery.
The embryo found inside one of the elongated
eggs has been identified as an oviraptor
(Egg Thief). The skull of one of these dinosaurs
was found crushed and on the top of a nest of what
was long believed to be a nest of Protoceratops
eggs. Osborn speculated that the skull was crushed
by an enraged Ceratopsian parent who caught the
unfortunate raptor in the act of stealing eggs. There
have also been evidences of a mother Oviraptor
shielding her nest, a caring parent defending
her nest !
Egg shell and climate: Studies of changes in egg
shell thickness over time show a gradual thinning
of the shells. Samples of pollen found in these
deposits show a concurrent change in vegetation.
This evidence shows a gradual change in climate
from humid subtropical to an arid temperate one
(Zhao and Yan 2000). Thinning of egg shells due
to environmental stresses in modern birds like the
brown pelican has been shown to significantly
reduce bird egg viability and could lead to eventual
extinction of the species. Changes in climate and
vegetation could have weakened the dinosaurs
ability to produce viable eggs and, hence, could
have led to their eventual extinction.
Percentage of oxygen in the atmosphere
measured in air trapped in amber went from a high
of 35 per cent to a low of 28 per cent, some 120
million years ago (Early Cretaceous) diminished
significantly over a period of 5 million years.
(G S A meeting, 1993). This may have a cause and
effect relationship with the gradual changes in shell
thickness observed by the Chinese scientists.
Pterosaur eggs: Three fossilized eggs of the
ancient flying reptiles, pterosaurs, two reported
from China (2004) and the third from Argentina
(Chiappe et al., 2004) resolve the debated point

whether pterosaurs laid eggs or gave birth to live
young one like mammals do. They also yield clues
to how pterosaurs lived.
The egg from Argentina is from Lower
Cretaceous, obtained from beds in association with
abundant fossils of Pterodaustro pterosaur and a
few fish fossils. The egg is thought to have had a
hard shell similar to those of birds and dinosaurs.
Chiappe identified the egg as belonging to the
flamingo-like Pterodaustro guinazui. The discovery
confirms the species’ communal lifestyle. It was
found amongst a bunch of other fossils of the same
species, fossils ranging from hatchlings to teenagers
to full adult individuals. That tells us that these
animals nested in a community and offered to their
young parental care.
Qiang Ji et al., (2004) described the new egg
from China. The egg is Lower Cretaceous in age
and is reported to have had a ‘soft and leathery’
shell similar to that of crocodile and turtle
eggshells. It belonged to Beipiaopterus.
Group living of Argentinian pterosaurs: Chiappe
et al., (2004) concluded that the Argentinian site
was unsuitable to all but this flamingo-like
pterosaur. The area was perhaps filled with
extremely salty lakes similar to those inhabited by
flamingoes high in the Andes mountains, Chiappe
speculated. Pterodaustro was a filter feeder,
just like the flamingo. He found the fossilized
Pterodaustro embryo among hundreds of other
fossils, ranging from young juveniles to full-grown
adults. Chiappe said that this suggests that the
reptiles nested in a colony and protected their
young one. Both Chinese pterosaur fossil embryos
are of similar age and were found in similar
environments. According to Bennett, this suggests
that a number of different pterosaurs may have
nested around this ancient lake-filled Chinese
environment.
Coprolites, etc. of fishes and reptiles: Coprolites
and fossilized gut contents are reported from the
Smoky Hill Chalk of Cretaceous age. The animals
are identified as sharks, teleost (bony) fish and
Chapter 18 Vertebrata of Chordata 297
marine reptiles. Coprolites may can tell us about
what kind of prey these animals ate and how their
digestive systems worked. Buckland (1829), Dekay
(1830), Mudge (1876) and Williston (1898)
(in Everhart 1999) were among the first to discuss
the presence and origin of coprolites in the Smoky
Hill Chalk.
Coprolites are fairly common and contain
partially digested bones (usually fish).  Sharks are
the source of the spiral coprolites, but most of the
others are more likely to be from marine reptiles
and large fish. Fossilized gut contents (Miller 1957,
Stewart 1978) and regurgitated prey material
(see Hattin 1996, Everhart, 1999) are the two other
categories of unusual biological materials
preserved in the same rocks.
18.11 Extinction of Dinosauria:
Reality of a Myth
Extinction of dinosaurs, the animals which reigned
over the surface of the earth for nearly 200 million
years with so many varieties of adaptations
producing comaparable number of kinds, has
haunted laymen as well as specialist
palaeontologists since long. Particularly intriguing
about it was the sudden and massive character of
the extinction event which coincided with the end-
Cretaceous (otherwise referred also as KTB, i.e.
Cretaceous-Tertiary Boundary) mass extinction.
The latter also brought about extinction of several
other groups of animals and plants (another
celebrity victim of this extinction was the
ammonoids). Numerous causes have been
advocated to explain such mass extinctions, though
many of them would appear to be wild guesses
even to laymen (see Factsheet 18.3).
18.11.1 Gradual or catastrophic
Scanning the list will, however, reveal that the
causes can actually be grouped into two categories
or general kinds of ‘causes’ or ‘theories’ or
‘models’. In one, the extinction, though massive,
298 Part Three: Miscellaneous

FACTSHEET 18.3
Suggested Causes or ‘Theories’ of Dinosaur Extinction (Based on Charig 1991)

EXTRINSIC: Events or changes in the external environment, sudden or gradual.

Extraterrestrial: Impact of meteorites, asteroids, etc. showering of high-energy and /or charged
particles from explosion of stars or other sources, passage of the earth through
interstellar dust clouds, etc.
Physical: Perturbations in gravitational constants, earth’s magnetic field, eccentricity of
the earth’s orbit, the position of poles.
Geological/geographical: Earthquakes, effusion and emission of volcanic dust and ash, lava, etc. continental
drift, mountain building, marine transgression or regression, floods, draining of
lakes and swamps, etc.
Hydrospheric: Sudden or gradual changes in salinity, acidity, etc. of sea.
Atmospheric: Increase in oxygen, acidic or poisonous ingredients; decrease in ozone layer,
changes in pressure.
Climatic: Sudden/gradual increase/decrease in temperature, humidity and precipitation,
etc.
Nutritional: Shortage of food due to changes in flora (rise of angiosperms and depletion of
pteridosperms, gymnosperms, etc.); due to competition with herbivorous animals;
due to rise of plant-eating caterpillars; due to lack of specific ingredients
(vitamins, etc.).
Pathological: Protozoan, bacterial or viral infections of terrestrial/extraterrestrial origin.
Other ecological: Faunal imbalance, inter necine fights, overcrowding, ‘overkill’ by predators.
INTRINSIC: Events or changes arising within the organisms themselves.
Disorders and diseases: Anatomical (vertebral slip-disc from overweight), metabolic (from gigantism
causing endocrinal/hormonal imbalances), neurological (dwindling of brain),
etc.
Genetic changes: Evolution into racial senility and over-specialization.
Reproductive: Eggshell thinning/thickening, various problems in copulation, etc.
INTRINSIC TRIGGERED
BY EXTRINSIC: Events or changes in the organisms, caused by extrinsic factors.
Physiological: Endo-/ectothermy, digestion, excretion, etc.
Reproductive: High or low ambient tempretures causing imbalance in sex ratio, sterilization, etc.
Genetic: Gene pool drainage (by mammalian egg-eaters).

is viewed as an outcome of a gradual process—
‘gradualistic model’ and in the other the trigger
is a geological or biological catastrophe —
‘catastrophic model’. Incidentally, knowingly or
not, this was an outcome of the influence of two
different philosophies or schools of thoughts, that
marked the history of development of geological
thoughts in the eighteenth and nineteenth
centuries.
The debate over the causes of dinosaur
extinction took a different turn in 1980, when the
American physicist Luis Alvarez and his geologist
son Walter declared dinosaur, or rather KTB mass
extinction to be the result of an asteroid or bolide
impact. Almost simultaneously, another group of
scientists attributed a massive volcanic activity, that
included the Deccan Trap volcanism in India, as
the cause of KTB mass extinction. Both the views,

however, hinged on one or another kind of
catastrophic event.
18.11.2 Bolide impact theory
The asteroid or bolide impact theory envisaged
an asteroid (coming from the asteroid belt between
Mars and Jupiter and falling on to the earth) about
10 km in diameter to have reached the earth at the
KTB. (There are variations in opinions about the
timing and duration of this asteroid shower). The
tremendous speed of the impact released huge
kinetic energy; it pulverized and vaporized matters
involved, threw back into the atmosphere a huge
mass of dusty matter that made a thick cloud above
the earth, taking months or years to settle down
and, thus, preventing sunlight to penetrate during
that period of time. This, in turn, brought about a
long spell of cold winter and arrested
photosynthesis. Plants were mortally affected, and
with that the food chain was shattered. Dinosaurs,
many of which were massive and, thus, dependent
on huge herbivorous diet or carnivorous, feeding
upon their herbivorous brethrens, might have been
strongly affected. The released energy from the
impact also generated heat that could have led to
high temperature reaction between oxygen and
nitrogen of the atmosphere to form nitrous oxide;
this gave way to acid rain, when the nitrous oxide
combined with water to produce nitric acid
aerosols. The process also destroyed the ozone
layer. The acid rain must have been equally
destructive to land-vegetation as also to
phytoplanktons and calcareous skeleton-bearing
microorganisms of the surface layers of seas, telling
upon the food chain there. In summary, the asteroid
impact brought about a devastating ecological
disaster that left its imprint on the then organic
world through the mass extinction at that juncture
of the earth’s history.
Alvarez and Alvarez built their explanation
on materials collected from the Gubbio sections
of Umbria region of Italy. There in an undisturbed
section at roadside cuttings are beautiful limestones
Chapter 18 Vertebrata of Chordata 299
used as decorative building stones. They included
certain foraminiferal limestones overlain by a 2
centimetre thick layer of dark clay; this is again
overlain by another set of limestone layers entirely
different in character and fossil content from the
underlying ones. It was a KTB section, as could
be known from the fossils and other evidences.
On precise chemical analyses (at parts per billion
level of measurement), this clay layer showed a
value of an element, iridium, that was abnormally
high. Such high iridium value was uncharacteristic
of the crust and was to be found in meteorites or
asteroids as well as mantle materials. This iridium
anomaly led to recognizing the clay layer as
meteoric or asteroid origin. There were other
evidences of impact: shock quartz or percussion
like features which could only be produced under
impact. A subterranean crater-like structure, 200
km in diameter revealed by anomalies in the
gravitational and magnetic fields near Chicxulub
village of Yucatan area of Mexico, was suggested
as the spot where the asteroid had hit the earth.
Alvarezs’ paper appeared in the journal
Science in 1980. Since then, there was a remarkable
surge of enthusiasm on the related topics, either
supporting or opposing the theory of bolide impact.
In 1984, Raup and Sepkoski even suggested a 26
million years periodicity of extinction of species
implying periodic impact of asteroids.
But the theory also invited strong criticisms.
In the main, it was contended that invoking
extraterrestrial cause for the iridium anomaly at
the KTB was hypothetical; searches should better
be made for more earthly causes. Besides, other
evidences were not unequivocal. Some of the
critics drew attention to the fact that right at this
KTB, there was a massive volcanic activity that
produced the extensive basaltic lavas now
occurring exposed over a large part of the present-
day Peninsular India. These lavas, as their
composition and settings indicate, might have
been derived from the upper parts of the earth’s
mantle and would thus, have been responsible for
the iridium anomaly. Such a volcanic activity must
300 Part Three: Miscellaneous
have had some other necessary global effects,
quite similar to those produced by the suggested
bolide impact.
18.11.3 Volcanism theory
Thus, some authors emphasized that such huge
volcanism must have injected ten times more
carbon dioxide into the atmosphere than the
atmosphere contains today. Oceans were warmer
than they are now and so could not dissolve this
massive amount of gas. Rather the gas
accumulated in surface waters and the
atmosphere. This hindered photosynthesis and
affected life of phytoplanktons and the food chain
too. In addition, excess carbon dioxide also
hampered production of calcium carbonates in
more than one way, chemically or biologically.
Single-celled algae or other organisms in seas and
oceans fix up a fair lot of carbon dioxide to build
their skeleton of calcium carbonate. When they
were exterminated due to the reduction of
photosynthesis and increase in acidity and other
types of toxicity of sea water from various
volcanic emanations, huge amount of carbon
dioxide could not be fixed up in calcium
carbonate. Carbon dioxide content was further
increased, leading to a greenhouse effect and
considerable rise in temperature.
Other scientists, however, suggest a different
scenario, much similar to that of the impact event
at some point or other. They argue the thick haze
caused by volcanic emanations (gases and volcanic
ash) prevented the passage of sunlight to cause a
prolonged winter. Also gases like sulphur dioxide
combined with water to form tiny droplets of
sulphuric acid. Dispersed throughout the upper
atmosphere they caused cooling, a partial
destruction of the ozone layer and, above all, acid
rain. In fact, recent studies on the chemistry of the
gas-bubbles and water caught up in polar ice caps
suggest that the sulphur content of an eruption may
have the determining effect of climatic impacts.
The mean atmospheric temperature decreases as a
function of the amount of sulphur coming out of
volcanic eruptions in the historical times and the
probable climatic effects (in terms of decrease of
mean atmospheric temperature) of much larger
volcanisms of the ancient times may be extra-
polated from these data of historical examples.
18.11.4 Debate changes to ‘how
catastrophic the event was’
In other words, the debate over dinosaur
extinction through ‘gradualistic’ or
‘catastrophic’ events took a rather different turn
in the last two decades of the twentieth century.
It came down to a heated tussle between
proponents of ‘asteroid impact’ and a ‘huge,
global volcanism’, both theories of catastrophic
events. The bone of contention seemed to have
changed at finding out how catastrophic the
event was. Asteroid impact was, no doubt, short-
lived. Different authors shared this view. One
measure from the amount and the constant rate
of accumulation of helium-3 from KTB rocks
of Italy and Tunisia suggested accumulation of
high iridium clays in only 10,000 years, as a
result of one single impact.
The proponents of the ‘volcanic’ theory, on
the basis of the same helium-3 measured from
microscopic bubbles caught up in the quickly
solidified basalts of the Deccan Traps, also
inferred that the Traps were erupted between 68.5
and 64.9 million years ago, though most of the
lava flows came up in about 0.5 million years,
sometime between 66 and 65 million years ago.
A French team of scientists working on
palaeomagnetism reached the same conclusion of
eruption having taken place about 0.5 million
years ago.
18.11.5 New facts and views
However, as discussed in Factsheet 18.2, at least
some dinosaurs are now considered as warm-
blooded, active, terrestrial animals like mammals
or birds rather than hybernating, semi-aquatic
 Chapter 18 Vertebrata of Chordata 301

reptiles. The relationship between dinosaurs and
birds are also better established. Phrases like ‘birds
are really the dinosaurs surviving today’ have also
come up in the literature. It brings many of the
suggested causes of dinosaur extinction, to
question.
There have also been changes in scientists’
views on sudden, all-out mass extinction. It is now
recognized that there may be at least three different
kinds of mass extinction scenarios. In one, a graded
mass extinction, species may have disappeared and
appeared regularly one after another. In a second
catastrophic mass extinction, there is an
instantaneous extinction of numerous species,
followed by a gradual reappearance of new species.
In the third type, there may be episodic rapid
succession of several events of extinctions, each
less intense than that of the catastrophic type
(Figure 18.6).
On reappraisal of fossil and sedimentary
records, it is also being accepted more and more
that even these three types of mass extinction may
be some apparent pictures of reality, being results
of a complex interplay of different factors.

(a)

G
F
E

(b)

(c)

Fig. 18.6 Alternative explanations of mass extinction.

(a) Mass extinction at one level (E) (catastrophic), (b) Mass extinction through a brief period at
different levels (E,F,G) (stepwise), (c) Mass extinction through a brief period (E to F) (Graded).
Vertical lines indicate vertical ranges of taxa involved (commonly genera or species); horizontal
lines show the level/levels of extinction.
302 Part Three: Miscellaneous
Firstly, the scenarios are reconstructed on fossil
records, which, in their turn, are largely controlled
by various taphonomic factors. Dinosaurs being
terrestrial organisms will stand less chances of
fossilization than marine ones. The record is often
very scanty, as we find in India. Besides, the
absence of fossils of any group in a record does
not necessarily mean that those organisms were
not present at that time; they may not have been
preserved or simply their fossils may not have so
far been discovered.
Secondly, sediments which are the most
common media that preserve fossils do not
accumulate at the same rate in different basins, nor
at different times in the same basin. Neither is
sedimentation a continuous process. Thus, there
may be sudden rapid influx in an otherwise slowly
accumulating process, for example, during a flood,
landslide or turbidity current, etc. On the contrary,
sedimentation may be considerably slowed down
or stopped. In the former case, organisms living
during rapid sedimentation or the slower one may
not be equally represented in the fossil record. In
the latter case, organisms of different successive
times will be condensed in the layer of sediments
that have been formed slowly over a long period
of time. Added to this, there are the effects of
erosion, that may wipe out the record of some
period of time completely or partially. Erosion may
also mix up the fossils of different times to be
reworked and redeposited in a younger layer. In
summary, fossil record seldom gives a continuous,
uniformly represented picture of the ancient
organic world. Even, a sequence of layers thought
to be continuous may prove to bear discontinuities
hitherto unknown or detected with more detailed
and improved studies. Inferences based on such a
record may then include uncertainties, over-or
under-representation, gaps, etc.
These are a few questions among many more
such, which reorient, in a sense, the ideas on
dinosaur extinction. There are reports, though
challenged for authenticity and thus requiring
checkup, of dinosaur fossils coming from rocks
younger than the KTB point of the geological
column, i.e. from Palaeocene rocks. From India,
too, such reports are available from occurrences
near Anjar in Kachchh in Gujarat. Closer
phylogenetic relations between birds and dinosaurs
have even prompted people to wonder “If birds
did evolve from dinosaurs, they would be members
of the dinosaur’s evolutionary lineage . Therefore,
dinosaurs would not really be extinct ... at least
not yet” (Dingus and Rowe 1997).
From the history of ideas on dinosaur and their
extinction, we may, thus, conclude as follows.
Dinosaurs and their contemporary world provide
ample proof of intricate ramifications in adaptation-
evolution-extinction scenario. The impact or
volcanism at the KTB may have had profound
effects certainly on the global environment. But
that may not make the impact or volcanism the
general or sole cause of extinction or mass
extinction. At the KTB many dinosaur genera and
species became extinct, but many others were
extinct earlier. Obviously they failed to adapt to
their respective contemporary environment. But it
is equally acceptable that there might have been
other dinosaur genera and species which did not
fail in adapting themselves and might have had
radically changed through that process into newer
kinds of organisms, for example, the birds. Besides,
on land, dinosaurs and other great reptiles as well
as large plants of the groups like pteridosperms,
gymnosperms, etc. and in seas ammonoids and
certain other groups were extinct at the KTB. But
mammals that had appeared earlier and were
evolving with an improved method of reproduction
and a more efficient metabolic and thermo-
regulatory mechanism of the body successfully
crossed the hypothesized impact or volcanic
calamity and radiated immediately after. In
addition, some other different groups of organisms
that made their appearance on the earth in late
Cretaceous time immediately before the KTB, like
angiosperms or flowering plants on land or
planktonic microorganisms as also echinoids and
corals with hard calcareous skeletons in seas,

successfully radiated by way of varied adaptations
immediately after the KTB. Of them, angiosperms
appeared with a more developed mode of
reproduction and propagation than those of any
other earlier land-plant group, and the hard
calcareous skeletons of the new zooplanktons and
their bottom-dwelling kins served more efficiently
as skeletal framework or for protection and other
vital functions of life, being perfectly adapted to
the physico-chemical conditions in seas that had
started to develop with modern trends before and
after the KTB. Hence, to hold catastrophe as the
cause of extinction, may really be putting one-sided
weightage on the ‘environment’ aspect of the
evolution-extinction process. It requires to be
appreciated that, catastrophe or not, extinction has
always been a part of organic evolution, a
particular aspect of the total evolutionary process
and phenomenon, often adding new vistas to
evolution as a whole. As and when dinosaurs
became extinct, a vast and varied environment
with comparable number of niches on the surface
of the earth was laid bare to land vertebrates, to a
relatively smaller, yet more developed group of
mammals. The latter found it favourable for its
phenomenal adaptive radiation that produced
much of the present-day land vertebrate fauna,
including our own selves.
18.12 Origin of Mammals
Living mammals are characterized by the following
set of characters:
1. Presence of hair
2. Large brains
3. Feeding of young from milk of mammary
glands of mother
4. Post-natal care for young for a considerable
period of time.
Mammals originated in Late Triassic, when the
distinction between mammals and non-mammals
was not great. A succession of Triassic carnivorous
Chapter 18 Vertebrata of Chordata 303
synapsids, the cynodonts, successively acquired
‘mammalian’ characters over a time span of 30-40
million years. The exact point of origin of mammals
is, however, not clearly defined. The first fossils
are incomplete. Adelobasileus, from Upper Triassic
of Texas, USA and Sinoconodon, from Lower
Jurassic of China might have been the most basal
(primitive) mammals. But small, shrew-like
morganucodontids of early Jurassic of Europe,
North America, China and South Africa present the
better-represented forms.
Traditionally on evidences of molars and
brain, mammals were thought to be diphyletic,
one leading to ‘therians’ (including Kuehne-
otherium, the symmetrodonts, marsupials and
placentals) and the other to be the ‘prototherians’
(morganu-codontids, tricodonts, docodonts,
multituber-culates and monotremes). Presently,
however, on cladistics, Mammalia is considered
monophyletic.
18.13 Vertebrates and
Palaeogeography-
Palaeobiogeography
It has been indicated earlier that the distribution
of Permian and particularly Triassic reptiles like
Mesosaurus and Lystrosaurus-Cynagnothus,
respectively help in reconstruction of the super-
continent Pangaea (Figure 18.7). Since the present-
day continents were then agglomerated into one
single supercontinent, the dispersal or rather
migration was not a problem for these large land
vertebrates.
Mammals present a slightly different story in
regard to their biogeographic importance. They were
evolving at a time when the supercontinent had been
fragmented and the continents were attaining their
present shape and position through rifting and
drifting. There were continental connections facing
breaches or continents becoming locked by seas on
one or several sides; on the other side, there were
new land bridge connections forming between two
304 Part Three: Miscellaneous
Mesosaurus Cynognathus
AFRICA
SOUTH
AMERICA
Fig. 18.7 Reconstruction of Gondwanaland on fossils.
continents.The separation of Antarctica and
Australia into virtual island position provides
examples of the first type; important instances of
the second are found with the Asia-Africa
landlocking and with the classic example of land
bridge between North and South America. These
developments naturally told upon vertebrate, rather
mammal distribution. Isolation led to endemism;
new land bridges supported migration.
For most of the Cenozoic, Australia and South
America were islands, sustaining geographically
restricted endemic mammalian faunas very much
different from those of other parts of the world.
Thus, South America had its own marsupial
mammals that looked like dogs, bears, sabre-
toothed tigers, rodents, ungulates like horses,
rhinoceroses, armadillos, sloths and others of the
northern continents, but were never the same or
Glossopteris
INDIA
Lystrosaurus
ANTARCTICA
AUSTRALIA
related. The KTB extinction affected the basal
mammalian stocks and particularly the marsupials
elsewhere in the world. In South America, however,
they were spared. That evolved into the specialized
fauna of the continent. They were joined by ‘waifs’
(poor, homeless children) from outside, which
consisted of small populations of a few groups of
placental mammals (for example, rodents in
Eocene, bats and primates in Oligocene) that
arrived at the continent by chance dispersal events
(Benton 2005). But biogeographically more
significant was the Great American Interchange
(GAI) that took place after the two continents were
connected by the Central American land bridge
connections. It affected emigration of opposums,
armadillos, ground sloths, etc. from South
America and immigration of horses, camels, deers,
mastodonts, etc. from North America. In both the

continents the situation was comparable. In both
the cases, some of the immigrants evolved and
many others became extinct. In South America,
however, the total number of mammal genera
increased markedly after the land bridge between
the two continents was formed. A comparable land
bridge connection between Alaska and Asia
established a migratory route at the northern end
of North America. Horses (equids) of China and
Indo-Pak subcontinent were introduced via this
route, along which even migration took place in
phases. Hipparion in Mio-Pliocene and Equus in
Pleistocene of the Siwalik Group of Formations in
India and Pakistan bear testimony.
It has been indicated in Chapter 5 that studies
on molecular evidence have now brought out the
importance of the role of biogeography in
controlling relationships of early placental
mammals. Though there is a debate on whether
molecular evidence reflect true picture or whether
eutherians arose in southern continents or northern
ones (see Hunter and Janis 2006), it is quite certain
that their distribution was later attained through
dispersal or migration from their point of origin.
By way of this distribution pattern, there developed
different distinct clades (monophyletic groups),
Afrotheria in Africa by the end of Cretaceous and
Xenarthra in South America by Palaeocene.
The African clade with common shared
derived characters included as different groups as
elephants (proboscideans), golden moles, tenrecs
and aardvarks (the latter three are insect-eating
mammals). Xenarthra included typical armadillos,
sloths and anteaters of South America.
After the divergence of Afrotheria and
Xenarthra, the stem of placental mammals gave
rise to Boreotheria which later diverged into two
sister groups Laurasiatheria and Euarchontoglires.
The former includes Chiroptera (bats), Artiodactyla
(cows, goats, giraffes, hippopotamuses, pigs, etc.),
Cetacea (whales), Carnivora and Perissodactyla
(horses, rhinoceroses, tapirs) and others. The latter
includes Primates, Rodents (rats, etc.), Lagomorpha
(rabbits, etc.), etc.
Chapter 18 Vertebrata of Chordata 305
Development of these African and South
American clades, indicate that whatever may be
the directions of dispersals, land connections
between Africa and Eurasia on one hand, and
between North and South America on the other,
were established in the end Mesozoic and the
earliest Cenozoic, respectively, which governed
distribution and subsequent evolution of mammals.
18.14 Indian Records
18.14.1 Non-mammalian vertebrates
In India, important records of non-mammalian
vertebrates, viz. the fishes, amphibians and reptiles
come mostly from the Palaeozoic-Mesozoic
successions of the Gondwana Group (/Supergroup)
of formations, a Permian Mamal Formation of
Kashmir, the Infra- and Intertrappean beds of the
Peninsular India. There are reports of these
vertebrates also from different Cenozoic units, in
which these groups are overwhelmed by the
mammalian counterparts. (see Factsheets 18.4 to
18.7) Of these, the Gondwana occurrences are the
best known and studied. So, they are being treated
separately in the next section.
18.14.2 Gondwana vertebrates
Gondwana Group/Supergroup of formations is a
huge thickness of essentially continental sediments
of Permian to Lower Cretaceous age. It is
characterized by dominantly siliciclastic rocks
(significant carbonates are found only in Jurassic
Kota Formation), with India’s richest coal reserve
and carbonaceous shales associated with them.
It is further characterized by typical terrestrial
floras (Pteridosperms and others in the lower parts
and cycadeoids and conifers in the upper parts; see
Chapter 19 for more details) and vertebrate faunas
(mainly amphibians and reptiles); fishes in
Gangamopteris Beds (now called Mamal
Formation) in Kashmir and Kota Formation in
Pranhita-Godavary (PG) Valley; micromammals
(in Kotas). Gondwana sediments were deposited
306 Part Three: Miscellaneous

FACTSHEET 18.4
Vertebrates from Indian Gondwana

Composition
Fossils occur in (i) 9 major faunal horizons between Permian and Jurassic and belong to
7 families of fishes, viz. chondrosteans, neopterygians, sarcopterygians and
elasmobranches
10 families of temnospondyl labyrinthodont amphibians,
26 families of anapsid, diapsid and synapsid reptiles, and
4 mammalian families, two therians and two nontherians;
and, (ii) lower Cretaceous coastal Gondwana equivalents.
Major host formations are:
Late Permian: Mamal Formation in Kashmir and Kundaram Formation in Pranhita-Godavary
(PG) Basin.
Lower Triassic: Panchet Formation in Damodar Valley basin.
Middle Triassic: Denwa Formation in Satpura basin and Yerrapalli Formation in PG Valley basin .
Late Triassic: Tiki Fmn in Son-Mahanadi basin and Maleri and Dharmaram Fmns. in PG Valley
basin.
Lower to Middle Jurassic: Kota Formation in PG Valley basin.
Lower Cretaceous: Raghavapuram Mudstone in Andhra Pradesh (with fish) and Umia Fmn in
Kachchh (with marine reptile).
Significance:
l Vertebrates show close relationships with Laurasian or Gondwanaland form, attesting to that India was an
integral part of that Pangaea supercontinent from Permian to the end of middle Jurassic.
l Early Permian fauna is absent indicating a climate hostile to vertebrates but congenial for a luxuriant flora.
The absence of herbivorous vertebrates meant minimal foraging by them; that caused uninhibited growth of
the flora which, in turn, could produce extensive coal deposits.
l Late Permian archaegosauroid amphibians of the Kashmir fauna, a typical lowland, aquatic fauna, may be
Laurasian in origin which migrated along the Tethyan coastline into India.
l Subsequent Indian, as well as other Gondwanaland fauna of late Permian assumed provincial characters.

l Early Triassic Panchet fauna was one of the most widely distributed terrestrial faunas of the Pangaea with
Lystrosaurus, Indobrachyops and others. It closely resembled the fauna of the Lystrosaurus Assemblage
zone of South Africa. It was also a lowland fluviatile and lacustrine type that flourished following the Permian
extinction event.
l Kota fauna consisted of:

(a) Upper horizon in limestone dominated lacustrine sediments, with fishes (Lepidotes, Paradapedium and
Tetragonolepis, a coelacanth, etc. resembling Liassic of Germany) and reptiles including a
crocodylomorph, a flying reptile pterosaur (Campylognathus indicus) and a kind of turtle.
(b) Lower horizon in a fluviatile sandstone-mudstone association with sauropod dinosaurs (Barapasaurus
and Kotasaurus) and five mammals (micromammals). The different genera and species are
palaeogeographically significant as they resemble those from Europe, Asia, Africa and America. The
dinosaurs also throw important light on the evolution of sauropods.
l Clupavas neocomensis of Raghavapuram Mdst. is a marker of Neocomian that help in fixing upper age limit
of the Indian Gondwanas and Plesiosaurus indicus of Umia Fmn. is the only marine reptile in India.
 Chapter 18 Vertebrata of Chordata 307

FACTSHEET 18.5
Vertebrate Record of Major Indian Basins and their Formations
Mahanadi Basin
Middle Jurassic
Early Jurassic
Late Triassic Amp: Buettneria maleriensis; Rep: three diapsids, a phytosaur, a rhynchosaur and
Middle Triassic Tikisuchus (identical to Lower Maleri fauna).
Early Triassic
Late Permian
Early Permian
Satpura Basin
Middle Jurassic
Early Jurassic
F: Ceratodus sp.; Amp: Parotosuchus; Stenocephalosaurus;
Late Triassic
Rep: undescribed species and genus.
Middle Triassic
Early Triassic
Late Permian
Early Permian
Damodar Basin
Middle Jurassic
Early Jurassic
Late Triassic
Middle Triassic
Early Triassic Panchet
Late Permian Raniganj
Amp: Lydekkerina panchetensis; Pachygonia incurvata, Indobrachyops
Early Permian Barren panchetensis, Gonioglyptus longirostis; Rep: Proterosuchus indicus,
Measures Different species of Lystrosaurus, Chasmatosaurus
Barakar
Talchirs

Pranhita-Godavary (PG) Valley

Middle Jurassic Yamanpalli -----
Early Jurassic Kota F: Lepidotes, Paradepium, Tetragonolepis; Amp:-; Rep:
Indochelys, Campylognathus, Barapasaurus tagorei,
Kotasaurus yamanapalliensis; Mm: Kotatherium,
Indotherium, Trishulotherium.
Late Triassic Dharmaram F: Ceratodus, Xenacanthus; Rep: phytosaurs, prosauropods,
plateosaurids.

(Cont...)
308 Part Three: Miscellaneous

FACTSHEET 18.5 (Cont...)

Vertebrate Record of Major Indian Basins and their Formations

Maleri F: Ceratodus (several species), Xenacanthus; Amp:

Buettneria, Compocerops, Kuttycephalus; Rep: Paradapedon
huxleyi, Malerisaurus, Massospondylus, Alwalkeria maleriensis.
Middle Triassic Bhimaram --------
Yerrapalli F: Ceratodus, Saurichthye; Amp: Stenocephalosaurus; Rep:
Wadiasaurus, Bethuisaurus
Early Triassic Kamthi ---------

Late Permian Kundaram Captorhinid, Gorgonopsid; dicynodonts: Endothiodon,

Emydops, Cistecephalus, Pristerodormatkayi, Kingoria,
Oudenodon
Early Permian Barakar ----------
Talchir ----------
NB: Periods and their subdivisions are represented only qualitatively; F stands for fishes; Amp for amphibians, Rep for
reptiles and Mm for mammals. Only a few important species names are given; otherwise generic or even identities
are preferred.

in fault bound basins, rather half grabens, elasmobranches, 10 families of temnospondyl

developed inland on the Indian craton along
previous weakness zones reactivated during
initiation of break-up of Pangaea. These sediments
are interpreted largely as of fluvial in origin
(in meandering or braided system of rivers;
opinions varying with authors), along with some
lacustrine and other types locally or in some parts
of the succession.
Recent studies on Gondwana vertebrates throw
much new light on the topic (Jain et al., 1975,
Bandyopadhyay 1999). Gondwana vertebrates are
known from several localities of Permian, Triassic
and Jurassic age showing significant close
relationships with those found in other parts of the
world, both from erstwhile Laurasia and
Gondwana, that is, the whole of the Pangaean
supercontinent. They, thus, show that India was
an integral part of that supercontinent from
Permian to the end of middle Jurassic.
Nine vertebrate faunal horizons are recorded
in the Indian Gondwana. There are a few other
isolated and less prominent occurrences. These
vertebrates belong to 7 families of fishes, viz.
chondrosteans, neopterygians, sarcopterygians and
labyrinthodonts, 26 families of anapsid, diapsid
and synapsid reptiles and 4 mammalian families,
2 therians and the other 2 nontherians. Major
formations that enclose these fossils are Mamal
Formation in Kashmir and Kundaram Formation
in the PG Basin (both late Permian), Panchet
Formation (Damodar basin; Lr. Triassic), Denwa
Formation in Satpura basin and Yerrapalli
Formation in PG basin (middle Triassic), Tiki
Formation in Son-Mahanadi basin and Maleri
Formation and Dharmaram Formation in PG basin
(both late Triassic) and Kota Formation in PG basin
(early to middle Jurassic).
A few interesting points about these faunas
may be discussed below. Early Permian fauna is
absent in the Indian Gondwana, though it
flourished in Laurasia. This indicates the
Gondwana climate might have been too in-
hospitable for the early terrestrial vertebrates,
though it favoured a luxuriant flora. This in turn
might have helped in uninhibited growth of the
flora and from them extensive coal deposits,
since foraging by herbivorous vertebrates was
minimal.
 Chapter 18 Vertebrata of Chordata 309

FACTSHEET 18.6
Distribution of Indian Dinosaurs

Indian dinosaur fossils are represented by

1. Skeletal remains; 2. Eggs and nests; 3. Coprolites: 4. Footprints.
Data are not available for interpretation of dinosaur behaviour based on the study of their nests, eggs, palaeoecology
of nesting sites and coprolites that may contain plant or fish remains reflecting diet of the animals.
Biostratigraphic study of dinosaur fossils may have a bearing on the resolution of the KTB problem and dino-
saur extinction event in India.
Skeletal remains are of two types : Pre-Late cretaceous and Late Cretaceous.
The former occurs in Gondwana localities particularly in the Pranhita-Godavari (PG) Valley in the neighbouring
districts of Andhra Pradesh and Maharashtra and the latter in Infratrappeans like the Lameta Beds, the Deccan
Intertrappeans and parts of Caveri basin formations.

Taxa Horizon Locality

Barapasaurus tagorei Kota Fmn. (Early Jurassic) Yamanpalli, Kota

Kotasaurus yamanapalliensis Kota Fmn. (Early Jurassic) Yamanpalli, Kota
Walkeria maleriensis Maleri Fmn (Late Triassic) Maleria
Titanosaurus indicus Lameta Formation Jabalpur
T. blanfordi Lameta Formation Pisdura
T. colberti Lameta Formation Dongargaon
Laplatasaurus madagascarensis Lameta Formation Jabalpur
Antarctosaurus Lameta Formation Jabalpur
Indosaurus Lameta Formation Jabalpur
Indosuchus (2 sps.) Lameta Formation Jabalpur
Lametasaurus indicus Lameta Formation Jabalpur
Jabalpuria teunis Lameta Formation Jabalpur
Camposuchus Lameta Formation Jabalpur
Laevisuchus Lameta Formation Jabalpur
Ornithomimoides (2 sps.) Lameta Formation Jabalpur
Megalosaurus Intertrappeans Nagpur
Massospondylus Ariyalur Fmn. Ariyalur
Bruhathayosaurus Kallamedu Fmn. Ariyalur
Brachypodosaurus Lameta Formation Jabalpur
Dravidosaurus Trichinopoly Fmn Trichinopoly
a
Also found two saurischians (a large plateosaurid and small thecodontosaurid) reported from Late Triassic Dharmaram
Formation near Maleri.

Some amphibian elements (archaegosauroids) Migration along the Tethyan coastline might have
of the late Permian Kashmir fauna, a typical led them to reach Gondwanaland. But thereafter
lowland, aquatic fauna, might have originated from the Indian, as well as other Gondwanaland fauna
early Permian European ancestors of Laurasia. of late Permian assumed provincial characters.
310 Part Three: Miscellaneous

FACTSHEET 18.7
Infra-and Intertrappean Vertebrates from Central and Western India

Localities
Asifabad A Marepalli M Naskal Ns Pisdura P Mumbai B
Dongargaon D Nagpur Ng Jabalpur J Timsanpalli T

Fishes
1. Restricted to Maestrichtian (uppermost Cretaceous) :
Igdabatis sigmodon AMNsP_NgJ_ Lepidosteus indicus AMNsPDNgJT
Pycnodus lametae AM__PDNgJ_ Belonostomus cinctus A_Ns___NgJ_
Apateodus striatus AMNs__ NgJT Stephanodus libycus AMNsP_ NgJT
2. Restricted to Campanian-Maestrichtian
Rhombodus sp. A
3. Restricted to Middle and Late Cretaceous
Lepidotes A______Ng__
4. Range not known
Raja sp. A Pycnodus bicresta A___P__NgJT
Lepidotes sp. A______Ng__ Phareodus sp. AMNs__NgJT
Enchodus sp. A___P__Ng__ Eotrigonodon wardhaensis A__P___T
Indotrigonodon ovatus AM__P____ T Palaeolabrus A______Ng__
Dormaalensis

Amphibia
Pelobatid frogs AMNsP__NgJT
Rana pusilla B

Reptiles
Booid snakes A__NsP_NgJ_ Anguid lizards A_Ns___Ng__
Pelomedusid turtles AMNsP_Ng__ Crocodilia AMNsP_Ng__T
Chelonid turtle
Hydraspis leithi B
See Factsheet 18.6 for list of dinosaurs

Mammals
Symmetrodont Ng Deccanolestes hislopi Ns

The early Triassic Panchet fauna is an
important assemblage with Lystrosaurus,
Indobrachyops and others, that closely resembles
the fauna of the Lystrosaurus Assemblage zone of
South Africa. In fact, it was one of the most widely
distributed terrestrial faunas of the Pangaea. This
fauna was also a lowland fluviatile and lacustrine
type, that flourished following the Permian
extinction event.
The Kota fauna consists of two horizons. The
lower horizon occurring in a fluviatile sandstone-
mudstone association yields sauropod dinosaurs
(Barapasaurus and Kotasaurus) and five mammals
(micromammals). The different genera and species
show resemblance to those from other continents,
viz. Europe, Asia, Africa and America. This adds
to their palaeogeographic significance. The
dinosaurs also throw important light on the

evolution of sauropods. The upper horizon comes
from limestone dominated lacustrine sediments. It
records fishes (Lepidotes, Paradapedium and
Tetragonolepis, a coelacanth, etc.) and reptiles
including a crocodylomorph, a flying reptile
pterosaur and a kind of turtle. Of these particularly
the fishes resemble those from Liassic of Germany.
18.14.3 Indian dinosaurs
Dinosaurs have been known since long in India
(see Jain et al., 1975; Bandyopadhyay 1999; Sahni
2001). William Sleeman had described some bones
in 1928, later to be assigned to dinosaurs. The
animals roamed on this piece of earth between
225 and 65 million years ago, from Triassic through
Jurassic to Cretaceous periods. Thus, earlier
records of dinosaurs come from different
Gondwana formations, whereas the Late
Cretaceous dinosaurs occur in what are known as
Lameta Formations or their equivalents in Deccan
Intertrappean or Caveri basin formations
(see Factsheet 18.6). The earliest species was
Alwalkeria maleriensis (now referred as Walkeria
maleriensis), a theropod dinosaur from Maleri
Formation. Another group, Sauropoda, was better
represented, with huge plant-eating Barapasaurus
tagorei (which had a thigh bone of more than 1.7
metres length; Figure 18.8) or slightly smaller
Kotasaurus yamanpalliensis (4.5 metres high; 14.5
metres long), in particular occurring as typical
examples (both form Kota Formation of PG valley).
In Cretaceous there were ferocious meat-eating
Indosaurus (may be about 4 metres high and 10-
12 metres long), and Indosuchus, both younger and
advanced theropods. There were as well giant
passive sauropod plant-eaters like Titanosaurus
and others. All these genera and species had
reptilian pelvic girdle or hip bone and they were
the more common or better recorded types in India.
Ornithischian dinosaurs with bird-like or avian
pelvic girdle were rare or ill represented on this
soil. Among them, there are a few uncertain
evidences of Stegosaurus, the genus familiar from
Chapter 18 Vertebrata of Chordata 311
its triangular bony plates on the back, or of giant
Diplodocus-like forms. Dinosaur egg fossils and
nesting sites at and around Jabalpur, north of
Nagpur, west of Indore, etc. of Gujarat, Madhya
Pradesh and eastern parts of Maharashtra are some
of the other interesting evidences of Indian
dinosaurs (Mohabey 2001: Figure 4). It has not
been possible to determine which kind of dinosaurs
laid these eggs, except for suggesting that some of
these eggs (called Megaloolithid, of more rounded
shapes) might be of titanosaurid sauropods while
some others (called Elongatoolithid; of more
elongate form) might be of abelisaurid theropods.
(Also see section 18.10.4 for some more details of
studies on dinosaur eggs). The former, i.e.
megaloolithid eggs occur in single layers of
sandstones without any definite arrangement
pattern, suggesting communal nesting and
gregarious habit of these late Cretaceous
titanosaurid sauropods and their preference to nest
in sands along riversides. The latter, i.e.
elongatoolithid eggs of abelisaurid theropods are
rarer and do not convey much information.
But, occurrence of theropod nest sites close to that
of sauropods may suggest carnivorous theropods
closely following herbivorous sauropods in order
to prey on them. In all the cases dinosaur fossils
were found in lake or river sediments. Particularly
in later parts of the history of Indian dinosaurs,
such sediments developed during intermissions in
the great volcanic activity which produced the
famous Deccan Trap Basalts in Peninsular India
as also similar kinds of volcanic rocks elsewhere
in the world. Godavari and adjacent valleys of
present-day south-central India were the main
abodes of Indian dinosaurs in this land, while
Narmada valley of west-central India was the
additional dwelling place in later parts. This record
of Indian dinosaurs includes important genera and
species that throw significant light on the
evolutionary history of the group, on breeding,
feeding and living habits of the concerned
organisms.
312 Part Three: Miscellaneous

Fig. 18.8 Dinosaurs: Disaggregated bone-fossils of a sauropod dinosaur mounted into a skeleton:
Barapasaurus tagorei; Jurassic; Kota Formation of PG Valley (courtesy ISI, Kolkata).

18.14.4 Vertebrates from Lameta Beds
and Deccan Intertrappeans
Vertebrates from the Lameta formation of the
Narmada Valley in central India and the different
intertrappean beds within the Deccan Traps of
central and western India include fishes, amphi-
bians, reptiles and rare symmetrodont mammals.
They are enlisted in Factsheets 18.6 and 18.7.
18.14.5 Important mammalian fauna of
India
The most important of Mesozoic mammalian
fossils of India are the micromammals, namely
the symmetrodont fossils, from the Kota Formation
of the Gondwana Group of formations. In Tertiary,
a few cetacean (whales) and sirenid (sea cows) are
reported from some Eocene marine or transitional
deposits of western India.
The most important mammalian faunas come
from younger stratigraphical units known as the
Bugti Bone Beds and the Siwalik Group of
formations that occur in Pakistan and northwestern
India. The former, though significant in housing
giant mammals like Indricotheres, are however,
restricted to parts of Pakistan. There have been
some recent studies on them and their equivalents,
that demanded some revision of ideas on them
(Lindsay et al., 2005). They will be discussed in
brief in a later section. The mammalian fauna

enclosed in the Siwalik Group is, however, much
more widely distributed and known. They will be
treated in more details. In an initial part, certain
observations on the fauna will be placed on the
basis of the faunal record, classically described.
In a later part, some recent additions and alterations
will be narrated.
Sudden appearance of this rich mammalian
fauna in Oligocene of this subcontinent may have
a palaeogeographic-cum-plate tectonic link. It was
in Oliogo-Miocene times that the African plate was
welded with the Eurasian plate as a result of which
the rich African fauna, including primates, spread
out to the Eurasian lands; the Indian fauna attests
to this. As mentioned, reported pre-Oligocene
Indian mammals mostly include Lower to Middle
Eocene marine forms.
The Siwalik Group of formations forms an
extensive belt of freshwater molasse, over-
whelmingly arenaceous clastic, deposits all along
the southern foothills of the Himalaya from Potwar
region in Salt Range of Pakistan to south and east
of the Jaldhaka River in Bengal of India. It is well-
known for a rich mammalian vertebrate fauna.
In north-western parts of India, the Group,
Middle Miocene to Middle Pleistocene in age,
overlies the Oligocene to Early Miocene sediments
grouped as the Murree, the Dharamsala and the
Dagshai-Kasauli formations mostly with tectonic
contact though conformable contact is reported in
Jammu region. They are exposed between Poonch
in the west to Paonta in the east. Equivalent of
Siwalik rocks are found in north-eastern India,
where they are given different names in different
localities. They are exposed in South Assam, South
Garo Hills in Meghalaya, Schuppen Belt of
Nagaland, Neogene folded belt of Tripura and
Mizoram, and Siwalik belt (sensu lato) of
Arunachal Pradesh. However, these deposits are
not as fossiliferous as their counterparts in western
parts of the occurrence.
In Pakistan, the Group has its equivalents often
with respective type-sections. Thus, the Potwar
1
The two dates of E. planifrons are different and do not seem to be confirmed as yet.
Chapter 18 Vertebrata of Chordata 313
Plateau of the Salt Range houses the type-sections
of the different formations of the Siwalik Group.
Bugti Formation (or the Bugti Bone Beds) that
underlies the Siwaliks, in Pakistan, also presents a
mammalian fauna, that may be considered as a
precursor to the Siwalik fauna.
The Siwalik Group has been divided into three
subgroups, viz. Lower (about 1300 metre to about
2000 metre), Middle (1400 metre to 2000 metre)
and Upper (1000 metre to about 2350 metre)
Siwalik subgroups. It ranges from 18.3 to 0.22 Ma,
that is from Middle Miocene to Pleistocene.
Traditionally and yet largely retained, the group is
subdivided into six formations, which are in the
younging order as follows: Kamlial, Chinji
(included in the Lower subgroup), Nagri, Dhok
Pathan (the two belonging to Middle Siwalik),
Tatrot-Pinjor and Tawi/Boulder Conglomerates
(Upper Siwalik).
Magnetostratigraphy, independent of
lithostratigraphic and biostratigraphic limitations
such as lateral lithofacies variations and faunal
diversity and, thus, proved an useful tool for
correlation, has been used for the Siwaliks rocks,
along with the isochronous occurrences of volcanic
ash (bentonised tuffs) during 3.0 to 1.5 Ma, 9.2
Ma and 13.2 Ma, and availability of marker fauna.
Even a magnetic polarity stratigraphy has been
suggested, though independently, for both Indian
and Pakistan rocks. [In India, it is for the Lower
and Middle Siwalik rocks from Haritalyangar area
by Johnson et al., (1983)]. It provides a datum
(7-7.5 Ma) for the last appearance of Ramapithecus
and Sivapithecus in Asia.
Elephas planifrons is reported at 3.6 Ma, the
earliest occurrence in the subcontinent.1
Barry et al. (1982) proposed four biostratigraphic
zones (interval-zones) for Middle and Upper
Siwalik Subgroups and dated them with magneto-
stratigraphic data. The zones are:
1. Elephas planifrons Interval-Zone (2.9-1.5 Ma)
(see footnote 1)
314 Part Three: Miscellaneous
2. Hexaprotodon sivalensis Interval-Zone
(5.3-2.9 Ma)
3. Selenoportax lydekkeri Interval-Zone (7.4-
5.3 Ma)
4. “Hipparion s.l.” Interval-Zone (9.5-7.4 Ma)
In Indian part of the subcontinent, vertebrate
faunas of the Upper Siwalik subgroup are
studied more extensively than those of the Lower
and Middle Siwalik subgroups. Lower Siwalik
subgroup fauna includes Deinotherium,
Dissopsalis, Conohyus, Dicoryphochoerus,
Anthracotherium, Dorcabune, Progiraffa; all of
Chinji affinity.
Typical Middle Siwalik faunas are known from
Haritalyangar of Bilaspur, Himachal Pradesh
(Prasad 1970, 2001; Vasishat 1985), one of the
most significant of which is the find of a new
species of Gigantopithecus bilaspurensis.
Nurpur Middle Siwalik fauna includes
Hipparion antilopinum, Cormohipparion
theoboldi, Propotamochoerus, Merycopotamus,
Bramatherium, Hydaspitherium, etc.
In Upper Siwalik, Hipparion and Proa-
mphibos typify Tatrot Formation, whereas Equus,
Rhinoceros, Cervus, Bubalus, Bos, etc.
characterize Pinjor fauna.
However, in view of controversies and
uncertainties in systematics and stratigraphical
positions of different fossils, it appears more useful
to consider broader aspects of the fauna.
The following are a few general points:
1. All the major groups (orders) are represented.
2. Fauna distinctly different in Lower-Middle
parts together and Upper part; the former is
more primitive and the latter more advanced.
3. Different formations vary in types of fauna.
(a) Thus, Chinji is characterized by primitive
proboscideans, Nagri by primitive
primates.
(b) Tatrot-Pinjor record both advanced
proboscideans and primate though in
lesser abundance.
4. In all the groups of mammals much of the
Kamlial-Dhok Pathan fauna (Lower and
Middle) is not found in Tatrot-Pinjor.
5. The latter marks the advent of a rich fauna of
more modern affinity.
Apart from the intrinsic character and
importance of the fauna, one interesting aspect is
its relation with later mammalian fauna of this
subcontinent. India and Pakistan have a rich
mammalian fauna presently living on this land.
There is also a fairly rich mammalian fauna in the
alluvial and cave deposits of Peninsular India.
Comparison of these faunas with the Siwalik fauna,
may lead to interesting conclusions on the pattern
of extinction and migration of the Siwalik mammals.
18.14.6 Palaeoclimatic interpretations of
Siwalik vertebrates
Palaeoclimatic conditions during the deposition of
the Siwalik Group may be reconstructed largely
on the basis of its rich mammalian fauna.
The Siwalik mammalian fauna is dominated
by five orders, viz. Perissodactyla, Artiodactyla,
Carnivora, Proboscidea and Primate. Their
elements, genera and species, do not only differ in
time, helping in zonation, but even suggest
palaeoclimatic variations. Some genera and species
of smaller or subordinate groups, too, act as good
palaeoclimatic indicators.
The Siwalik fauna includes at least three
ecological types, viz. (i) river-bank or aquatic,
(ii) forest, (iii) grassland.
Proboscideans (elephant-like forms), rich in
lower, also significant in some upper parts, were
forest elements. Of these, older forms (e.g.
Deinotherium, Trilophodon or Gomphotherium
Serridentinus, etc.) show adaptation (teeth) for
succulent plants and warm, humid lowland near
water; higher genera (Tetralophodon, Stegodon,
Elephas, Archidiskodon, etc.) were fitted to a more
rugged country and harsher vegetation (indicated
by stronger teeth).

Perissodactyls (horses, rhinos and their kins)
were varied. Rhinocerotids (Aceratherium,
Gaindatherium, etc. in lower parts; Rhinoceros
in later parts) suggest warm, humid, swampy
lands, whereas equids (Hipparion in lower and
Equus in upper parts) indicate grasslands of
increased aridity. Between the two, rhinocerotids
were more common in lower parts, the latter in
upper.
The ant-bear, Orycteropus, which now inhabits
African deserts, indicates that in Dhok Pathan
times, rainfall might have been scanty and
conditions desert-like; Camelus, too, in middle or
upper parts suggests aridity. Antelopes (family
Bovidae) and deer (Cervidae) suggest drier
grasslands, whereas goats (Capra), Ovis (sheep),
oxen and cows (Bos, Bubalus, Bison, Boselaphus)
a less arid climate. Water-deer or tragulids
(Dorcabune, etc. in earlier formations, Tragulus)
were river-bank dwellers; hippos, Hippopotamus
also; weasels too, indicated the presence of water
bodies (Martes, Lutra, Mustela) whereas pigs
(Listriodon, Propotamochoerus in older
formations, Sus in younger), and dogs or wolves
(Canis) were forest dwellers. Giant apes
(Dryopithecus, Sivapithecus, Ramapithecus,
Brahmapithecus even Gigantopithecus) in Middle
Siwaliks (in Nagri Formation) and monkeys and
apes of upper formations (Semnopithecus, Papio
or baboon, Pan, or chimpanzee, Pongo, i.e.,
orang-utan, Macaca or rhesus monkey), as also
the vast number of carnivores like Amphicyon,
racoon-like forms, Hyaenelurus, Crocuta, or
hyaena, Felis and Panthera (big cats such as tiger,
lion, etc.), Indarctos, Ursus and others (bear or
their kins) required adequate forest cover. The
earlier giraffids, okapi-like Giraffokeryx,
Hydaspitherium, might have been forest forms,
the giraffes, Giraffa, coming later might have
preferred open land with scattered trees. Rodents
represented by different genera like Rhizomys, etc.
and porcupine-like Hystrix indicate varied
environment.
Chapter 18 Vertebrata of Chordata 315
A recent study (Sehgal and Nanda 2002) of
faunal assemblages from Ramnagar (Jammu and
Kashmir) and Nurpur (Himachal Pradesh) in
India corroborates the scenario given above
(Factsheet 18.8). Sedimentological studies have
indicated a meandering fluvial regime with broad
flood plains at Ramnagar and a broadly similar,
yet of higher energy environment, as evident from
multistoried sandstones at Nurpur. Both the
assemblages are dominated by ‘terrestrial’ forms,
which include forest and grasslands types;
associated are aquatic-semiaquatic forms and
arboreal primates. The Ramnagar rocks of Chinji
equivalence (Lower Siwalik: Astarcian) house
suids, tragulids, bovids and giraffids pointing to a
tropical humid environment with prominent
swampy conditions. The Nurpur assemblage
(younger in age: Middle Siwalik: Turolian)
contains hipparionine equids and larger giraffids,
in addition to suids, tragulids and bovids
suggesting a more open condition with reduction
of forest cover and development of wider
grasslands interspersed with tree canopy.
Another study (Scott, Kappelman and Kelley
1999) of a diverse mammalian fauna in association
with the hominoid Sivapithecus parvada from the
Nagri formation of Pakistan (ca. 10 million years
in age) has thrown new light on
palaeoenvironment. The fauna is dominated by
tragulids and bovids. Based on a functional
morphological study of femoral characters of
extant bovids in relation to their habitat, it is
suggested that a forested habitat with varying
degrees of cover existed during that time. A global,
and also local in that area, predominance of
grasslands occurred between 8 and 6 million years.
The evidences of the Nagri Formation suggests
change towards that from a thicker forest canopy
during the earlier environments of the Chinji
Formation. Sivapithecus here is akin to
Kenyapithecus of Africa found from Fort Ternan
in Kenya. The latter occurrence is also dominated
by bovids, though other evidences suggest a lighter
and less continuous canopy of forest cover there.
316 Part Three: Miscellaneous

FACTSHEET 18.8
Palaeocommunities in Ramnagar and Nurpur (N) assemblages
(After Sehgal and Nanda 2002)

Community Carnivores Herbivores Omnivores

Aquatic and Crocodylus (N) Trionyx (N)
Semi-aquatic Gharialis Lissemys
Terrestrial Dissopsalis (N) Aceratherium (N) Dicoryphochoerus (N)
Percrocuta Gaindatherium (N) Conohyus
Amphicyon (N) Brachypotherium Propotamochoerus
Eomellivora Anthracotherium (N) Listriodon
Deinotherium Chilotherium Sus
Dorcatherium (N) Dorcabune (N)
Giraffokeryx (N) Giraffa
Hemimeryx Progiraffa
Protragoceras (N) Gazella
Miotragoceras (N) Hydaspitherium (N)
Hipparion (N) Cormohipparion (N)
Merycopotamus (N) Bramatherium (N)
Arboreal Sivapithecus (N) Ramapithecus

Besides mammals, crocodiles, pythons, turtles,

FACTSHEET 18.9
aquatic birds, freshwater fishes and molluscs of
the Siwalik fauna represent forest- or stream- Siwalik Climate (through time)
dwelling forms.
Boulder Conglomerate cold/glacial
In summary, it may be visualized that for most
Tatrot-Pinjor wet, warm
of the Siwalik time (Mid. Miocene to Mid.
Pleistocene), there was a luxuriant forest and open Dhok Pathan dry, warm
grasslands, with plenty of streams and lakes. Nagri drier than before
Climate would have been a dominantly warm Kamlial; Chinji wet, warm
humid one.
There was variation (see Factsheet 18.9); earlier effects of late Pleistocene glaciation. Striated and
parts (Lower Siwaliks: Kamlial, Chinji and even faceted pebbles of the Tawi (Boulder) conglomerate
parts of Nagri in Middle Siwalik) were probably Formation are attestations of this.
wetter; in the middle (particularly Dhok Pathan)
climate became dry. With post-Middle Siwalik 18.14.7 Bugti Bone Beds of Pakistan
orogeny, parts of the land went drier. But in Upper Though entirely restricted to Pakistan, the famous
Siwaliks, wet condition might have returned, in Bugti Formation or the Bugti Bone Beds have an
which more modern proboscideans, artiodactyls, important standing in Indian geology and
carnivores and others thrived. Towards the end, vertebrate palaeontology. With that in view, a brief
climate went cold ultimately coinciding with cooling note is added here, based on some updated data.

Bugti Bone Beds occur in Baluchistan of
Western Pakistan. It is reputed for abundant fossils
of some primitive artiodactyls and perissodactyls
that stand as precursors of the corresponding
members of the Siwalik fauna. The fossils are also
famous for their giant forms, particularly of the
gigantic indricotheres, Baluchitherium.
Though in the last few decades of the twentieth
century, a number of scientists (including among
others Raza et al., 2002, Downing et al., 1993,
Downing and Lindsay 2005, Welcomme et al.,
2001) have worked on the Bugti Beds or their
equivalents, particularly on those of the Zinda Pir
Dome of the Sulaiman range near dera Ghazi Khan,
where there is a lot of controversy on the age and
boundaries of the beds.
New collections reported a new group of
rodents, Baluchimyinae, from Baluchistan and
Zinda Pir Dome; rich micromammals taxo-
nomically similar to Baluchimyinae; proboscideans
and rhino fossils from Zinda Pir Dome from
Siwalik-equivalent units overlying the Bugti Beds.
Associated beds also include fossil wood in some,
or marine microfossils, bivalves and gastropods in
others. There are also reports of Thalassinoides
and Skolithos burrows in Zinda Pir Dome, which
indicate sand-dominated shore and backshore
environments (Lindsay and Downs, 1998, 2000;
2005).
These palaeontological data have been
correlated with magnetostratigraphic data and
sedimentological studies. On the basis of all these,
it is now inferred that the Bugti Beds or their
equivalents may represent a relatively stable
shoreline deposition along the vestiges of the
receding Tethyan seaway during much of the
Oligocene and early Miocene. This was followed
upwards by a fluvial system in the Siwalik-
equivalent rocks of the Zinda Pir Dome.
Additional palaeocurrent analyses indicate a
mean drainage to the southeast, agreeing with the
generally inferred palaeodrainage trend from the
early Eocene through the Miocene in the
Himalayan Foreland Basin.
Chapter 18 Vertebrata of Chordata 317
In the faunal list of smaller mammals rodents
dominate, with both primitive and more derived
forms. They are associated with fossils of bat,
shrew, hedgehog, different kinds of squirrel and a
primate Bugtilemur mathesoni.
Larger mammal fossils include large
indricothere rhinocerotids and amynodontids in the
lower part. They are succeeded by proboscideans.
Large anthracothere artiodactyls are also
characteristic of the Bugti fauna. They inlcude
Parabrachyodus hyopotamoides. Primitive
ruminants including the oldest bovids, tragulids are
found in the upper parts. Carnivorous mammals
are rare and are represented mostly by larger taxa
like amphicyonids and creodonts.
The baluchimyine rodents and indricothere
rhinos belong to the Oligocene part of the beds, dated
between 28.5 Ma to as late as 25 Ma. The appearance
of proboscideans is dated as earliest Miocene
(23 Ma). Hence, the upper parts may be equivalent
to the lowest parts of the Siwalik (Kamlial) rocks.
18.14.8 Extinction of Siwalik mammals
The rich Siwalik fauna, particularly its mammals,
were largely extinct by the end of Pleistocene.
Several causes, alternative or in combination, are
affixed. But they remain still unfounded and
unsubstantiated. Hence, a reassessment of the
question seems necessary.
The first point that makes the Siwalik
vertebrate (rather mammalian) fauna attractive to
palaeontologists, is its richness in number of genera
(or even family) and consequent diversity of
morphological-taxonomic and ecological varieties.
Almost all the major groups (orders) are important
in this regard. It is often concluded, and rightly so
that a very favourable environment was available
at that time in this part of the subcontinent. It
included luxurious forests, developed in the
foothills of the rising Himalaya, abundant water
bodies from streams coming down from the new
mountains, with associated lakes, ponds, swamps,
etc. as also occasional grasslands, particularly in
drier times. This environment could sustain a rich
318 Part Three: Miscellaneous
herbivorous fauna, which provided the trophic
level for abundant secondary and tertiary
consumers, the smaller and larger carnivores. In
addition, immigration and emigration via different
routes to and from the continents of Eurasia, Africa,
and North America were held as causing addition
to or subtraction from the fauna of this
subcontinent.
At the same time, the communities,
indigenous or immigrant, could evolve rapidly in
this congenial ambience, so much so that it could
make the region a centre of adaptive radiation and
produce the variety and diversity, observed in the
fossil record. However, it is also argued that this
evolution produced overspecialization that coupled
with drastic environmental changes could have
triggered the extinction of the Siwalik mammals.
The Pleistocene glaciation causing climatic
changes, particularly those in temperature and the
Himalayan tectonics and consequent unrest, are
held as major causes of environmental
deterioration.
To all intents and purposes, the whole gamut
of suggestions has truth inherent. But their
uncritical appreciation may not be helpful for
finding the real picture. Tectonics, as such cannot
cause unrest to destroy and drive away the living
community of the area concerned. At best, it may
cause changes in geomorphology, the latter
themselves acting as causes of change in wind and
precipitation pattern, and thus in humidity and
aridity. In short, a climatic change may be ushered
in by drastic tectonic activities. This is definitely
expected with the rise of the Himalaya, which took
place in phases though. However, the main phase
of uprising is held to have taken place by the Mio-
Pliocene and, hence, it rather precedes to the
Siwalik mammalian fauna than succeeding it.
Secondly, it is true that there are evidences of
‘glaciation’ cited with the Boulder or ‘Tawi’
Conglomerates. But as such evidences of Pleistocene
glaciation from the terrestrial, even marine rocords
are not fully supported in recent studies. So, severe
climatic deterioration may be a localized
phenomenon not enough to cause the extinction of
a fauna living for over 15-20 million years.
Ideas on palaeogeography have also changed.
With that have changed opinions about migratory
routes. At one stage of knowledge, Hipparion
was considered as evidence of a link between
Indo-Pak subcontinent and the Pikermi Island of
Greece. Subsequent details of equid evolution has
preferred immigration of Hipparion and Equus
(at different times though) from North America
to the Indo-Pak subcontinent, via the newly
formed land bridge between Eurasia and North
America, and southeastern portions of China
(Yunnan Province). Recognition of the
Afrotherian clade on molecular evidence suggests
proboscidean evolution in Africa, followed by
radiation to Asian countries (which were by that
time joined to the African continent). Hence, the
rich proboscidean fauna of the Siwaliks require a
reassessment, if it is indigenous or immigrant and
if the Siwalik forests served as the centre of
radiation of proboscideans or as a favoured
receptacle of roaming populations. Of course,
emigration from the African continent was also
correlated with tectonic activities along the Great
African Rift Valley, which also included strong
volcanic activities along the belt.
Over and above these considerations, the
Siwalik fauna may need be judged in relation to
the fairly rich mammalian fauna that exists in the
subcontinent at present. There is also an interesting
fauna in between the two, namely, the Siwalik
fauna and the recent fauna; that is, the mammalian
fauna of the alluvial and cave deposits of Middle
Pleistocene to Subrecent age that occur in the
Narmada, Krishna and other river valleys and
Kurnnol cave (in Andhra Pradesh) of Peninsular
India. Even a broad survey may pose leading clues
for future consideration on what might have been
the pattern of Siwalik extinction. Factsheet 18.10
presents some data on this.
In Siwalik stratigraphy (and palaeontology)
fixing Neogene-Quaternary boundary (i.e.
between Pliocene and Pleistocene) is a problem.
 Chapter 18 Vertebrata of Chordata 319

Without going into details of that, it may be said
that the fauna includes two parts: one more
primitive and largely of Middle Miocene-Pliocene
age and the other of more modern affinity, broadly
of Lower to Middle Pleistocene age. The Lower
and Middle Siwalik Subgroups, that is the
formations from Kamlial to Dhok Pathan, enclose
the older fauna and the Upper Siwaliks the
younger one. The categories under which the
number of genera of different orders are classified
in Factsheet 18.10 point to the relative importance
of the older and newer fauna, how much of the
older fauna was absent from the newer fauna, i.e.
apparently exterminated by the end of the Dhok
Pathan Formation and how much was the new
faunal turnover in Tatrot-Pinjor. Similarly, for the
fauna of the alluvial and cave deposits together
and for the recent fauna, categories were classifed
to show much of the older forms continuing and
how significant (in generic number) was the new
appearances.
The raw data of the study may need some
refurbishing; systematics and stratigraphic
information may have undergone some changes.
But they do not alter the broad scenario at the
generic level. The first point to note is that a large
portion of each of the orders present in the lower
fauna was absent in the upper fauna. There may
be two reasons, both partially valid. One is
extinction and the other is emigration. There was
no drastic change of environment, due to
glaciation or tectonics, at the boundary between
the two faunas. Hence, a catastrophic mass
extinction of these forms may not be the answer.
As discussed in connection with the dinosaur
extinction, it was a part of general evolutionary
pattern. In the alluvial-cave deposits of Peninsular
India, the hang-overs of the Siwalik fauna

FACTSHEET 18.10
Siwalik, Alluvial-Cave and Recent Mammalian Faunas of India Contrasted
(On Number of Genera Per Orders)
(After Ray and Baksi 1978, Ray and Roy Moulik 1979)

Mammal groups Siwalik Alluvial-Cave deposits Recent

S1 S2 S3 S4 S5 A1 A2 A3 A4 A5 R1 R2 R3 R4
Carnivora 35 26 18 09 44 00 03 00 05 08 10 04 13 27
Artiodactyla 56 33 48 25 81 00 06 02 10 18 09 02 11 22
Proboscidea 12 08 08 04 16 00 01 01 01 03 01 01 00 00
Primate 12 12 04 04 16 00 00 00 04 04 02 00 05 07
Perissodactyla 09 08 05 04 13 00 00 00 02 02 03 00 00 03
Rodentia 07 05 04 02 09 00 02 00 02 04 03 00 35 38
Tubulidentata 01 01 00 00 01 — — — — — — — — —
Lagomorpha 00 00 01 01 01 00 00 00 01 01 01 00 02 03

Number index:
S1–Kamlial-Dhok Pathan forms
S2–Kamlial-Dhok Pathan forms not in T/P
S3–Total of Tatrot-Pinjor
S4–New forms in Tatrot-Pinjor
S5–Total Siwalik (1+4)
A1–New forms, now extinct R1–Siwalik forms
A2–New forms, still living R2–Alluvial deposit forms
A3–Siwalik forms, now extinct R3–New forms
A4–Siwalik forms, still living R4–Total recent
A5–Total in alluvial deposits
320 Part Three: Miscellaneous
(e.g. Stegodon, Hippopotamus) dominate. Only
artiodactyls and carnivores present sizable
fraction of new forms. In the recent fauna, too,
the last mentioned two groups include both
considerable amount of Siwalik forms and newer
ones; the rodents present dominant newer forms
(it may be a reason that rodents were not studied
with due attention for Siwalik or other deposits).
Particularly dwindling were the proboscideans
and the primates. The former, as a group has
dwindled all over the globe (see Chapter 5); of
the latter some of the Siwalik primates are now
found in Africa (e.g. Chimpanzee or Pan); some
like orang-utan represented by Sivapithecus or
Ramapithecus or Papio are found in the Far East
forests of Malayasia, etc. Certainly whether they
migrated out of this land to their present abodes
or simply were exterminated here, seems yet
undecided.
In the end, it may be added that the rich
mammalian fauna of the Siwaliks, did record
extinctions, but that took place, might be in phases,
and apparently not causally linked with the
catastrophic environmental changes of the area
(i.e. Himalayan rise) or of the time (i.e. glaciation).
At least, one may not be prompted to look for any
massive drastic extinction of this huge fauna. It
made its way in natural manner, either losing
evolutionary battle in due course or shifting from
adversities to find a new recluse.
 19
19.1 Introduction
Plants in common usage include algae, fungi and
land plants without vascular tissues, i.e. bryophytes
such as liverworts, mosses, etc. and those with
vascular tissues. However, the classification of
plants is beset with differences among authors
(see Factsheet 19.4). To do justice to modern ideas,
yet not create confusion, it is desirable to work
with one standard recent scheme, pointing out
important variations as it may be necessary. In
recent usage, Planta refers to a taxonomic category
at the rank of Kingdom in the organic
classification; it is one of the five such kingdoms
of the organic world (Whittaker 1969, Whittaker;
see Factsheets 1.7 and 19.4). By that definition,
plants are autotrophic, eukaryotic, multicellular
organisms, popularly including land plants,
referred above. They are the organisms which carry
out metabolism by assimilating nutrients from
around and by synthesizing them with the help of
the sunlight energy: a process known as
photosynthesis.
The process is, however, not unique to the
members of Planta. Many protoctistan groups
(eukaryotic, yet unicellular) and cyanophytic monera
(prokaryotic, unicellular) are autotrophic and
photosynthetic. Hence, it is the three-character
321
Planta
combination: autotrophic, eukaryotic with nucleated
cells and multicelled body, which is critical for the
identity of plants (Factsheet 19.1). By this measure,
though heterotrophic (parasites: acquire nutrients
from a living organism/saprophyte: acquire nutrients
from dead remains of decayed products, e.g. fungi
and bacteria), neither they, nor the stromatolite-
producing cyanophytes or dinoflagellate
Zooxanthellae, symbiotic with corals, can be
regarded as plants; cyanophytes belong to the
entirely different kingdom Monera and
Zooxanthellae to Protoctista. Similarly, sapropelic,
heterotrophic fungi are also separated into the fifth
Kingdom, separate from both autotrophic Planta and
heterotrophic Animalia.
Plants have various significant uses in
palaeontology-geology. Classically they found use
in biostratigraphy. For instance, in India, the
Gondwanas were classified largely on Glossopteris
and Ptilophyllum floras or their respective generic
and species constituents at lower level stratigraphic
units. Weichselia and Onychiopsis, the two plant
fossils of Himmatnagar Sandstone Bed, an
equivalent unit of the Uppermost Gondwanas, were
used as Wealden (Lower Cretaceous) marker to help
fix the upper age limit of the Gondwanas. Later,
palynofossils were utilized for finer resolution
biostratigraphy.
322 Part Three: Miscellaneous

FACTSHEET 19.1
Plants and their Record

What plants are?

Plants are autotrophic (photosynthetic), eukaryotic (with nucleated cells) and multicelled organisms belonging
to the Kingdom Planta of the organic world. They include thallophytes, bryophytes and tracheophytes (vascular
land plants), and multicelled algae.
Ecological distinction
Being autotrophic, plants are the primary producers, thus occupying the apex of the food web or trophic chain of
organic community.
Unity of opposites in plant fossil record
Plants originated in water, where there might have been sedimentation and, hence, fossilization.
Yet they did not have preservable hard parts to ensure preservation.
Higher plants are largely land-dwelling and vascular.
They have preservable hard constituents like cellulose and lignin.
Yet they lived and live on land where sedimentation and, hence, chances of fossilization were, or are, scarce.
Preservation is enhanced when, as in Rhynie Cherts of Devonian, diagenesis introduces resistant material
(in the case of Rhynie Cherts hot spring silica) to render plant remains particularly preservable.
Higher land plants have large bodies with fairly distinct parts, like, roots, stems, leaves, seeds and flowers, etc.
that are likely to be taxonomically significant, that is, helpful for identification and systematics.
Yet plant bodies are very seldom preserved in entirety; the parts are disaggregated and may be removed from the
place the plant lived. Thus, the trunk may stand on the ground with roots driven underneath. But the leaves fall
out from above on the ground and may be carried by storms or streams. Generally, plant parts are preserved
separate and are, thus, recognized as form-genera. Glossopteris is the leaf of a plant organism and not the
organism itself. It is, thus, not a genus like Arca (a bivalve), it is a leaf-genus.

In addition, the two floras mentioned,
particularly the Glossopteris flora proved to be one
of the most important tools for palaeogeographic
reconstruction of the Gondwanaland as also for
arriving at palaeoclimatic conclusions. These have
been discussed in Chapter 20 dealing with fossils’
use in palaeogeography and palaeoclimatology.
19.2 Appreciation of Plant Fossil
Record
However, the present-day appreciation of the utility
or importance of plants in palaeontology-geology
includes a few more understanding.
Being autotrophs and so the primary producers
of food, plants lie at the apex of the trophic chain
of any community. Hence, they are vital and
inalienable component of the latter. It has two
apparent bearings. First, it is seen in the geological
record that major innovations in animal life are
preceded by some or other kind of major
innovations in plant life too. Thus, life on land
began with the advent of land plants towards the
end of Silurian; the advent of land animals followed
suit. It took place in Devonian when first tetrapods
appeared to live on land. Likewise, large
herbivorous sauropod reptiles found their abode
filled with large pteridospermous and
gymnospermous plants which had appeared in
Carboniferous-Permian times. Angiosperms
brought in their wake, so to say, birds and
mammals. It means the plants, particularly the
higher vascular plants played a vital role in
structuring and restructuring the then organic
communities, making room for ushering in of

newer and newer groups of animals. Details of this
relationship remains out of scope of the present
chapter, but nonetheless, that does not undermine
the unique importance of the plants.
However, this relationship between the floral
and faunal components of the community is often
not very apparent from the fossil record. For
example, the prolific Siwalik mammalian fauna of
India includes a large number of herbivorous
members. There were both forest dwellers and
grassland inhabitants. This implies then the
presence of a rich vegetation in that part of the
country, including both forests and grasslands.
Unfortunately, this likely-to-be rich vegetation of
the Siwalik times is not represented in the fossil
record, so far known.
Here comes the second important under-
standing on plants. The reason of this apparent lack
of knowledge of fossil record of Siwalik flora, may
simply be the lack of interest on them. Gondwana
floras found profound attention on account of the
huge coal reserve associated with them, whereas
the varied and large Siwalik animals proved to be
the main attraction, studies on any associated flora
being sidetracked. However, at the same time, it
may also be the taphonomy of plant fossils that may
have come in the way of preserving them in the
record. As discussed in Chapter 2 and
Factsheet 2.4, particularly land plants consist of
roots, stem and branches with leaves, flowers,
seeds, fruits, cones, spore-sacs, etc. After death,
even during lifetime, any of these may be
disaggregated from the main parental body, to be
fossilized, if at all, as separate entity. It is almost
invariably difficult to find out which leaf fits with
which flower or stem, and so on, resulting in
uncertainties about deciding the actual plant
organism to which they may have belonged. As
discussed in Chapter 2, in the rich Gondwana flora
of India, there is only one plant genus recognized,
Williamsonia, from one occurrence of the leaf
Ptilophyllum, stem Bucklandia and the reproductive
organ Williamsonia, attached together. In all the
other cases, we have either the leaf genus, for
Chapter 19 Planta 323
example, Glossopteris, or the stem (rhizome) genus
Vertebraria, which are referred as form-genera.
Besides, the higher plants being basically
land plants, they live on areas where there is no
deposition or little of it. As a result, the remains
of plants, either as a whole or disaggregated, stand
little chance of preservation, until and unless they
are carried to some basins of deposition.
Secondly, the remains, thus, preserved may be of
plants of the areas around or may have been
brought from distance. Once-debated issue
whether Gondwana plant remains of India were
indigenous or derived, hinged on this fact. It was
not merely academic too. The indigenous or
transported nature of the plant remains was
important in reconstructing the environment,
including knowing about the nature of water
bodies, whether there were rivers or lakes in the
area, where did fossilization take place, etc.
Sedimentological studies proving the character
of the host sediments, fluviatile, lacustrine or
otherwise, may cast light. Nature of preservation,
fragmented or complete, evidence of in situ
preservation, like root system preserved at right
angles to bedding, etc. may come as additonal
evidences to work upon.
Another important aspect of plant taphonomy
is the presence of preservable resistant material in
plant bodies. They are rare, and less durable than
the mineralized hard parts of animals. Hence, the
preservation of plants takes place differently and
is generally inferior to preservation of animal
fossils (see Factsheet 2.4). Basically plant remains
are preserved either as impressions on the
enclosing sediments without any original organic
materials preserved, or as compressions in which
the plant remain is represented by its negative
imprint, or petrifaction or permineralization in
which material from outside premeates the cells,
changes their composition or fills up the interstitial
void space (Factsheet 2.4), in cases where remains
are rapidly buried before the cell structures are
prevented from decay.
324 Part Three: Miscellaneous
Plants are found from Precambrian onwards;
but their record becomes substantial only with the
advent of vascular plants at the end of Silurian.
Components like cellulose and lignin, are among
the common resistant materials in these plant
bodies, that make them more preservable than the
earlier water-dwelling plants. However, vascular
plants were the innovation towards living on land.
This, in turn, told upon their chances of
preservation, as stated. In sum and substance, this
vital constituent of the organic world, the primary
producers of the organic community are deprived
from the due importance in the geological record
of fossils on account of taphonomic pecularities
(see Factsheet 19.1). There are, however, a few
taphonomic windows or fossil Lagerstätten that
provide interesting and fascinating data about
ancient plant life. One of them is the Rhynie Cherts
of Devonian age obtained from Aberdeenshire of
Scotland, that has been mentioned in some relevant
details in Chapter 2, as also in Factsheet Apx.1.4
of Appendix 1. Here, in one of the earliest land
plant record, preservation was enhanced with
diagenesis introducing resistant material (in case
of Rhynie Cherts hot spring silica) to render plant
remains particularly preservable.
19.3 Scope of the Chapter
At this point it must be made clear that there cannot
be any fruitful and all-pervading discussion even
on the different important aspects of plants, in the
span of one single chapter. Only a complete book
on the branch of palaeobotany, can do some justice
to the subject. Hence, the present chapter will cover
only a few most signifcant points in the plant record
that may throw some light on the geological
importance of the group. Topics or issues like plant
morphology or details of plant evolution, including
Indian examples will be touched upon, only in
course of that discussion or as background
materials presented in factsheets. Thus, Factsheets
19.2 to 19.5 provide essential background materials
for the necessary morphological terms associated
with the understanding of plant body and
description of plant fossils, brief classification and
particularities of the plant life cycle. They should
be treated as constant companion for any further
discussion. Textbooks on palaeobotany such as
Arnold (1947), Taylor (1981), Meyen (1987) may
provide more of relevant materials.
19.4 Indian Record of Land Plants
A fossil record of a region need not be judged merely
as a load of names. Rather it is best appreciated when
it is judged with a view to understanding different
aspects of the geology of the region and the
sediments to which it belongs. The following
discussion on the Indian floral record is done with
this approach and understanding.
The Indian record of land flora is
characterized by the presence of a rich flora,
known as the Gondwana flora developed during
the time span of Permian to Lower Cretaceous. It
is actually a succession of at least two, according
to some authors three, floras. The two are the
Glossopteris flora of Permian and the Ptilo-
phyllum flora of Triassic to Lower Cretaceous age,
while the third is a Triassic component, often
called Dicroidium-Thinnfeldia flora. The whole
of the Gondwana flora stands like a ‘water-divide’
separating the pre-Permian Pre-Gondwana and
post-Lower Cretaceous Post-Gondwana floras,
which all differ from each other in composition,
distribution, abundance and importance.
However, judged in the background of the
evolution of land vascular plants with three
distinct stages (see Factsheet 19.6) and of the
taphonomic particularity of plant fossils, this
entire record fits perfectly with the general
scenario, of course bearing the mark of
peculiarities of its development in this part of the
earth. Elaboration follows next. Factsheets 19.7
to 19.14 provide some details of information on
the record.
Advent of plants on land is not adequately
represented in India. The earliest land plant from
 Chapter 19 Planta 325

FACTSHEET 19.2
Background Information:
Parts of Vascular Land Plants or Tracheophytes
Vascular plants are characterized by a system of tubular passages that carry water from the underground
parts to the rest of the body. Special vascular or water-conducting cells include Xylem that convey water
from the root, and Phloem that carry food from the leaves. Life activities in plants include vegetative
functions performed by root, stem, leaf and reproductive functions performed by fructification (reproductive)
apparatus.
To present the morphology in a biologically significant way, it is necessary to follow the ontogeny of the plant.
Thus, a seed of a plant produces the shoot, its vegetative parts. The latter, at a certain stage of life, i.e. ontogeny,
give way to reproductive organs and processes. It results in fertilization and thence formation of fruits and seeds
for the next generation.
A seed is a detachable vehicle of reproduction, a mature hardened structure fairly large in size (relative to
body) which carries the fertilized spores and is covered by a sort of tough integument. It, thus, carries a new
individual, an embryo which remains dormant for a period and then germinates. In gymnosperms, the ovule
(megasporangium: the container of spores) is naked (phanerogams) and not covered in a fruit. In angiosperms
the ovule or ovules are enclosed (cryptogams) within a case formed by carpel/carpels; it is the fruit.
Angiosperm seeds may be dicotyledonous as in pea or gram or monocotyledonous, as in rice, wheat or
maize.
A plant (Figure 19.1) has:
(a) shoot system, negatively geotropic, the axis ascending from the ground with stem, leaf, etc. it is green or
chlorophyll-bearing; a shoot originates from a bud developed by a seed; and
(b) root system, positively geotropic, below the ground, which acquires water and nutrients from the soil
and provides it the anchorage to hold the plant erect.
Branch shoots, exact but smaller copies of the original shoot, develop from axillary or lateral buds.
Stem, generally aerial, is the principal skeletal part of the shoot, giving it the strength, as well as providing a
passage for conducting water and nutrients from the root to different parts of the plant body and, thus, serving as
the major trunk-link of the water vascular system of the plant. It has nodes from which the leaves originate.
Nodes are separated by internodes. Stems may be branching into two (dichotomous) or three (trichotomous) as
found in the earliest tracheophytes, the Psilophytes. Stems may otherwise be monopodial with a dominant axis
giving of branches at intervals, as found in lycopsids.
Pith cavity formed by dissolution of parenchymatous cells of the birth or naturally developed in aquatic plants.
Rhizomes are underground dorsiventral stems or branches growing horizontally under the surface of the soil;
divided into nodes and internodes as in stems. For example, potato : a tuber; onion or garlic : bulbous; Vertebraria:
a fossil rhizome from the Lower Gondwanas.
Leaves produce ‘food’ from the nutrients and water with the help of chlorophyll that absorbs sunlight and, thus,
acquires the energy to perform the biochemical reactions; they also help in respiration and transpiration. Leaves
originate from nodes on stems. They may be of different kinds :
1. A cotyledon of a seed
2. Scale leaves with underground stems or aerial parts (in bamboos).
3. Bract leaves on floral buds at their axils.
4. Prophylls, the first few leaves of a branch.
(Cont...)
326 Part Three: Miscellaneous

FACTSHEET 19.2 (Cont...)

Background Information:
Parts of Vascular Land Plants or Tracheophytes

5. Floral leaves and sporophylls, sepals, petals, stamens and carpels are specialized leaves; stamens and carpels
are also called sporophylls.
6. Foliage leaves, leaves in the common parlance, which perform photosynthesis, transpiration and respiration;
protect buds, serve for vegetative propagation, etc.
In many plants (xerophytes) the function of leaves is taken up by the stem, which remains green, flat and leaf-
like. On the other hand, there are plants in which there is no stem, the whole plant is a leaf (Lemna).
Phyllotaxy is the mode of arrangement of leaves on the stem. When there is one leaf at a node, the phyllotaxy
may be spiral/alternate/acyclic. When there are two or more leaves at each node, it is cyclic/verticillate/ whorled.
When there are two at right angles, phyllotaxy is opposite. For example, we have opposite phyllotaxy in
Schizoneura (Lr. gondwana), spiral in Elatocladus (Up. Gondwana).
A foliage leaf has three parts:
Hypopodium/leaf-base; Mesopodium/petiole; Epipodium/leaf-lamina or blade;
Phyllopodium is the entire leaf axis.
Base may be petiolate or sheathing or decurrent, etc.
Sheathing leaf-base is prominent, winged and clasps and sheaths the stem.
Decurrent leaf-base extends down the stem as two wings.
Petiole is the stalk that connects the leaf-lamina with the stem and is, thus, the vital link in the water vascular
system of the leaf. When there is no petiole the base or the plant is called sessile. The sessile leaf (or the leaflet,
later explained) may be attached to the stem (or the rachis) by the whole of its base or a part.
Leaf-lamina is described or identified on certain criteria. More important in fossils are shape, base, margin,
apex and venation.
Shape varies and may be acicular (pine), linear (grasses), lanceolate (karabi), oblong (banana), ovate (jauba),
cordate (betel), reniform (thankuni), obovate (kanthal/jackfruit).
Margin of the lamina may be entire (mango), serrate (china-rose/jauba), dentate (water lily), lobed (radish), etc.
Apex may be acute, acuminate, obtuse, truncate, etc.
Veins are linear, tube-like passages, which are actually parts of the vascular tissue syetem of a leaf-lamina that
traverse the latter. The system enters the leaf-lamina at the base, through the petiole and then branches out or
ramifies. These ramifications are called veins and their arrangement venation. Venation serves as a circulatory
system for water and other raw material as well as prepared food throughout the leaf and as a skeletal structure
of the leaf to give mechanical strength.
Type of venation is basically two: reticulate and parallel. The type is best determined from the smaller veins and
not the main ones. In reticulate type, smaller veins also ramify, whereas in parallel veins, the smaller veins are
unbranched and run parallel to one another.
Further types are:
Reticulate venation:
1. Unicostate/pinnate: with a midrib, which is the main vascular passage running from base to apex and
giving out secondary branch veins on either side, as in a feather, that in turn may give out smaller
anastomosing veins.
(Cont...)
 Chapter 19 Planta 327

FACTSHEET 19.2 (Cont...)

Background Information:
Parts of Vascular Land Plants or Tracheophytes

2. Multicostate/palmate: Here the vascular system at the base of the lamina breaks up into a number of
equally prominent major veins or costae, without a midrib and radiating from the base; they may again
converge towards the apex (convergent palmate) or diverge and spread out (divergent palmate).
Parallel (or striate) venation:
1. Unicostate/pinnate with a midrib giving off parallel branches that may be joined by still smaller transverse
veinlets parallel to each other.
2. Multicostate/palmate with a number of costae, convergent or divergent.
Leaves may be simple or compound. To distinguish them, the following points are relevant:
Leaf-lamina margin may be entire or broken or incised. The degree of incision varies. Much incised margin
makes the leaf lobed, incision running upto the midrib in leaves with pinnate venation or upto the base in
those with palmate venation.
Finally, the lobes are so complete that there is no laminar connection between them and each of them is independent
of one another. They are then called leaflets and the leaf itself, a compound leaf, pinnate or palmate.
Thus, in a simple leaf, howsoever dissected, the lobes have laminar connection with the adjacent ones. This
makes the leaf look like a complete entity.
In a compound leaf, the lobes are not connected and are independent. Hence, it looks like a leaf made of many
smaller parts.
A simple leaf has an axillary bud and a stipule at its termination.
In a compound leaf there is no axillary bud or stipule at the termination of leaflets. It is present at the base of the
whole leaf.
A simple leaf is attached to the stem.
In a compound leaf, the leaflets are attached to rachis, which is the stem-like feature, but is actually the main axis
of the vascular system of the leaf, usually equivalent to the midrib (in the case of pinnate compound) or the
petiole (in the case of palmate compound) of the simple leaf.
In most of the fossil samples of leaf genera, the base is not preserved. Thus, only the look as a complete form or
made of smaller units, is the only criterion to work immediately upon.
Flowers, fruits, seeds, cones and spores are concerned with reproduction.
Sporangia are the capsules in which the spores occur.
Sporophylls (sporangium-bearing leaves) are fertile, scaly leaves that cluster to form cones, for instance in
lycopsids as also the stamens and carpels in flowers of angiosperms.
A flower, a reproductive organ, may be defined as a shoot with sporophylls. It means it is a modified shoot for
the special purpose of reproduction. An ordinary shoot has an axial stem with nodes and internodes and the
foliage leaves occurring at nodes in alternate or cyclic phyllotaxy. In a flower, the thalamus is equivalent to stem;
it is so condensed that nodes are too close spaced to be rocognized. But they are there and the floral leaves
(modification of foliage leaves) are borne at these nodes in spiral or cyclic phyllotaxy.
328 Part Three: Miscellaneous

Apical bud

Axillary bud

Floral bud

Leaf

Branch stem

Node

Branch trace Node

} Internode

Leaf trace Xylem

Phloem }
Cortex
Pith

Shoot system
Stem

Epidermis

Tap root

Root system
Branch root

Root hair

Origin of
branch root Root cap

Fig. 19.1 Parts of a plant.

 Chapter 19 Planta 329

FACTSHEET 19.3
Background Information:
Leaf and its Origin

A leaf is an areally limited, flattened in a plane, generally green outgrowth of stem or its branch, developing
from the node and having an axil there at the point of emergence (see Figure 19.2).
In early tracheophytes leaves were small and scale-like.
Shoots were differentiated into a central axis with regular offshoots; they became confined to a plane; webbed
to form a leaf (Rhacopteris) in ferns.
Rachis: Main stem of fern frond ; first–order branches are primary pinnae; second-order branches are secondary
pinnae; Pinnules are the smaller subdivisions.
Morphological parts relevant for description and also for identification of fossil leaf genera.
1. Leaf base (hypopodium) is the part of attachment with stem (or branch).
It may be petiolate or sessile; a petiole (leaf-stalk) generally protects the small bud at the axil (Glossopteris;
generally not found).
Sessile base may be broadened into a flattened extension that sheaths around and covers the stem, to form
a sheathing leaf base (Elatocladus).
Decurrent leaf base (Dictyozamites).
2. Petiole (mesopodium) is a rod-like structure to support the base; it may be attached to the base of the
lamina, in the same plane with the lamina axis (as in Mango leaf/ Glossopteris) or may be attached to the
centre of the lower surface of the lamina at right angles to the lamina axis (as in ......).
3. Lamina/leaf-blade (epipodium) is the flat, expanded green part of the leaf, traversed by a network of
veins. Characteristically, it has a point of attachment (base), the end opposite to the base (apex), the margin
which defines the areal extension, the two surfaces, of which the dorsal or the upper surface is greener and
with veins embedded on it, while the ventral of the lower surface is paler with veins raised on it; a mid-vein
or midrib may run from base to apex.
A few common shapes of lamina are : acicular, linear, lanceolate, spathualate, oblong, ovate, reniform, fulcate,
elliptical, orbicular, etc. The margin may be entire, wavy, serrate, dentate, lobed, etc. The apex may be acute,
obtuse, acuminate, etc. Venations are basically of two types: reticulate or parallel; each may be pinnate or
palmate. The two are contrasted below:

Reticulate Parallel

A. Pinnate: A. Pinnate:
single midrib, many lateral branches from base Prominent midrib or not; lateral branches
to apex. parallel.
B. Palmate: B. Palmate:
>1 prominent rib, many lateral branches spread >1 prominent ribs run from the petiole towards
out like fingers from base to apex. margin and apex.
(i) convergent: converge towards apex veins (i) yet curved towards apex
parallel.
(ii) divergent: diverge towards apex and margin. (ii) veins parallel, yet curved towards margin
330 Part Three: Miscellaneous

Apex
(i) Vein (i)

Margin Lamina
or
Epipodium

Petiole
Lamina base or
Mesopodium (ii)
Axillary bud Stipules
(ii)
Leaf-base
or
hypopodium
(a)

Carpel
(Gynoecium) Stamen
(Anoroecium)

(iii) (iii)

Petal
(corolla)
Sepal
(calyx)
Thalamus

(b)

(iv) (iv)

(d)
(c)

Fig. 19.2 Parts of a leaf and a flower and simple to compound leaves.
(a) Typical parts of a leaf, (b) Typical parts of a flower, (c) and (d) Stages of development from simple
to compound leaf, (c) Simple to pinnate compound, (d) Simple to palmate compound
(i) Simple leaf; (ii) Simple leaf with indented margin; (iii) and (iv) Compound leaves .
 Chapter 19 Planta 331

FACTSHEET 19.4
Background Information:
Plant Classification
NON-VASCULAR Taylor 1981 Meyen 1987
PROKARYOTES
Algae* Bacteriophyta
Thallophyta (algae, (Seven divisions) Cyanophyta
fungi, etc.) EUKARYOTES
Fungi* Pyrrophyta Chrysophyta: cocoolithophores, etc.
(Five divisions) Phaeophyta Bacillariophyta
(* non-formal groups) Rhodophyta Chlorophyta
Charophyta
+ Fungi; Acritarcha and Cryptarcha
HIGHER PLANTS
Bryophyta Bryophyta Bryophyta
(liverworts, mosses) (non-vascular)
VASCULAR Rhyniophyta, Zosterophyllophyta Propteridophyta
Trimerophytophyta, etc. Rhyniopsida
Trimerophytales, Psilophytales, etc.
Psilopsida
Psilophytes Pteridophyta
Lycophyta Lycopodiopsida/Lycopsida
Lycopsida Sphenophyta Equisetopsida
Equisetales, etc.
Sphenopsida Pteridophyta Polypodiopsida/Pteropsida/Filicopsida
Marattiales
Polypodiales/Filicales, etc.
Progymnospermophyta Progymnospermopsida,
Pteropsida Noeggerathiales, etc.
(Arnold 1947) Pinophyta/Gymnospermae
Ginkgoopsida,
Ginkgoales
Pteridospermophyta Arberiales
(equal to Glossopteridales), etc.
Ginkgophyta Cycadopsida
Cycadophyta Cycadales
Cycadeoidophyta Bennettitales
Coniferophyta Pinopsida/Coniferopsida
Cordaitanthales
Pinales(conifers)
Anthophyta Magnoliophyta/Angiospermae
Also see Factsheets 1.7 and 15.6
Following International Code of Botanical Nomenclature, for bryophytes and vascular plants, the suffix-phyta indicates
Division, - opsida stands for Class; -ales for Order.
332 Part Three: Miscellaneous

FACTSHEET 19.5
Background Information:
Two Generations of Plant Life Cycle

Three important stages (rather revolutions) of evolution of vascular plants involved new innovations towards
life on land, more and more freeing plants from dependence on moisture around and adopting to life away from
water. It was foreseen in Devonian spore assemblages. All tracheophytes were homosporous at the outset; towards
the end of Devonian heterospory became common, seeds developed from that in Upper Devonian. Seeds existed
in cones in gymnosperms, in angiosperms they were placed in flowers and then fruits. Development of seed-
coat as a protective covering was also an innovation to protect seeds and help them function independently of
water from outside.
Vascular land plants reproduce in two ways, producing two generations and types of offsprings.
One, the sporophyte, reproduces asexually producing wind-borne spores in millions (e.g. in pteridophytes).
The other, gametophyte, reproduces sexually and results from germination of these spores. Here male sperm
reaches female organ and the latter, thus, fertilized leads to the growth of new sporophyte. For this to take place
the gametophyte must be covered by a film of moisture to allow the sperm to swim to the female organ. It makes
such plants live near water, in damp places (mosses, ferns, etc.). It also meant that even the sporophytes, more
independent of water, could not live away from moist places to keep them near the gametophytes.
Mosses and Psilophytes of tracheophytes are homosporous. Here, the spores which are released by the sporophyte
and which give rise to the gametophyte generation are the same. The gametophyte possesses both male and
female organs on the same plant.
Some lycopsids, sphenopsids and ferns are heterosporous (Sphenopsids are homosporous, but sometimes function
as heterosporous). Smaller microspore gives rise to a gametophyte with male organs, and the larger megaspore
forms a female gametophyte.
However, in all these groups, spores kept in sporangia or formed in cones or strobilus (in Lycopsida), which are
clusters of fertile, scaly leaves or sporophylls, carried out reproduction.
Seed plants broke away from dependence on water. Here female gametophyte remained on the sporophyte
(captive: it is called). It developed within ovule, a structure. It was fertilized by microspores (known as pollen)
Pollen reached female stigma, grew a long pollen tube, the equivalent of gametophyte, which runs down to the
ovule. Fertilization produced new sporophyte, which takes the form of seed, an embryo inside. Through various
dispersal mechanisms, seeds give out sporophytes to disperse. Seeds may be borne in woody cones, as in conifers,
from which they are dispersed by winds.
Seeds may lack a thick, leathery coat, as in gymnosperms (naked seed) also called phanerogams, or may have
a thick, leathery coat from the covering of the ovule, as in angiosperms (hidden seed : cryptogams).
A flower, which is virtually a little more than a colourful cone, houses both male and female organs, anthers
or microsporangia (male) borne on slender filaments and carpels (female organs or captive femle gametophytes).
It produces cryptogam seeds, where the seed-coat develops from the covering of the ovule.
Since flowers produce protected seeds, rapid dispersal and self-fertilization therefrom become rare. Thus, flowers
encourage and ensure outbreeding or cross-pollination whereby, in turn, there is a good amount of genetic
mixture within the population, helping in wider adaptations.

India is also the psilopsid, but represented by
Psilophyton only from Muth Quartzite of Devonian
age. There are reports of Ordovician and Silurian
ages, which are however not confirmed, nor
universally accepted.
Earliest questionable report of lycopsid from
India is Protolepidodendron sp. from Late
Devonian of Kashmir or Lepidodendropsis
(a cosmopolitan genus) from Early Carboniferous
of Spiti/Kashmir; Bothrodendron from Barren
 Chapter 19 Planta 333

FACTSHEET 19.6
Three Stages of Evolution of Vascular Plants and
the Major Groups Produced Thereupon

1. Advent on land and vascularity

2. Seeds and spread on land
3. Flowers and conquer on land

Three stages Major Plant Groups

1. Development of vascularity Palaeozoic plants without seeds

and colonization of land
In late Silurian, followed by
Devonian diversification.
The new plant groups were
devastated through end-
Permian extinction.
2. Appearance of new seed-
bearing groups
In Permian-Triassic, followed
by acme in Jurassic-Lower
Cretceous.
These groups were forced to
subordination (in abundance)
with the advent of flowering
plants.
3. Flowering plants including
grasses
In Cretaceous
Colonization of land near-
complete
NB. Within parentheses are shown the stratigraphical range of the group, followed by the period of
maximum development.
Psilopsida/Psilophytes (Upper Silurian-Recent)
Lycopsida (Upper Silurian-Recent)
Sphenopsida (Lower Devonian-Recent: Carboniferous)
The ferns:
Pre-ferns (Middle Devonian-Upper Permian)
True ferns (Filicopsida: Devonian-Recent: Mesozoic)
Seed ferns (Pteridospermales: Upper Devonian-Jurassic)
Seed-bearing plants:
Palaeozoic:
Pteridospermales: (Upper Devonian-Jurassic: Permian)
Cordaitales (Carboniferous-Triassic: Permo-Carboniferous)
Coniferales (Upper Carboniferous-Recent: Jurassic-Cretaceous)
Non-flowering seed plants of Permian/post-Permian
Ginkgoales (Permian-Recent: Permian)
Cycadales (Triassic-Recent: Mesozoic)
Bennettiales (Triassic-Upper Cretaceous: Early Mesozoic) Dominant
over cycads in Mesooic
Flowering plants
Angiospermopsida (Triassic/Cretaceous-Recent: Recent)

Measures of the Lower Gondwana is the earliest
member of the group from the Gondwanas.
Sphenopsids find more significant
representation. Two orders of this division,
Sphenophyllales and Equisetales are represented
in the Gondwanas of India, the former important
in Lower Gondwanas, whereas the latter having
important genera and species in both Lower and
Upper Gondwanas.
Filicopsida, the other major group of non-seed-
bearing vascular plants have two orders
representing it in the Gondwanas. Of them Marat-
334 Part Three: Miscellaneous

FACTSHEET 19.7
Land Plant Record of India:
Three Stages of Plant Evolution and Indian Scenario

Three stages Indian scenario

1. Development of vascularity and
colonization of land in late Silurian
Diversification in Devonian
2. Appearance of new seed-bearing
groups
In Permian-Triassic, followed by
acme in Jurassic-Lower Cretaceous.
These groups were forced to
subordination (in abundance) with
the advent of flowering plants.
3. Flowering plants including grasses
In Cretaceous
Colonization of land near-complete
Silurian appearance and Devonian diversification not represented;
earliest in Spiti in Devonian
Pre-Gondwana Permo-Carboniferous flora of Spiti and Kashmir,
Cosmopolitan lycopsid, Lepidodendropsis flora of Lower
Carboniferous of Kashmir may just predate the more known Spiti
flora of Sphenophyllales and Filicales in Thabo Stage of Po Series
of Middle Carboniferous.
Prolific pteridosperm (seed-bearing ferns/cycadofilicales) records of
the Lower Gondwana formations point to the advent of seed-bearing
land plants in India. They were accompanied and finally succeeded
by different seed-bearing plant groups of gymnospermous affinity,
particularly Cycadeoidales/Bennatiales and Coniferales in Mesozoic
parts of the Gondwanas, i.e. Upper Gondwanas.
Flowering parts or angiosperms are recorded in India since late
Cretaceous in Deccan Intertrappean flora (essentially swampy);
subcrops and disjunct Cenozoic outcrops from different parts of India,
including those of the Tertiary lignites (largely paralic), the Siwaliks
of northern India (forest or grassland, mountainous or plainland) and
Karewas of Kashmir.

tiales has Marattiopsis in Upper Gondwana
formations, whereas the other one, viz. Filicales,
the true ferns, are overwhelmingly dominant and
figure from Palaeozoic pre-Gondwana as well as
Lower Gondwana formations, but are much more
prominent in Mesozoic, after the decline of the
seed-ferns (Pteridospermales) (Factsheet 19.8).
They continue, though subdued by the angiosperms
in Cenozoic, with fern spores continuing from
Palaeozoic to Cenozoic. A special mention may be
made of Azolla, a water-fern occurring in Deccan
intertrappeans of Eocene.
In a more recent account, however, reports
are made of Taeniocrada sp., Protolepidoden-
dron sp. questionably from Late Devonian of
Kashmir and Rhacopteris, Lepidodendropsis
and Triphyllopteris from Early Carboniferous
of Spiti/Kashmir. All the three genera in the last
case are cosmopolitan. In addition, Early
Permian rocks of Kashmir are said to have
Rajaria, Sphenophyllum and Cathaysian
elements; of Salt Range to have Gangamopteris,
Glossopteris, Ottokaria , Samaropsis and
palynoassemblage; late Permian of Salt Range,
Malla Johar in UP to bear typical palyno-
assemblage; rocks of the same age from Sikkim,
Darjeeling, Arunachal Pradesh having megafossils
of Gondwana aspect. Thus, these Permian rocks
have been considered as equivalents of the
Gondwanas formed along the periphery of the
Gondwanaland proper.
Though seed-bearing plants had their acme in
middle to later parts of Mesozoic, efficiency and
supremacy of seed-bearing habit towards living on
land, was established by the appearance and
development of pteridosperms in Carboniferous
 Chapter 19 Planta 335

FACTSHEET 19.8
Land Plant Record of India:
By Group and Through Time

l Psilopsids represented by Psilophyton from Muth Quartzite (Devonian)

l Earliest lycopsid from India questionably Protolepidodendron sp. from Late Devonian of Kashmir or
Lepidodendropsis (a cosmopolitan genus) from Early Carboniferous of Spiti/Kashmir; Bothrodendron from
Barren Measures of the Lower Gondwanas
l Sphenopsids: Sphenophyllales and Equisetales are represented in Gondwanas of India; of the former
Sphenophyllum speciosum important in Barakar and Raniganj Fmns. Sphenophyllum also from Early Per-
mian of Kashmir.
Of the latter, Schizoneura gondwanensis and Phyllotheca indica are important in Karharbari, Barakar and
Raniganj Fmns.
Equisetites rajmahalensis in Rajmahal Fmn.
l Filicales: In Palaeozoic: Rhacopteris ovata and Sphenopteridium furcillatum, the earliest ferns in
India from Thabo Stage; Rhacopteris and Triphyllopteris, two cosmopolitan genera are also reported from
Early Carboniferous of Spiti/Kashmir
Alethopteris, Pecopteris, Sphenopteris common in Talchir and Raniganj
Callipteridium in Karharbari
In Mesozoic, i.e. in Upper Gondwana Pecopteris, Sphenopteris, Cladophlebis, Gleichenites, Weichselia,
Matonidium, Protocyathea.
Also present is Marattiopsis of Marattiales in Upper Gondwana formations
Fern spores from Palaeozoic to Cenozoic
Water-fern Azolla in Deccan intertrappeans of Eocene.
l Pteridospermales Cycadofilicales
Gondwanidium validum characteristic in Karharbari
Glossopteris, Gangamopteris, Vertebraria all through Lower Gondwana. Barakaria dichotoma and
Walkomiella indica characteristic in Barakar Fmn.; Palaeovittaria in Raniganj Fmn. Dicroidium in Panchet
equivalents.
Salt Range have Gangamopteris, Glossopteris, Ottokaria, Samaropsis (a cordaitale) also from Permian.
l Among gymnosperms Cycadeoidales/Bennatiales include Taeniopteris, Ptilophyllum, Bucklandia,
Williamsonia, Zamites, Otozamites, Dictyozamites, Pterophyllum all from Upper Gondwanas.
Cycadales: Macrotaeniopteris Pseudoctenis, Nidia?
Ginkgoales: Psygmophyllum haydeni in Gangamopteris Beds (Mamal Fmn.) of Kashmir, Rhipidopsis
ginkgoides in Barakar Fmn., Ginkgo lobata from Jabalpur Fmn., species of Ginkgoites from the coastal
Gondwanas of Andhra Pradesh.
Cordaitales: Dadoxylon from Lower Gondwana formations as also coastal Gondwanas of AP and Umia
Beds of Kachchh. Noeggerathiopsis hislopi, a common species from Lower Gondwanas, Cordaicarpus and
Samaropsis are seeds, common in Talchir.
Coniferales: Buriadia sewardi/Voltzia (Palaeozoic conifers) characteristic from Karharbari Moranocladus
from Karharbari, Elatocladus from Rajmahal, Jabalpur, Umia formations,
Aurocarites, Indostrobus, Takliostrobus are fruit cones from upper Gondwanas
Brachyphyllum, Pagiophyllum, Podozamites systematics uncertain, important in Upper Gondwana.

(Cont...)
336 Part Three: Miscellaneous

FACTSHEET 19.8 (Cont...)

Land Plant Record of India:
By Group and Through Time

Pollen of different groups of conifers are common in Tertiaries of Assam, etc.

Angiosperms:
There may be a long list of angiosperm fossils.
Here only a few geologically or palaeontologically significant points are included.
Sahnioxylon: earliest angiosperm in India, Rajmahal Intertrappeans
Angiosperm pollen of Eocene age come from Cauvery basin and West Bengal subcrops, Kachchh and Khasi
Hills
Tura flora of Palaeoc.- Lr. Eoc of Meghalaya, Deccan Intertrap. flora Eoc. of Deccan,
Cuddalore flora Mioc. AP, Karewa flora Pleistoc Kashmir, Siwalik flora Mioc-Plioc.
Monocot:
Palmoxylon: petrified stem : Kachchh Cretaceous and Tertiaries elsewhere.
Palm-leaf Kasauli Beds of Early Miocene in Simla
Nipa: Palm fruit in Deccan Intertrap MP
Dicot:
Fossil wood from
1. Deccan Intertraps, MP
2. Upper Tertiary Assam, Tripura
3. Cuddalore and equivalent beds of Coromandal Coast
4. Kasauli Beds and Siwaliks
Leaf fossils from the Siwaliks

and Permian. In fact, the Lower Gondwana flora
of Permian is named and characterized by members
of this group, viz. Glossopteris, Gangamopteris,
Vertebraria in particular, which were often prolific,
though varying in their relative abundance
(Factsheet 19.11). Association of these fossils with
huge deposits of coal and carbonaceous shale also
attest to the presence of fairly well-developed forest
at that time, which would have provided the
enormous amount of ‘carbon’-matter required. Of
course, pteridosperms were associated with other
gymnospermous groups like Cordaitales
(e.g. Dadoxylon, Noeggerathiopsis), Ginkgoales or
Equisetales like Schizoneura and Phyllotheca
among others to make the flora richer.
Whether typical pteridosperms of Permian,
viz. Glossopteris continued into Triassic in India
is debated though the genus is said to coexist with
Ptilophyllum in the Panchet formation in the type
area. In general, however, the Upper Gondwanas
ranging through Mesozoic, is characterized by the
gymnosperms, and particularly by the cycad-like
(Cycadeoidales/Bennatiales) and conifers
(Coniferales). Their relative importance varies,
conifers assuming more prominence in the younger
formations; nevertheless they remain the major
groups of non-flowering seed-bearing plants
flourishing in India.
Post-Gondwana floras were the latest
Mesozoic-Cenozoic in age. Though it is now
suggested that angiosperms might have appeared
as early as in Triassic, they were no doubt worth
notable only in late Cretaceous. The earliest
angiosperm in India (Sahnioxylon) is, however,
reported from Lower Cretaceous Rajmahal
Intertrappeans. Subsequent distinct Deccan
 Chapter 19 Planta 337

FACTSHEET 19.9
Background Information:
Gondwana Group/Supergroup of Formations: What It Is

l Gondwana Group/Supergroup of formations is a huge thickness of essentially continental sediments.

l It is generally agreed stratigraphic range is Permian to Lower Cretaceous, though some authors like Veevers
and Tewari (1995) mark the initiation of the Gondwana basins at Upper Carboniferous.
l It is characterized by dominantly siliciclastic rocks with significant carbonates found rarely as in Jurassic
(Kota Formation), with India’s richest coal reserves and carbonaceous shales associated with them.
l It is further characterized by typical terrestrial floras (Gondwana floras, as mentioned) and vertebrate faunas
(mainly amphibians and reptiles, occasional fishes).
l Sediments were deposited in fault bound basins, rather half grabens, developed inland on the Indian craton
along previous weakness zones presumably reactivated during initiation of break-up of Pangaea.
l Sediments are interpreted largely as of fluvial in origin (in meandering or braided system of rivers; opinions
varying with authors), along with some lacustrine and other types occurring locally or in some parts of the
succession.
l The generalized succession is, in younging order, Talchir/Karkarbari/ Barakar/ Barren Measure/ Raniganj /
Panchet-Parsora-Mangli/ Mahadeva-Yerrapalli/ Tiki-Maleri/ Kota/ Rajmahal/ Jabalpur Formations, but there
is controversy on the boundaries between units and their correlation.
l Lower parts of the succession are developed particularly in Damodar-Koel valley, Son-Mahanadi valley,
Pranhita-Godavari valley, Satpura- Narmada valley ; and the upper parts occur in widely apart basins
including the mentioned ones as also in Rajmahal basin and in a few isolated basins (with equivalents) in
coastal areas of Andhra Pradesh, Orissa, West Bengal, in eastern India and Kachchh, Jaisalmer, Caveri
basins, etc.
l Different basins show different history and with successions, differing significantly.
For instance, a boundary conformable somewhere is represented by break elsewhere.
Breaks are also represented in different magnitudes in different basins.
A thick unit of one basin is either reduced or represented by different facies in another.
A single classification of the succession is debated in result.
l Classically, a two-or three-fold classification is proposed by different authors primarily on flora as also on
other criteria. The two-fold classification is generally favoured with a lower unit of Permian and an upper of
Triassic to Lower Cretaceous age. In three-fold version, a middle unit is raised for Triassic formations with
Dicroidium-Thinnfeldia flora. Presently, however, a single succession of formations is preferred.
l Four major facies associations, viz. (1) fluviatile, the most dominant (channel + flood plain) and occasional
lacustrine producing sandstone-shale combination with or without coal or carbonaceous shale; (2) red bed
facies (Permo-Triassic) with a hemaitic cement in coarse or fine sandstone as well as siltstone ; (3) marine
intercalation or alternations found inland during Permian (e.g. Umaria, Manendragarh) and along the coast in
Lower Cretaceous (east and west coast); (4) intertrappeans particularly in Rajmahal basin which are of fluvial-
lacustrine-swampy origin.

Intertrappean flora characterized by its swampy
attributes developed in and around water bodies
on low-lying or flat topography of the
subhorizontal lava-flows. It included an important
part of the angiosperm flora. Other representations
of the latter come from different subcrops and
disjunct Cenozoic outcrops from different parts
of India, including those of the Tertiary lignites
(at Neyvelli in Tamil Nadu, Palana in Rajasthan,
Panandro in Kachchh and in Assam), the
338 Part Three: Miscellaneous

FACTSHEET 19.10
Background Information:
Problem of Fixing Age of Gondwana Dormations

Biostratigraphically important evidences that help fix the age of Gondwana Formations or its lower or upper age
limits include (i) association or intercalation with marine-fossil bearing rocks, (ii) age-diagnostic fossils,
(iii) stratigraphic position, and (iv) other evidences.
Marine intercalations occur at different stratigraphic positions in Talchir and Karharbari Formations, at different
places like Umaria, Manendragarh and Anuppur (MP), Rajhara near Daltonganj (Jharkhand), Sikkim, Arunachal
Pradesh, etc.
On the strength of fossils like Eurydesma, Pleurotomaria, Spirifer, Productus a broad Lower Permian age may
be fixed for these, setting the lower age limit of the Gondwanas at that level.
Marine associations or intercalations also occur near or at the top of the succession, particularly in what are
called coastal Gondwanas. Raghabapuram mudstone with Clupavas neocomensis (fish), ammonoids and
foraminifers of Lower Cretaceous age occur intercalated with sandstones containing Ptilophyllum. Ptilophyllum
with Palmoxylon occur in Umia Plant Beds in Kachchh associated with Trigonia beds containing T. ventricosa
and T. crassa of Lower Cretaceous age. This determines the upper age limit.
Being continental and largely provincial, most of the plants and larger vertebrates are not helpful for age
determination. However, there are a few important fossils.
Thus, the assemblage with Lystrosaurus, Indobrachyops and others, closely resemble the fauna of the Lystrosaurus
Assemblage zone of South Africa. This fauna, one of the most widely distributed terrestrial faunas of the Pangaea,
was a lowland fluviatile and lacustrine fauna that flourished following the Permian extinction event and fixes
early Triassic age for the host Panchet Formation.
Fishes like Lepidotes, Paradapedium and Tetragonolepis that occur also in the Liassic or Lower Jurassic of
Germany, in association with marine fossils, determine the same age for the Kota Formation. Large sauropods
and micromammals of the same formation also suggest Jurassic age.
Rhinesuchus from equivalents of the uppermost parts Raniganj Formation is a Permian form.
Onychiopsis from Bansa and Jabalpur Formation in MP and Umia in Kachchh Weichselia (in association with
Matonidium) from Himmatnagar Sdst., Dhrangadhra Formation of Gujarat and Bansa Formation of MP mark
Lower Cretaceous or more precisely Wealden age. Onychiopsis and Weichselia are found associated with
Ptilophyllum also in Japan and different parts of Europe (Sengupta 1988). Dicroidium, a pteridosperm is held as
Triassic marker; however, it has now been reported from Upper Permian of Dead Sea region, which was in the
then tropics (Kerp et al., 2006). The genus, it is held, might have spread southwards subsequently, to be found
in India or the Gondwanaland slightly later in Triassic. This may suggest a clue to the association of Permian
Glossopteris and the so-called Triassic Dicroidium in Nidpur Belt.
Talchir formation earlier interpreted as glacial, periglacial, glacio-fluvial and glacio-lacustrine deposits, and
now suggested as glacio-marine in a storm dominated epeiric sea environment, being affected by a Permian
glaciation. However, some authors have suggested Upper Carboniferous age of this glaciation. In fact, Permo-
carboniferous glaciation is characterized by heterochroneity. It started at different times at different places,
continued for different spans. Thus, even though it started in Pennsylvanian (Upper Carboniferous) in many
southern continents, it might have taken place in India in Permian (Stanley 1989, 1993). This is corroborated by
other evidences.
There are a few cases in which age may be suggested from stratigraphic position or correlation on the basis
of similarities with other stratigraphic units.
(Cont...)
 Chapter 19 Planta 339

FACTSHEET 19.10 (Cont...)

Background Information:
Problem of Fixing Age of Gondwana Dormations

Thus, Lower Jurassic age is said to be based on similarities between Rajmahal intertrappeans and Yorkshire
flora (Seward 1933 in Sengupta 1988); latter dated from intercalated marine beds with characteristic
ammonites.
Ptilophyllum floral assemblage with elements similar to Rajmahal flora from Bhutan, are found below a marine
Jurassic bed. Similarly, Lathi Formation in Rajasthan, subcrop ‘infra- and lower intertrappeans from Malda
and Birbhum, West Bengal (Sengupta 1988).
Particularly in central and western India, Jabalpur Formation with Ptilophyllum flora is overlain by the Deccan
Traps, though there are no intertrapppeans with Gondwana plant fossils so far reported from the Deccan Traps.
At Jabalpur, the formation is capped by a clay bed to be succeeded by the Lameta Beds of Turonian (Middle
to Upper Cretaceous) age with typical dinosaurian vertebrates.
A last set of evidence comes from the radiometric ages of basaltic trap rocks between which the intertrappeans
yield Gondwana plants. Thus, Ptilophyllum flora from the Rajmahal intertrappeans occur between trap rocks
dated at 110+ Ma, i.e. Lower Cretaceous. However, stratigraphic position of samples are often not clear and
hence the data remain only estimates.
In more recent approach, ten palynoevents have been recognized, based on multiple criteria, viz. FAD, LAD,
abundance, absence, dominance and diversity of palynomorphs.

FACTHSEET 19.11
Land Plant Record of India:
Variation of Major Genera of Lower Gondwana

Raniganj: Glossopteris overwhelming,

Glossopteris retifera, G. conspicua abundant and characteristic
Barren Measure Glossopteris dominant
Barakar: Gangamopteris + Noeggerathiopsis << Glossopteris
Karharbari: Gangamopteris + Noeggerathiopsis > Glossopteris:
Glossopteris of more importance
Talchir: Gangamopteris + Noeggerathiopsis > Glossopteris

FACTSHEET 19.12
Land Plant Record of India:
Dicroidium, Characteristic of Triassic in Different Associations:
In Parsora Formation: Dicroidium hughesi, Noeggerathiopsis hislopi,
Vertebraria indica, Pterophyllum sahnii and Neocalamites toxii
In type area of Panchet Formation
Glossopteris, Schizoneura, Dicroidium, Podozamites
In Daigaon Beds
Glossopteris, Samaropsis, Cladophlebis, Pterophyllum, Dicroidium,
340 Part Three: Miscellaneous
FACTSHEET 19.13
Land Plant Record of India:
Lower and Upper Gondwana Floras Compared and Contrasted
Item Lower Gondwana
Range Up. Carboniferous-Permian
Formations Raniganj/Barren Measures/
Barakar/Karharbari/ Talchir
Group Pteridosperms preponderant
Sphenophyllales, Equisetales
and Cordaitales important
Filicales significant, rarer
Ginkgoales present
In widespread, continuous outcrops:
Distribution Damodar-Koel valley
Son-Mahanadi valley
Pranhita-Godavari valley
Satpura- Narmada valley
Glacial signature in Talchir Formation
Climate Rich coal and carbonaceous shale
in Barakar and Raniganj suggesting
humid, sapropelic condition
Iron carbonate in Barren Measures
aridity; red beds in central and southern
Indian basins in equivalents of Raniganj
suggest aridity or seasonal variation of
heavy rainfall and dry, hot spells.
Vertebrates make less important
companion of the rich Glossopteris flora.
NB: For generic and specific names of plant fossils of different groups see Factsheet 19.8.
Siwaliks of northern India and Karewas of
Kashmir. Both monocotyledons and dicotyledons
are represented. Evidence of grazer mammals
from the Siwalik deposits (such as equids, viz.
Hipparion, Equu, etc.) point to the presence of
grasslands (grasses representing monocot angio-
Upper Gondwana
Triassic-Lower Cretaceous
Jabalpur/Rajmahal/Kota/Panchet
Bennetiales in lower part;
Coniferales in upper;
Pteridosperms rare
Equisetales relatively poor;
Filicales many important genera
Marattiales,Cordaitales
and Ginkgoales present
Occur in widely apart basins
Mainly Triassic in DK valley
Less important in SM valley
Jurassic important in PG valley
Mainly Lr. Cretaceous in Satpura basin
In Rajmahal basin
In a few coastal and inland basins in
Andhra Pradesh, Orissa, West Bengal,
in eastern India and Kachchh,
Jaisalmer and Cauveri basins,
Red beds in central and southern Indian
basins particularly in Triassic part of the
succession suggest aridity or seasonal
variation of heavy rainfall and dry, hot spells;
large reptiles suggest presence of big water
bodies and, thus, stand against total aridity.
Some amount of coal in uppermost parts
suggest return of humidity in Cretaceous.
Vertebrates significant in many different
parts; in Triassic Lystrosaurus, in Jurassic
sauropods
spermous ground-plants) in addition to the forests
in which the Siwalik mammals thrived. Forests
continue to grow in India in quite large volumes
and in their different types (tropical rain forests,
estuarine mangrove forests, mountainous forests
of northern, eastern and southern India, etc.).
 Chapter 19 Planta 341

Asses living in the Raan of Kachchh and Kiang in
Tibet continue as remnants of the equid fauna in
their now restricted habitat.
19.5 Some Relevant Questions
on Gondwana Stratigraphy
Details on the composition of the Gondwana floras,
relative importance of different major groups as
well as major genera, differences between the
Lower and the Upper Gondwana floras are all
entered in Factsheets 19.6 to 19.8, 19.14, and 19.11
to 19.13. It is, however, felt necessary to present
and discuss, in brief, a few important questions on
Gondwana Stratigraphy that have close relevance
with a discourse on floras.
Classically Gondwana succession is
considered largely continental with occasional
marine associations and intercalations. Broadly the
depositional environment of the different
formations may be summarized as below.
Talchir: Glacial valleys in southern uplands
and a broad delta plain crossed by tidal channels
in the low-lying northern Son-Mahanadi area.
Karharbari: Braided bars of low sinuosity
river succeeded by coalescing channel bodies topped
by fining upward cycles with coal capping them.
Barakar: Sandstones are channel deposits, with
interbedded shales corresponding to vertical accretion
of leaves and the coal as deposits of peat swamps in
flood plain and lakes of meandering streams.
Barren Measure: Sandstone-shale-iron-
stone repetition interpreted as channel and flood
plain deposits of meandering streams; the absence
of coal and the presence of local phosphorite remain
unexplained.

FACTSHEET 19.14
Summary Information About the Major Groups

Psilosida: Psilophytes, the earliest vascular land plants may be discussed on two genera, Rhynia and
Asteroxylon. Of them Rhynia had no true roots, its aerial shoots being without leaves, rose from a horizontal
subsurface stem (rhizome); they branched into two or three (dichotomous/trichotomous); the plant was
homosporous.
Asteroxylon was with an upright stem of a single dominant axis which grew straight up from a growing tip
(monopodial) and sideshoots; stem and sideshoots were covered by leaf-like scales; some sideshoots bore
sporangia, of homosporous type.
Lycopsida: Lycopsids include modern club-mosses; they are heterosporous, with small leaves. Spores formed
in cones that were clusters of fertile, scaly leaves(sporophylls); branching was combination of dichotomy and
monopodial.
Lepidodendron is a lycopsid plant with secondary wood. In it, at the foot of the stem there were branching root
system with spirally arranged rootlets, which when shed, left characteristic scars in the stem, the form genus
Stigmaria. Form genus Lepidophylloides is the leaf of the plant, Lepidostrobus is the large cone. Giant lycopsids
were among the main components of the European Coal Measures. Lycopsid spores are abundant in Palaeozoic
and Mesozoic and are helpful in zonation.
Sphenopsida: Sphenopsids are jointed plants, Equisetum (horsetails) being the extant example. It has a
rhizome system, vertical stem from it and radially arranged appandages (actually jointed stems, not leaves) at
nodes; leaves very reduced; homosporous, but functionally heterosporous. Sphenophyllum (Carboniferous)
had delicate stems and fan-shaped leaves in whorls. Calamites were giant sized with secondary wood with
large central pith cavity. Calamites and Annularia (radiating leaves) common in Coal Measures of the northern
hemisphere.

(Cont...)
342 Part Three: Miscellaneous

FACTSHEET 19.14 (Cont...)

Summary Information About the Major Groups

Filicopsida or Filicales: Filicales were, and are, true ferns, frond-like leaves grow from a central crown, close
to the ground or at a height on trunk; they have fibrous roots; they are heterosporous, spores kept in sporangia
underneath the pinnae, gametophyte, whose spore gave rise to, is innoccuous; sperms from the male organ
swim in films of moisture to the female, which on fertilization grows into the spore-producing fern again. They
were subordinated to pteridosperms in Permo-carboniferous, though significant in Mesozoic.
Pteridospermales: also called cycadofilicales because of the fern-like leaves of these plants, but with development
of seeds as in cycads; seeds borne upon the fronds either laterally or apically, but not in cones, and always
singly. Found from a rare specimen of frond and seed together; microporangia not on undersides of pinnae (as in
ferns); some arborescent/creeping/herbaceous/larger ones with enough secondary wood within their trunks.
Gymnosperms:
l Including Cordaitales and Coniferales of Palaeozoic, it covers the non-flowering seed plants. It also includes
Ginkgoales, Cycadales, Bennnettiales (Cycadeoidales belonged to this group).
l A seed is basically a dispersal mechanism. It dates back to Upper Devonian, though seed-bearing plants
(seed-ferns) became dominant in Permo-Carboniferous. The sexual generation develops on the parent
plant. Male sex cells modify into pollens, which travel by air, at the tip of claws of birds or otherwise to
the fixed female organ on the cone or flower of another plant. After fertilization, an embryo plant is
formed.
l Conifers have woody cones, from which seeds are dispersed by wind. Bennettiales (Ptilophyllum;
Williamsonia) are extinct relatives of cycadales. The two differed on the way they bore cones.
Extant Ginkgo is native to China.
Coniferales : Pollens have wings or air-sacs, leaves are typically needle-like ad spirally arranged (Elatocladus)
(largest trees living or fossil : Redwoods 100 metre height); pine, larch, etc. Conifers became important only in
Mesozoic, were overshadowed by angiosperms in later periods.
Cordaitales : produce dense secondary wood, extensive root system, probably lived on drier grounds in comparison
to the earlier other groups; they are, thus, generally drifted; reproductive structures borne on axils of leaves and
seeds in cone-like clusters.
Angiosperms:
l The most abundant group of the Tracheophytes, angiosperms appeared in Cretaceous, but there are possible
angiosperms in Jurassic, and even in Triassic.
l Early angiosperms were woody, shrubs and climbers; soft herbs or groundplants appeared in Mid Tertiary
(Oligo-Miocene: see discussion on evolution of Equidae/ Hominidae).
l Seeds protected by a thick coat (cryptogams); may have two seed leaves (dicots: leaves with mesh-like
venation) or one seed leaf (monocot: with parallel venation; grasses and palms).
l Evolution of flower based on meagre evidence. Flowers are specialized structure, little more than a colourful
cone, generally bearing both male and female organs, viz. microsporangia, called anthers set on thin
filamentous parts, together with carpels (female organs, female gametophytes that capture the pollen for
fertilization).

Raniganj: Braided to meandering streams Kota: The lower clastics are fluviatile, while
from bottom to top. the upper carbonates are lacustrine.
Panchet: Bed load deposits of braided streams. These largely continental Gondwana rocks with
Supra-Panchet: Same as of Panchet. terrestrial fossils lack adequate definite evidence for
 Chapter 19 Planta 343

FACTSHEET 19.15
Recent Evidences of Marine Transgression in the Gondwanas

1. Foraminifer from Talchir and Kulti Formations of West Bokaro and Raniganj basins, respectively
2. Marine pelecypods from Raniganj basin
3. Palynoflora, microfauna and trace fossil from Barakar Formation of Saharjuri basin
4. Marine bivalves and brachiopods from Talchir Formation of Son-Mahanadi basin
5. Marine coccoliths from Talchir, Karharbari and Barakar Foramtions
6. Abundance of whole rock Boron and P2O5 in sediments and organic sulphur in Barakar Formation coals
of Wardha Valley
7. Shallow marine ichnofacies
8. Shallow marine cryptalgal stromatolitic bioherms from Talchir Formation of Talchir Basin

age fixation. Biostratigraphically important bedrock-trough confined glacigenic facies,

evidences that help fix the age of Gondwana
Formations or its lower or upper age limits include
(i) association or intercalation with marine-fossil
bearing rocks (facies 3 in Factsheet 19.18), (ii) age-
diagnostic fossils, (iii) stratigraphic position, and
(iv) other evidences (Factsheet 19.10). More recently
palynofossils have been used for biostratigraphy of
Gondwana rocks. They have also been used in
deducing the then climates (see section 20.4.3).
Further, important modifications have been made
in regard to palaeoenvironment palaeogeography
of the Gondwana successions. They relate mainly
to marine incursions (also see Factsheet 19.15).
In the background of these, two stratigraphic
questions, yet undecided, may turn out to be
important leads for future work. Of them, one may
be framed as : How far important are the marine
associations in the Gondwanas ?
As mentioned earlier, marine associations or
intercalations help fix the lower and upper age
limits of the Gondwana Group/Supergroup. In
recent years there have been increasingly more
suggestions of the Gondwana successions being
associated with or containing marine sediments.
For example, Talchir formation was earlier
interpreted as glacial, periglacial, glacio-fluvial
and glacio-lacustrine deposits. Subsequently, it
has been suggested that the formation may be
glacio-marine with three facies assemblages, viz.
shoreface facies and slope facies. Occurrence of
hummocky cross-beds, domal sand waves, wave
ripples, ripple cross-laminations, etc. in repeated
succession has been accepted as indicative of a
storm dominated epeiric sea environment. Even
the coal basin development has been reinterpreted.
Prolific coal seam cycles through time and through
repeated adjustment in base level in coal swamps
plus widespread paralic signatures are believed
to stand against a craton-interior setting of
Gondwana coals. Other new evidences of marine
transgression in the Gondwanas have come up
recently (see Factsheet 19.15).
However, preponderance of terrestrial
vertebrate fauna and land flora in most of the
important stratigraphic units bear strong evidence
of a dominantly continental character of the
Gondwana successions even where they are
associated with contemporary marine rocks. Thus,
the issue ‘How far important are the marine
associations in the Gondwanas?’ needs more
attention and is yet to be viewed as settled.
The second question, or rather the second set
of questions crop up from recent suggestions
concerning definition, age fixation and
classification of the Gondwanas. They are : Which
rock body should be taken as the Gondwana
Group/Supergroup? From when did it start and
when did it end? How should this be classified?
344 Part Three: Miscellaneous
The two-fold or three-fold classifications of
the Gondwanas are based on recognition of breaks.
However, controversies galore on the nature,
importance and extension of the breaks and thus,
their significance in classification.
In an added suggestion, Sarbadhikari (1978)
suggested a major stratigraphic break at Jurassic
in the Indian Gondwanas. He contended that the
only definite Jurassic formation was the Kota. Rest
of the formations considered Jurassic, in part or in
full, do not provide any definite evidence of that
age. The opinion was not accepted by many
authors. For example, Sengupta (1988) stood for
Jurassic age for Rajmahal Formation on
similarities between Rajmahal intertrappeans and
Yorkshire flora (as held by Seward 1933 in
Sengupta 1988); latter was dated from intercalated
marine beds with characteristic ammonites.
Moreover, Ptilophyllum floral assemblage with
elements similar to Rajmahal flora occurring in
Bhutan, are found below a marine Jurassic bed ;
Lathi Formation in Rajasthan, subcrop ‘infra- and
lower intertrappeans from Malda and Birbhum,
West Bengal also belonged to that age (Sengupta
1988). More recently, however, authors from
different lines seem to converge on the opinion of
a Jurassic break. Thus, Bandyopadhyay (1999)
while reviewing the Indian Gondwana vertebrates
stops at Middle Jurassic (Kota) . Again Veevers
and Tewari (1995) in their still broader review of
tectonics, stratigraphy and other aspects of the
Indian Gondwanas follow the same line.
Implication of this suggestion is that, should then
the Lower Cretaceous parts of the Gondwanas, viz.
Jabalpur Formation, Rajmahal Formation and and
those of the east and west coasts be included in
the Gondwanas or not. In this regard, it may be
mentioned that Cretaceous Gondwana units are
much more scattered in comparison to the relatively
more continuous occurrences of Permian to Middle
Jurassic, rather Permian to Triassic formations.
There have already been suggestions if the coastal
Gondwanas (in Andhra and elsewhere in the east
or in Kachchh in the west) which are largely
associated with marine rocks should be included
in the Gondwanas at all, simply and solely on the
ground of their containing Ptilophyllum or other
fossils. In Rajmahal basin, too, the Gondwanas
occur as a completely different facies as
intertrappeans and the same question as that with
coastal occurrences, may be raised with the
Rajmahal formation too. The Jabalpur formation
with siliciclastics is overlain by the Lametas. The
latter include fluvial sediments in the lower parts
that provide evidences of pedogenesis or
calcretisation and occur conformably and in
continuum with the Jabalpur siliciclastics. This
suggests that the Jabalpur formation may really be
more closely related to the Lametas than they are
to the Gondwanas (Sarbadhikari 1978).
This question “Should the Indian Gondwanas
be considered to have ended in Middle Jurassic ?”
albeit fundamental, then remains a moot question.
To answer it, we must not lose sight of the fact that
the Ptilophyllum flora starts from Triassic and
continues through Jurassic to Lower Cretaceous.
If Lower Cretaceous be excluded from the typical
Gondwanas, occurrence of the Ptilophyllum flora
in the relevant formations must be adequately
explained. It should also suggest how the two-fold
(Lower of Permian and Upper of Triassic to Lr.
Cretaceous) or three-fold (Permian/Triassic/
Jurassic-Lr.Cretaceous) schemes of classification
should be treated.
The beginning of the Indian Gondwanas is
more universally accepted as Permian, though in
more recent times, Veevers and Tewari (1995)
marked the initiation of the Gondwana basins at
Upper Carboniferous.
19.6 Brief Critical Appraisal of
Indian Record
The above discussion of the Indian record of land
plants shows:
1. The whole course of plant evolution is
represented in India, though not with uniform
development of all parts or in perfect
continuity.

2. This is a meagre record of earlier parts, viz. of
psilophytes and lycopsids.
3. Prolific representation of pteridosperms and
gymnosperms mark the floras.
4. Relatively less prominent record of angio-
sperms demands attention.
With this, there remains to add:
5. There were definitely rich flora present on this
land, as evident from:
(a) coal reserves and carbonaceous shales in
Palaeozoic- Mesozoic and in Tertiary; and
(b) the presence of large herbivorous
dinosaurs and other reptiles in Mesozoic
and mammals in Cenozoic.
A critical appraisal of the Indian continental
flora must take into account answers to the question:
why was not then the likely-to-be rich floral record
represented in uniform prominence all through. The
answer has been provided at the beginning of the
chapter. We can reiterate with a few more words.
Till late Palaeozoic, the Indian craton, formed
in Precambrian, existed as a stable land mass. Hence,
the preservation potential of the early land flora was
poor. It was only when India suffered a tectonic
readjustment at the beginning of the break-up of
Pangaea, that the Gondwana basins started to form.
In fact, the basins may be viewed as the precursors
of the tectonism that produced rifting of the Indian
plate. Continued tectonism favoured synse-
dimentational subsidence of the basins resulting in
thick clastic sequence of the Gondwanas.
At the same time, these basins were formed
on a peneplained land mass on which the river
systems did not have sharp gradients. They were
braided or meandering, as have been found from
sedimentological work of scores of authors.
It implied that the clastics had a considerable
percentage of fine argillaceous components. It also
favoured rather swampy condition on a low-lying
plain that could develop adequate and widespread
humus from the vegetative matter, to be
transformed into coal subsequently.
This accounts for the formation of huge coal
reserve in the Gondwanas and also considerable
Chapter 19 Planta 345
thickness of relatively impermeable shale and
siltstones for plant fossils to be preserved in good
condition.
It, thus, appears that the rich Gondwana flora
was the result of an interplay of floral growth,
tectonic background and taphonomically
favourable ambience for preservation.
This combination never repeated itself in later
times. Immediately after the development of the
Gondwana Group (or Supergroup), there was the
widespread volcanic activity that produced the
Deccan Traps. It again gave way to a flat-topped
geomorphology that was never going to be suitable
for strong river systems. The intertrappeans, thus,
hosted only swampy, paludal flora and fauna that
could be preserved in the relatively smaller and
shallower lakes and ponds in occasional depressions
on the flat trap country. The floral record did not
reach abundance, howsoever abundant might have
been the vegetation.
Barring the sporadic occurrences of Cenozoic
deposits in the coastal belt of India, the next
imporant continental deposit of the country was the
Siwalik Group (or Supergroup). Having been
derived from the rivers coming down from the
newly emergent, collision-tectonics-derived folded-
belt mountains, the Siwalik clastics were more of
coarser grains, with fine components represented
in lesser amount. Naturally, both the energy
condition at the depositional locales and the
preponderance of coarser clastics, did not prove
ideal for preservation of plant materials, even
though the region had both extensive and thick
forests (as proved from the presence of fossils of a
rich proboscidean, primate and carnivorous fauna)
and grasslands indicated by the equids and giraffids,
in particular. In summary, the rich gymnospermous
flora of Indian Mesozoic might have evolved into
an equally important angiospermous flora. But the
tectonic and taphonomic conditions did not prove
favourable for adequate and good preservation of
the floral remains.
It is felt that an understanding of the Indian
flora should take into account these factors and
developments. (see Factsheet 19.16)
 346
FACTSHEET 19.16
Land Plant Record of India:
Some Common Genera of the Indian Gondwanas
Leaf genera
Glossopteris Schizoneura Noeggerathiopsis Pecopteris Cladophlebis Gleichenites
Simple/compound Simple Simple Simple Compound Compound Compound
Bipinnate*1 Bipinnate*1 Tripinnate*1
Arrangement Two at a node Opposite Alternate Alternate
Shape (of pinnules Spatulate Lenticular Spatulate Subelliptical Fulcate Orbicular to
for compound leaves) subdeltoid
Margin Entire Entire Entire Entire Entire Entire
Part Three: Miscellaneous

Base Narrow, Acute, Narrow, Broad, Broad, Broad,

Attachment Petiolate Sessile Sessile Sessile Sessile Sessile
Apex Broad, Sharp, Broad, Broad, Sharp, Broad,
rounded acute rounded rounded acute rounded
Venation Branching Parallel Parallel Mid vein, Mid vein Single vein
Reticulate*2 Convergent Divergent lateral veins lateral veins may be
with mid-vein rare bifurcation at angle forked bifurcating
( *1Sengupta (1988); * 2lateral veins branch out and anstomose to form mesh or reticulation)
Ptilophyllum Otozamites Dictyozamites Pterophyllum Taeniopteris Elatocladus
Simple/compound Compound Compound Compound Compound Simple Simple
pinnate pinnate pinnate pinnate
Arrangement Alternate Alternate Alternate Alternate Spiral about stem
Shape (of pinnules Fulcate, long Fulcate, short Fulcate, long Curved rectangular Spatulate Acicular
for compound leaves)
Margin Entire Entire Entire Entire Entire Entire
Base Broad Broad Broad Narrow Petiolate Broad, sessile
Attachment Sessile Sessile Decurrent Sessile Overlapping
Apex Sharp, Broad, Broad, Broad, Broad, Sharp,
acute rounded rounded truncate truncate acute
Venation Parallel Parallel, a few Parallel in the Parallel Parallel with Parallel
bifurcating middle, reticulate a mid vein,
along margin lateral veins at rt<s

(Cont...)
 FACTSHEET 19.16 (Cont...)
Land Plant Record of India:
Some Common Genera of the Indian Gondwanas

Leaf genera
Thinnfeldia Dicroidium Sphenopteris
Simple/compound Compound Compound Compound
pinnate pinnate bipinnate*1
Arrangement Alternate Alternate Alternate
Shape (of pinnules Subtriangular- Fulcate, short Elliptical
for compound leaves) broad elliptical
Margin Lobed (3) Entire Slight wavy
Base Narrow Broad Narrow
Attachment Decurrent Sessile Sessile
Apex Broad, Broad, Narrow,
rounded rounded rounded
Venation Branching and Parallel, a few Branching and
forking bifurcating forking

*1
(Sengupta 1988)
Stem genera include:
Vertebraria, a rhizome, rectangualr parallel sided, divided into two or more columns of rectangular blocks by longitudinal and transverse
grooves; blocks themselves are striated longitudinally and transversely.
Schizoneura is a common stem of Lower Gondwanas rarely found with two leaves attached at the node at acute angles. Stem is long, parallel
sided, articulated into nodes and internodes. Nodes very sharp and internodes long; parallel grooves run continuously across the nodes. The
genus differs from Phyllotheca, principally on the continuity of grooves across nodes; in Phyllotheca grooves are offset across nodes.
Dadoxylon is fossilized wood, often quite large and generally preserved in permineralized form (silicified); shows well-formed annular rings
striations on the surface and pith cavity.
Brachyphyllum is an elongate, narrow leafy shoot (twig); it branches repeatedly; the shoot is covered by small leaves of subelliptical or lozenge
shaped, arranged spirally and imbricating, i.e., overlapping; leaves have their long dimensions parallel to the shoot-axis, acute apex entire
margin and curved base.
Chapter 19 Planta

Bucklandia is an irregularly branching stem with leaf bases marked on it.

347
348 Part Three: Miscellaneous

19.7 Calcareous Algae 1. They are photosynthetic, with chlorophyll

Though calcareous algae do not belong to the
higher plants, they warrant a brief treatment on
account of their geological importance.
Algae form a large and diversified informal
category of aquatic, non-vascular, often chlorophyll-
bearing plants, ranging from microscopic cells, a few
micron in size to enormous benthonic seaweeds tens
of metres in length.
Calcareous algae are various kinds of these
organisms whose thalli (sing. thallus) contain
biochemically precipitated calcareous skeletal
material. In additon, deposits of calcium carbonate
caused by algae, which are usually an interaction
between biological and physical (sedimentological)
processes are also grouped with the calcareous
algae. They include stromatolites-oncolites and
algal-laminated sediments.
Three genetic categories may be recognized;
they are skeletal calcareous algae, laminated
calcareous structures and freshwater calcareous
tufa (see Factsheet 19.17).
Biologically important calcareous algae are
characterized by the following features:
but other pigments often mask its effect.
2. They evolve oxygen during photosynthesis and
require it for respiration.
3. Thallus (the plant body) is non-vascular and
not differentiated into root and shoot.
4. Some are prokaryotic (cyanophytes), others
eukaryotic.
5. Thalli may consist of single cells, colonies of
single cells, filaments may be branched or
unbranched, cellular tissue.
6. Identified on growth form and external
geometry, internal structure and skeletal
microstructure.
7. By growth form, they may be encrusting or
spread over the substrate, attached to it, or erect
plants or filamentous.
8. Major groups of fossil benthonic skeletal
calcareous algae are shown in Factsheet 19.16.
9. Skeletal mineralogy differs, as shown in
Factsheet 19.18.
10. They live in aqueous or moist environment.
Indian record of calcareous algae is
summarized in Factsheet 19.19.

FACTSHEET 19.17
Calcareous Algae: Categories

Skeletal calcareous algae: Taxonomically diverse types; calcium carbonate is deposited as a result of metabolic
and biochemical processes in various habits and concentrations within and upon the entire plant body or may be
localized in only a portion of the plant.
Laminated calcareous structures: Produced primarily by mechanical (rather sedimentological) accumulation
of fine-grained (micritic) calcium carbonate particles, on organic films or mats generated by colonies of filamentous
algae or planktic coccoids (both without skeleton). Periodic interaction of algal mats and physical sedimentation
as also metabolic secretion of calcium carbonate, produce distinctly laminated organosedimentary structures,
stromatolites (or oncolites : on shifting or moving substrate, e.g. pebbles or tests of foraminifera subject to
current action) where the organism is cyanophytes or blue-green algae, or algal laminated structures produced
by any other type of algae, like crustose red algae or melobesioid rhodophytes.
Freshwater calcareous tufa: Concentrically laminated and densely so, because of biochemical origin, produced
by photosynthesis of non-skeletal algae (e.g. blue-green algae) .
 Chapter 19 Planta 349

FACTSHEET 19.18
Calcareous Algae: Types and Mineralogy

Division Family Subfamily Mineralogy Environment

CYANOPHYTA .... Calcite Nonmarine-marginal
(blue-green algae)a
RHODOPHYTAb Corallinaceae Melobesioideae Calcite Marine, cold to warm,
(red algae) (crustose) shallow to deep,
Corallinanoideae Calcite Marine cold to warm,
(articulated) shallow to deep restricted
niche of genera and species
Solenoporaceae Aragonite Marine
CHOLOROPHYTAb Codiaceae Aragonite Marine, shallow, warm
(green algae) Dasycladaceae Aragonite Marine, shallow, warm
CHAROPHYTAb Calcite Non-marine
CHRYSOPHYTA Coccolithophoraceae Calcite Marine, planktonic
PHEOPHYTAb Aragonite Marine
(brown algae)

a Now referred as cyanobacteria.

b included in the Kingdom Planta.

FACTSHEET 19.19

Calcareous Algae: Indian Record

l Common groups are

Rhodophytes (marine red algae), both crustose (melobesioids) and articulated (corallinoids)
Chlorophytes (marine green algae) and
Charophytes (non-marine green algae)
Cyanophytes (blue-green algae) in Stromatolites in different Precambrian units
l Marine groups occur in Upper Cretaceous rocks of central India (Bagh Beds) and southern India (Caveri
Basin) and
in Tertiary (both Palaeogene, particularly Middle Eocene and Neogene) rocks of
Kachchh, Assam, Andaman Island, Samana Range Pakistan
l Non-marine Charophytes
reported chiefly from Deccan Intertrappeans of central, western and southern India
l Cyanophytes in stromatolites in different Precambrian units
l Some commonly reported genera:
Melobesioideae: Melobesia: Eocene-Recent Lithothamnium Jurassic-Rec.
Lithoporella Jurassic-Recent Archaeolithothamnium : Jurassic-Rec.
Mesophyllum Palaeocene-Recent Lithophyllum Jurassic-Rec.
Corallinanoideae: Corallina Jania
Dasycladaceae: Acicularia Jurassic Recent Cylindroporella Jurassic-Cretaceous
Neomeris Cretaceous-Recent Acetabularia Palaeogene-Recent
Diplopora Permian-Triassic Macroporella Carboniferous-Jurassic
Codiaceae: Halimeda Cretaceous-Recent
Charophyta: Chara Gyrogonites
 Part four
Some Synthesis
 20
Fossils and
Climate, Geography,
Biogeography, and
Ecology of the Past
20.1 Introduction
Physical geography deals essentially with land-sea
distribution and associated effects on the biosphere
or the atmosphere; palaeogeography covers
corresponding aspects of the geological past. Hence,
the primary task of palaeogeography lies in the
reconstruction of the land-sea distribution of a region
or the globe for a particular segment of geological
time or through different geological periods.
However, this branch of palaeontology, or rather of
geology, assumed special importance over and above
being a descriptive or reconstructive science, during
the course of its development.
The recognition that the earth’s geography
changed continuously through time, even led
geologists to modify their way of viewing the earth
itself. Continents and oceans or such other features
on the surface of the earth were no longer to be
353
considered as fixed in time-space continuum.
Rather they had their origin, and even death, and
were constantly changing their positions on the
surface of the earth. In the wake of this recognition,
palaeogeography gave rise to radically marked
concepts of continental drift, utilizing tangible
materials or evidences from any and every line of
studies to reconstruct the geographies of different
times of the geological past. Fossils make one of
the most important set among these materials and
evidences for palaeogeographic reconstructions.
At the same time, as we know from the recent
biosphere, each living species has a geographic
range, varying from a single pond to an entire
continent. Thus, ourselves, the species Homo
sapiens have a virtually worldwide range; the
kangaroo is native to Australia and Tasmania, the
present-day Hippos to Africa. Studies on
geographic distribution of modern organisms are
354 Part Four: Some Synthesis
covered in biogeography, which brings out the
existence of biogeographic provinces with typical
faunal and floral elements. It also explains the
barriers that are set to these organisms to maintain
the provinciality, as also the instances or situations
in which the constituent organisms migrate out of
some province or into another.
Knowledge of fossils reveal that the organisms
might have had different geographic distribution in
the past; thus hippos, of our earlier example, were
very much there in India, in the Siwaliks, or even
the Peninsular India in Subrecent times.
Palaeobiogeography takes care of these patterns and
their changes of biogeographic distribution of
organisms in the geological past. It is a multi-
disciplinary approach and serves both geography
and palaeontology. Thus, palaeobiogeography has
brought out the presence of ancient oceans and
island complexes, microcontinents to super-
continents, their connections and breaches. On the
other hand, it studies variations in morphology or
diversity with differences in geographical settings.
The subject is an upcoming branch of palaeontology,
though its essence was recognized for over centuries.
Similarly, palaeoclimatology, often considered
as a part of palaeogeography is another freshly
innovated, important branch of palaeontology in
reconstructing or understanding the earth’s history.
Intrinsically, it is concerned with the study of
climatic patterns in the geological past, again on
the basis of tangible criteria of lithology, fossils
and any such others.
The present chapter will deal with the three in
a reasonably brief outline. Since the three along
with palaeoecology are largely interlinked, their
understanding and treatment are likely to bear
occasional overlaps.
20.2 Palaeogeography
Palaeogeography and the theory of continental drift
are interwoven in their growth. So, we may begin
our discussion with a brief history of how the
theory came into life.
20.2.1 Brief history from continental
drift to plate tectonics
The changing face of the earth was amply
recognized in the later half of the 15th century.
In Leonardo da Vinci’s words “... above the plains
of Italy where flocks of birds are flying today, fishes
were once moving in large schools”.
About one and a half century later, Francis
Bacon noted that the shorelines of western Africa
and eastern South America were similar, though
he did not suggest any continental fit.
Alexander von Humboldt, about two centuries
later attributed these similarities to erosion.
Antonio Snider-Pellegrini in his book Creation
and its Mysteries Revealed, suggested, on the
evidence of similar plant fossils in Europe and
North America, that all continents were linked
together during Carboniferous (Pennsylvanian) and
were later split apart. The cause was, in his opinion,
the deluge described in the Bible.
During the late 19th century, Edward Suess
brought out the similarities in Upper Palaeozoic
flora of India, Australia, Africa, Antarctica and
South America, as also the evidence of glaciation
in the stratigraphic succession of that age. He
proposed in 1885 the name, Gondwanaland for the
supercontinent of these southern hemisphere
landmasses. He thought that the continents were
connected by land bridges that were later
submerged.
In 1910, Frank Taylor proposed a theory of
continental drift from a large polar continent, in
which the fragmented parts moved towards the
equator to assume their present position and shape.
He ascribed the cause to gigantic tidal forces,
which was generated when the Earth captured the
Moon about 100 million years ago. He also
suggested that the mid-Atlantic ridge, discovered
by the Challenger expeditions during 1872-1876,
might mark the line along which an ancient
continent had broken apart to form the present-
day Atlantic ocean.
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past
It was not until Alfred Wegener in 1915
(in his book The Origin of Continents and Oceans)
followed by Alexander du Toit in 1937
(Our Wandering Continents) that the theory of
continental drift took real shape. Wegener proposed
a single supercontinent Pangaea (Greek: all land)
on a vast amount of evidences, geological,
palaeontological and climatological, whereas du Toit
is to be credited with developing the idea of Laurasia,
a northern supercontinent with North America,
Greenland, Europe and Asia minus India, in addition
to the southern Gondwanaland, the former
reconstructed on the evidence of contemporary coal
deposits of Carboniferous age. Du Toit maintained
that the glacial deposits of the Gondwanaland
pointed to its presence near the South Pole, whereas
the Laurasian coal deposits indicated their near-
equatorial positions. Both the sets of continents, thus
moved from their original positions to the present
ones. Wegener also gave a series of maps showing
the fits of continents and different positions as they
gradually broke apart through time.
In spite of overwhelming evidences in favour
of the movement of continental land masses on the
surface of the earth, the theory of continental drift
faced vehement opposition for want of a suitable
mechanism that could have made the continents
move. It was even suggested in 1944 that “further
discussion of it merely encumbers the literarture
and befogs the minds of students”.
The advent of the concept of Plate Tectonics
by Harry Hess in his landmark paper in 1962,
brought doom to the theory of continental drift,
but finally vindicated its content, “the continetal
land masses have grown and have constantly
changed their positions through geological time,
at least during a major part of the Phanerozoic,
when the crust had grown adequately thick and
large to behave in association with an upper part
of the mantle as rigid ‘lithospheric’ plates and the
earth’s interior has attained a thermal condition to
sustain a process that could break apart the brittle
lithosphere into plates and could drag them along
on the surface”.
355
The concept of Plate Tectonics is so much
entrenched in geology today, that we need not
discuss details of it here. We mention here in the
Factsheet 20.1, evidences that come in support of
the aggregation and disaggregation of continents.
Two more Factsheets 20.2 and 20.3 are included
in this discussion, though they do not have direct
bearing in palaeontology. They relate to the
lithological types that are found useful for regional
palaeogeography and steps involved in the
reconstruction of palaeogeography of a region,
respectively. They are added only to indicate some
practical asepects of palaeogeographic studies.
20.3 Palaeobiogeography
Palaeobiogeography of ancient organisms was also
obvious to scientists for long. Thus, 19th century
biologists and geologists recognized similarity
between Scottish Lower Palaeozoic faunas with
those of North America, while those of the same
age of England and Wales were alike the faunas of
Europe and Scandinavia. This, along with other
evidences, led Scientists to conceive an early
Palaeozoic ocean system dividing Europe and
North America. There were also the classic
examples of similar reptilian fauna and flora of
the southern continent.
20.3.1 Modern biogeographic divisions
The present-day earth is said to have six main
biogeographical regions. These are:
1. Palaearctic: The eastern part of the northern
hemisphere beyond tropic areas, including
Iceland, Europe, the northernmost Africa above
the Tropic of Cancer and Asia without its three
southern peninsulas, viz. the Arabian, the Indian
and the Malaysian; it also includes the eastern
Atlantic and the western Pacific Oceans.
2. Nearctic including North America and
Greenland, eastern Pacific and western
Atlantic, thus covering western part beyond
tropic of northern latitude areas.
356 Part Four: Some Synthesis

FACTSHEET 20.1
Evidences of Aggregation and Disaggregation of Continents

l Fit of shorelines of continents, modified into fit along continental slopes where erosion would be minimal;
best fit of Pangaea at about 2000 metre depth, confirmed by latest ocean basin data.
Now on widely separated continents, the occurrence of:
l Similar rock sequences of Carboniferous to Jurassic age in five Gondwana continents and mountain ranges
of same age and deformation style of North America and Canada (Appalachian Mtns) and Greenland,
Great Britain and Norway.
l Contemporary continental glacial deposits of Permian age, evident from tillites and glaciated pavements
in the Southern Hemisphere and virtually contemporary coals and tropical plants in the Northern Hemisphere
indicating similar palaeoclimatic zones.
l Similar extinct fauna and flora; continental Glossopteris flora, freshwater Permian reptile Mesosaurus
(from South America and South Africa), Lower Triassic land reptiles like Lystrosaurus (Africa, India and
Antarctica) and Cynognathus (South America and Africa), all point to the absence of intervening oceans
between these lands of the Gondwanaland. In fact, northern and southern continents (Laurasia and
Gondwanaland, respectively) were characterized by different sets of dinosaurs and smaller reptiles.
l Palaeomagnetic studies indicate that apparent movement of the magnetic pole of a single continent through
time can be explained either assuming the continent fixed and magnetic pole moving, or the magnetic pole
still and the continent moving or both the continent and the pole moving. Analysis of all continents in their
present position, indicated each continent had its own series of magnetic poles, an unlikely proposition, that
could be resolved only with the assumption that the continents moved through time.
l Evidence of sea floor spreading suggests that the continents and the oceanic crust move together as the
sea floor separates at oceanic ridges where new crust is formed by upwelling magma. It not only attests to
the movement of the continents; it leads to conceiving the mechanism of such movement, from the formation
of new crust-sea floor spreading- subduction cycle to the thermal convection cells beneath them.

FACTSHEET 20.2

Lithological Types Useful for Regional Palaeogeography

(Ziegler et al., 1985)

Clastic-carbonate sediments
Conglomerate, Sandstone, Mudstone, shale Carbonate
Climatically significant sediments
Tillite and glacio-marine beds Peat.coal Dolomite Reefs
Gypsum, anhydrite (Evaporites) Halite and bittern salts (Evaporites)
Oceanographically significant sediments
Bedded chert, radiolarite, diatomite Phosphorite Chamosite Glauconite
Ferromanganese nodules and concretions Limonite, goethite or hematite
NB: In addition, acid to basic volcanic rocks or sequences like rhyolite-rhyodacite-trachyte-latite or basalt-phonolites-
basanites-dolerite dykes or acid to basic intrusive rocks or sequences or ‘cooling ages’ of uplift and unroofing may
be used in palaeogeographic interpretations of igneous-metamorphic terrains.
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past 357

FACTSHEET 20.3
Reconstruction of Palaeogeography of a Region: Steps Involved

l Observation:
On beds, their rock and fossil contents, thickness, primary structures, composition, texture, sedimento-
logical parameters, including vertical and lateral variations of all these aspects.
l Preparation of maps:
lithological/geological; facies; isopach, etc.
l Drawing up the succession
l Interpretation of:
Relief pattern of the basin, provenance of the sediments, current directions and depositional settings.
l Reconstruction of the geological history:
When did the succession start? On what basement?
Under what conditions? How they changed?
l Enumerating palaeoenvironment variation
l Reconstruction of basin history
l Similar information and study for other localities.
l Correlation on rocks and fossils
l Reconstruction of palaeogeography-palaeobiogeography and palaeoclimatology of the region.

3. Australasian with Australia, New Zealand
and the ocean around.
4. Oriental including the Indian and the
Malayasian Peninsulas and the East Indies
islands (Indonesia).
5. Ethiopian with the Africa south of the Tropic
of Cancer and the Arabian Peninsula.
6. Neotropical including South America, the
Carribean Islands and Falkland Island. The two
former regions are aligned east-west and
include temperate to polar climatic zones,
whereas the four latter are disposed north-south
from east to west, respectively. Obviously they
cut across the latitudinal climatic belts and
include all the four of humid tropics, dry
subtropics, warm and cool temperate and polar
zones. Antarctic region merges into the
southernmost ends of all the four and is not
regarded as a separate biogeographic region.
Apparently, in addition to the climatic control,
these biogeographic regions have also other factors
to define them. One important factor is the
continuity of land; any break on account of the
presence of large water masses (between Nearctic
and Neotropical or between Palaearctic and
Australasian, as also between the southern regions
themselves) or hostile terrains of high mountain
belts (between Palaearctic and Oriental, as also
partly Ethiopian) or deserts (between Palaearctic
and Ethiopian), acts as a major barrier to help
define the regions.
Changes in biogeographic distribution take
place when organisms move across their region on
account of some drastic and sudden changes in
their environment, again primarily climatic
(e.g. glaciation), as also otherwise, such as develop-
ment or removal of barriers or prolonged
environmental adversities like volcanism. There are
two models at present preferred, to explain these
dispersals. Before getting into them, it is advisable
to look through a few relevant definitions and
concepts (Factsheet 20.4).
20.3.2 Two models of biogeographic
changes
The two models to explain dispersal, particularly of
ancient organisms differ in assuming the relative
movement between the organism and its geography.
358 Part Four: Some Synthesis

FACTSHEET 20.4
Some Concepts and Definitions

Biogeographic province: Basic unit of the studies: defined on shared, i.e. same set of, orgainsms (taxa)
restricted to a particular geographical region; such taxa are called endemic. A province may be demarcated of
maps drawn in workable scale and is thus equivalent, in a sense, to lithostratigraphic formation.
Realms: Large regions with strong biotic contrasts at the level of higher taxa.
Taxa may be:
Eurytopic: Ecologically tolerant and thus living in a wide range of habitats.
Stenotopic: Ecologically sensitive and, thus, living in a narrow range.
Endemic: Restricted to a single geographical area, generally an island chain or a mountain range or a small
geographical territory.
Pandemic: Distributed widely across regions.
Cosmopolitan or global: Worldwide, though rare among land animals.
Disjunct: Distributed in widely separated areas either due to some tectonic processes, e.g. rifting or sea-floor
spreading, or due to development of some environments, such as anoxia or chemosynthetic environments.
Barriers to faunal and floral migration may be of three types (vide Simpson on examples of mammals):
Corridors: Restricted passages though open at all times;
Filters: Allowed selected movement to chosen groups or at particular times and
Sweepstake routes: occasionally open and access random.
In oceans and sea barriers are set by:
Expanse of oceans, ocean currents and salinity
The presence of ocean ridges and islands, and
Temperature gradient across latitudes producing four major belts, viz. tropical, warm-tropical, cold-temperate
and cold that may be interrupted by continental plates, ocean ridges, hydrothermal vents, etc.
Thermocline: A vertical barrier in oceans that varies in depth according to substrate topography, latitude,
seasons and water currents.

Thus, dispersal model presupposes that most
organisms originate from a point and gradually
spread out across continents or ocean basins,
enlarging their geographical ranges.
In contrast, the vicariance model assumes that
the geographical range is disjuncted from
fragmentation of the biogeographic province, due
to rifting or otherwise.
Dispersal is often followed by a period of
adaptive radiation. The organisms involved are
placed before new opportunities that trigger
radiation. Oceanic islands are useful stop-overs
during migrations for mobile organisms, even
larvae of benthic organisms. Such mode is called
island-hopping. In contrast, rafting, i.e. adapting
to a pseudoplanktonic mode of living, being
attached to some floating objects is another mode
of dispersal.
Vicariance model hinges on the plate tectonic
theory. Geographical ranges of the taxa are
modified by the break-up and rifting of both
continental and oceanic crust. Sea-floor spreading
in oceans and mountain building or development
of rift valleys on land may lead to disjunct ranges.
On the other hand, development of new land
connection due to collision of plates, may help
extend the geographical range.
Several instances of importance of
biogeography in palaeontology are discussed in
Chapters 5, 12, 14 and 18. Here an additonal
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past
example is given that may be considered,
particularly with reference to the migration in
equids that involves migration of these forms from
their centre of radiation in North America to Asia,
more so to the Indian Siwaliks.
Anourosoricini, a tribe in the family Soricidae
of Mammalia, which includes a kind of shrews with
carnivore-like specialization in dentition and
mandibular structure, provide a well-documented
pre-Quaternary example of southward migration
under the influence of climate.
Palaeoecological interpretation shows that
Anourosoricini avoid areas with low levels of
precipitation (a minimum of 600 mm/yr is about the
threshold value) and low soil humidity. On this basis
a proposed palaeobiogeographic model presumes
that southward mammal migrations are driven by
changes in the precipitation regime rather than the
temperature regime of the Mediterranean region.
The available evidence also suggests eastward
dispersal of a primitive Crusafontina species from
Eurasia into North America at about the same time
when Hipparion crossed the Bering Strait in a
westward direction (van Dam 2004).
Another added instance shows palaeo-
geographic changes modifying palaeobiogeography.
The early diversity and endemism of nautiloid
species during Late Bathonian-Early Callovian times
in Kachchh was interrupted by dominance of
cosmopolitan forms in later times. This is interpreted
(Halder 2000) as a consequence of the rifting of the
Gondwana supercontinent. A shallow basin that
developed with initiation of the drifting of the Indian
plate triggered early speciation events. Subsequently,
as sea routes developed with rifting, free faunal
exchanges across the Tethys gave rise to pandemic
faunal character.
20.4 Palaeoclimatology
Ever since the days of Charles Lyell (1797-1875;
the author of The Principles of Geology, in three
vols. 1830-1833 (see Summerhayes 1990), it has
359
been appreciated that climate is one of the first-order
controls in stratigraphy. In fact, it is one of the three
allocyclic factors (those factors in which changes
in energy and materials are external to the
sedimentary system) that control sedimentation and,
hence, stratigraphy (Cecil and Edgar 1994), the other
two being tectonics and sea level. Tectonic processes
put continents where they are and create topography.
Tectonic framework creates space for sediments to
accumulate in basins and itself modifies the climatic
regime. Sea level modifies land-sea configuration
and based on that climatic signals. Climate then
governs the kind of weathering, development of soil,
rates of erosion, means of transport (e.g. wind in
arid, river in humid), conditions in the environment
of deposition, nature of biota and, thus, much of the
development of stratigraphic record. Tectonic effects
are largely intrabasinal, whereas eustatic processes
are interbasinal; the two can be identified from each
other through correlation of basins. In the later parts
of the last century, factors controlling climate change
have also been more clearly understood. It is now
appreciated that the first-order effect of climate on
stratigraphy and sedimentation comes as a result of
movement of continents through latitudinal climatic
belts with time. There are modifying effects of other
factors such as orbital-forcing cycles, mountain
building, ocean circulation and variations in
atmospheric ‘greehouse’ gases.
Again, palaeogeography controlled by tectonic
processes has an important modifying effect on
climate. Distribution of climate-sensitive deposits,
on whose basis palaeoclimate is generally
interpreted, depends both on suitable and
favourable climatic regime and on a favourable
palaeogeographic-cum-physiographic setting.
Thus, in Triassic all the continents were held
together in a single supercontinent Pangaea with
mountainous coasts and high interiors. As a result,
coastal regions were dry or wet depending on the
climatic zone, but the vast interior was largely dry.
This may have been one of the reasons for the
development of extensive red-beds during Triassic.
360 Part Four: Some Synthesis
By contrast, in later periods, for instance in Middle
Cretaceous break-up of Pangaea led to the
formation of several discrete continents separated
by sea-ways that provided moisture for rain to pour
on the formerly dry interiors of the supercontinent.
20.4.1 Some facts about climate
The climate of an area is the characteristic weather
pattern extended over a long period. It depends on
the following:
1. Latitude that controls temperature and
distinction between seasons. In its turn,
latitudinal position determines the angle at
which the sun’s rays are incident on the earth’s
surface. At and around the equator, the rays
are at high angle, are focussed and
concentrated over a smaller area and, thus,
produce higher temprature. At higher latitudes
near the poles, the rays are incident at low
angles are, thus, spread and dissipated over a
larger area and, thus, cause lesser temperature.
This causes the temperature gradient across
the latitudes on the surface of the earth.
2. Climate also depends on the moving air masses
that prevail in the area from local or regional
causes. In general, temperature gradient also
gives rise to pressure gradients; hot air rises
up and the cooler, dense and heavy air fills in
the low pressure area rushing in from around.
This may bring in moisture to cause more
precipitation, i.e. rainfall or may be dry causing
aridity. Added effects come from the Coriolis
effect caused by the earth’s rotation, which
deflects winds to the right in the Northern
Hemisphere and to the left in the Southern
Hemisphere.
3. Climate is also influenced by the relative
distribution of land and sea, high ground and
low, the presence of forests, lakes, valleys,
glaciers and such others. Thus, coastal belts
tend to have more rainfall, a largely maritime
climate with seasons less differentiated;
whereas deep inland areas are drier, with
sharply marked seasons. The presence of great
reliefs causes high precipitation zone in front
of it towards the sea and a rain shadow zone
behind it. The present-day Himalayas in India
producing the wet belt south of it and the dry
region to its north is an example. Similar
example is also found in the west coast of
North America of Palaeogene times, that
caused the grassland to develop to the east of
the newly rising Applachian mountains,
providing a favourable niche for the evolution
of Equidae.
4. The present-day climatic variation is further
related to what is known as psychrosphere,
which developed and at the same time,
triggered the formation of polar ice caps
(see Factsheet 20.5).
20.4.2 Methods and examples
Palaeoclimate may be interpreted in various ways.
The most common, yet fairly dependable criteria
are provided by a host of climate-sensitive deposits.
Of them, particularly significant are coals,
evaporites, carbonates and tillites. Coals indicate
a moist climate that may be tropical or temperate
in particular. Evaporites indicate aridity, generally
hot. Carbonates are produced in large quantities
mainly in shallow warm water of the tropical belt.
A major amount of carbonates of the geological
past is biogenic and marine. Marine calcareous
biota, in turn, is more prolific in the shallow sea
water within the photic zone and they produce the
major portion of the calcareous deposits. These
climate-sensitive deposits have a distribution
pattern much similar to the pattern as found today.
It is marked by tight latitudinal zonation. Hence, it
may be inferred that the earth’s climatic system
has operated in the past much in the same way as
now. Hot air rose at the equator and moved away
before descending at middle latitudes under the
influence of the Coriolis force generated by the
earth’s rotation.
Plant fossils are used for palaeoclimatic
reconstruction. There are two methods generally
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past 361

FACTSHEET 20.5
Psychrosphere and Pleistocene Ice Age

Psychrosphere is the cool, deepest bottom layer of the modern ocean, where waters are nearly freezing. This
came into being in Late Eocene as cold, dense, polar water sank to the deep sea, as an effect of the Antarctic
icecap build-up. It then moved to lower latitudes, where on being heated it rose up to the surface. It is held that
the upwelling of psychrospheric water affected climate in many parts of the world. As found from isotopic
signatures (lighter O16 isotope is concentrated more in polar ice), the psychrosphere formed rather quickly in less
than 100,000 years during which time, bottom water temperature dropped perhaps by 4°C to 5°C.
One of the most profound change during Cenozoic is the Pleistocene glaciation. It, however, came about through
a series of events snow-balling into increasing cooling almost through the whole era (also see Factsheet 20.6).
The greenhouse warming effect of Cretaceous largely due to excessive carbon dioxide emanated with extensive
volcanic activity in different parts of the earth and continuing till Middle Eocene, was reduced, for reasons
unknown, in later parts of Eocene leading to Terminal Eocene cooling event. Besides, the only major exclusively
Cenozoic event of continental rifting, the separation of Australia from Antarctica, brought about alteration of
wind and water circulation near the South Pole, which, in turn, had effects on the whole of the globe itself.
Before separation, Antarctica centered over the South Pole remained relatively warm, because its shores were
bathed in relatively warm waters from lower latitudes. When Australia, broke away and drifted northward, a
cold current formed between the two continents deflecting the warm current that flowed towards Antarctica
from Australia. During Oligocene, a cold circumpolar current was established around Antarctica, adding to
further cooling. This resulted in the formation of sea-ice around Antarctica, sinking of near-freezing water to
deep sea and its spreading to the north, forming the psychrosphere.This climate deterioration in Late Eocene and
Oligocene fostered further deterioration later in Cenozoic time.
Psychrosphere in the northern hemisphere was also developed around Middle Palaeogene (late Eocene), when
mid-Atlantic rift proceeded northwards along two forked-arms, splitting Greenland from North America on the
west and Greenland from Eurasia in the east.

followed (Chaloner and Crebar 1990). In one,
climate is interpreted from plant physiognomy. A
limited number of leaf characters that show strong
correlation with present-day climatic parameters
are taken help of, such as leaf size (greater or
smaller than 10 cm length), thick cuticle, leathery
evergreen or thin cuticle, soft texture, an entire or
non-entire margin of leaf and the presence or
absence of drip-tip, and so on. In these characters
typical deciduous temperate leaves contrast with
those of lowland tropical evergreen rainforest trees.
However, this method is particularly de-
pendable only with angiosperm dominated leaf-
floras. The group appeared only in Upper Cretaceous
and, hence, the method is not very effective in older
ones.
Xeromorphic features point to aridity or
‘water-stress’ environments. However, certain
plant groups, such as confiers show broadly
xeromorphic features regardless of environment.
Moreover very thick cuticled pteridosperms of
Middle Jurassic in England (genus : Pachypteris)
was a plant adapted to salt-marshy condition, rather
than aridity.
Another plant feature used for palaeoclimatic
studies is the annual ring found in permineralized
or silicified wood fossils. They indicate seasonal
variations between humidity and aridity, the former
producing the growth ring and the latter putting
stops to them.
In another method called NLR or Nearest
Living Relative method, the principle of actualism
is utilized, i.e. climate is interpreted on the basis
of flora comparing the latter with any known
present-day flora. There is, however, problem in
this method as adaptation, particularly local
362 Part Four: Some Synthesis
adaptation of plants may have changed through
time. Any interpretation of the past climate on
calculations based on present flora may be fraught
with mistakes.
Coral reefs flourish in shallow tropical warm
normal seawater. Ancient coral reefs enclosed in
rock records are quite successfully used in
demarcating tropical belts of respective time.
However, present-day coral reefs show perfect
symbiosis between corals and photosynthetic
zooxanthellae, a dinoflagellate popularly known
as brown algae. This relationship cannot be verified
for the ancient counterparts as the preservation
potentiality of zooxanthellae is naturally low.
20.4.3 Indian examples
As an Indian example of palaeoclimatic
interpretation, we may cite the case of Gondwana
Group of formations. As discussed in Chapters 18
and 19, this huge thickness of continental clastic
sediments ranging in age from Permian to Lower
Cretaceous, is very rich in flora and fauna,
particularly in certain parts. Fauna includes large
or small terrestrial vertebrates, namely amphibians
and reptiles, with occasional fish recorded from
different units of the group. Major floral elements
are pteridosperms in lower parts and cycads
(gymnosperms) or related plants (cycadeoids) and
conifers in upper parts. In addition there are
Filicales (ferns) and Ginkgoales, etc.
Gondwana flora is classified into a lower
Gondwana or Glossopteris flora and on the upper
Ptilophyllum flora. Some authors prefer the third
middle Thinnfeldia-Dicroidium flora. Glossopteris
flora is dominated by broad leaf pteridosperms,
fossil wood with annular growth rings and is
generally associated with rich coal reserves or
carbonaceous shales. The flora is also associated
with tillites or glacial-periglacial deposits at the
beginning. It is of Permian age. This whole
combination indicates a moist or humid climate,
possibly of temperate to high tropical region,
prevailing at the initiation along with a Permian
glaciation.
The middle flora of Triassic time is dominated
by xerophytic forms indicating aridity.
Triassic formations of Panchet as also of the PG
(Pranhita-Godavari) Valley are, however
characterized by large reptiles like Lystrosaurus.
It could not have survived without large water
bodies. Abundant red sandstones and shales of
Triassic formations, too, indicate enough water to
carry iron cement into the sediments in their
intergranular pores and again enough aridity to
precipitate them. It is more likely that a seasonal
variation between humidity and aridity marks the
Triassic time.
Upper Gondwana (Jurassic–Lower Cretaceous)
containing Ptilophyllum flora, big land vertebrates
like dinosaurs (sauropod : Barapasaurus tagoreii),
occasional coal deposits indicate more humid
condition than in Middle Gondwana times. Cycads
and conifers, however, suggest not as humid a
climate as it was in Lower Gondwana.
There have also been attempts to correlate
palynological assemblages with climate, which
conforms with palaeoclimatic interpretations based
on larger plant fossils. Thus, the Talchir assemblage
is said to indicate extreme cold to cold, low to
medium humidity conditions; the Karharbari cold
to very cold, medium to high humidity; Barakar
cool to warm, high humidity; Kulti/Barren Measure
warm, medium humidity; Raniganj warm, very
high humidity; Panchet cool to warm, low to
medium humidity.
20.4.4 Some recent studies
More critical appreciation of recent data throws
additional interesting lights on the topic, as well
as calls for some revisions on this or that aspects.
For instance, distribution of carbonates in
general and reefs in particular was considered to
be controlled by temperature, and hence climate.
However, Ziegler et al., (1984) hold that the
limiting factor is the light penetration to the sea-
floor. Year-round light refraction drops markedly
at about 35 degrees from the equator, the present
pole-ward limit of the tropical coral-reef
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past
environments. This depositional system relies on
algal fixation of calcium carbonate, either directly
or indirectly as in the case of the symbiotic
zooxanthellae of corals, and hence is limited by
light penetration. However, within the tropical and
subtropical zones, light penetration may be reduced
by turbidity associated with clastic runoff from land
or resuspension due to energy variations and high
surface productivity.
Interpretation of climate from fossil wood is
also judged in an additional perspective (Creber
and Chaloner 1984). Fossil wood occurs widely
throughout the geological record from Upper
Devonian to the present. In Upper Devonian and
Carboniferous, wood had very weakly defined
growth rings consistent with growth in moist
tropical palaeolatitudes. In late Palaeozoic,
Gondwana wood from temperate palaeolatitudes
show marked growth rings.
In Mesozoic, very high palaeolatitude trees
show substantial growth increments. It may be due
to either of the following. Firstly, the possibility of
changes in the obliquity of the earth’s axis of rotation
may be the mechanism by which a very different
latitudinal pattern may have prevailed in the past.
The second alternative is the higher mean global
temperatures in Cretaceous permitting the growth
of abundant large trees in very high latitudes. As
global temperature declined in middle Tertiary, these
high latitude forests became increasingly restricted
to the latitudes where they exist now.
Tertiary palaeotemperature curves are drawn,
among others, on palaeobotanical evidence of
central European sequences. But later studies
revealed that the fossil groups on which these
curves were based appear to be the result of
taphonomic or depositional-preservational effects,
rather than real ecological associations. An
alternative grouping of palynological taxa is
obtained by Principal Component Analysis of large
quantities of high quality percentage frequency
data. The new stratigraphic distribution of these
groups helps create a new palaeotemperature curve
(Boulter 1984).
363
Though suggestive of diversities of
taxonomies and morphologies, palaeofloras cannot
provide details of the structure of the vegetational
habitat to provide additional details about climate.
Fossil ungulate communities are particularly
valuable in this regard. Recent studies have brought
out correlations between ungulate morphology and
foraging and reproductive strategies, which, in
turn, are highly dependent on structural details of
the vegetational habitat (Janis 1984).
An example is obtained from the study of
North American late Miocene fauna and flora.
They both come out to have formed a savanna
biome similar to that of the present-day Africa.
The ungulate faunas of the two (North America
and Africa) are similar in morphological types,
though the North American fauna is taxonomically
more diverse with lesser number of horned
ruminant artiodactyls and in the greater taxonomic
and morphological diversity of equid
perissodactyls. Distribution of postulated social
behaviours (territoriality as evidenced by the
possession of horns in artiodactyl males) and
foraging behaviours (fibre content of the diet as
evidenced by the degree of hypsodonty in equids)
suggest that the North American Miocene savanna
vegetation was of lower nutritional value and more
dispersed in its structure than in the African
savannas of the present. This could reflect a
difference in the overall amount and seasonal
distribution of rainfall.
An interesting scenario is suggested by
Brenchley (1984) in his studies on late Ordovician
extinctions of families, genera and species in
relation to the Gondwana glaciation (Hirnantian
in age). In the earliest phase of extinction, deep-
shelf benthic faunas of trilobite-cystoid-gastropods
are affected by falling sea level. Contemporaneous
ocean temperature changes may have affected the
plankton belts and caused extinction among
graptolites.
The last phase of extinction may have
occurred under rapidly rising sea level. Deep
flooding of continental shelves and restriction of
364 Part Four: Some Synthesis
shallow marine fauna to a narrow strip adjacent
to land may have contributed to the extinction of
some bentic faunas. A temperature rise of extra-
tropical waters with cooling of tropical waters
may have caused planktonic extinctions;
alternatively, a drop in salinity in the surface layer
of oceans from the influx of melt-water may have
been the cause.
Appearance and extinction of species and
higher taxa in Upper Palaeozoic may have been
controlled by climate (Dickins 1984). A lower
Carboniferous global equable climate changed to
a colder one in Upper Carboniferous. By the
beginning of Permian, a distinct ‘Gondwana biota’
developed consisting of the Eurydesma fauna
(marine invertebrate) and the Glossopteris flora
(terrestrial plant). During Upper Permian with
warming restored, cosmopolitan marine faunas
were re-established.
In a broader perspective, Scotese et al., (1999)
find from palaeogeographic-palaeoclimatic maps
drawn on lithological indicators of climate, such
as, coal, evaporites, bauxites and tillites, that major
climatic zones of humid tropics, dry subtropics,
warm and cool temperate and polar run across the
Gondwana supercontinent. Variations in location
and width of these climatic zones are now
considered to have been effected by both the
latitudinal movement of the Gondwana continents
and the changes in global climate between Ice
House and Hot House conditions.
Palaeoclimatic studies have also been
significant in ‘economic’ purposes too. Climate
is a major control on organic productivity. Thus,
from ancient climate patterns, the occurrence of
conditions necessary for the formation of type III
kerogen may be predicted. This kerogen is the
basic precursor to coal and certain petroleum
deposits. Potential petroleum reservoir rock
(siliciclastic or carbonate) can also be located
from an acurate understanding of palaeoclimate,
among others.
Peat forms in relatively wet intervals and
limestone during drier periods. Types of
palaeosoils (ancient soil) can also indicate wet and
dry cliamtic cycles, as the two produce different
kinds of soils.
20.4.5 Stable isotope studies in
palaeoclimatology
Stable isotope geochemistry now forms a rich and
promising branch of geological studies. A few
examples of such studies on Indian materials are
provided here, not as evidences of what has been
done, but as instances of what it is possible to do
with fossils, in this line. Hence, conclusions of the
studies cited here may be contested and must be
taken only as probable inferences.
Reconstruction of environment (Sarkar, Ray
and Bhattacharya, 1996) was attempted from
oxygen and carbon isotope analyses of bulk
samples of some foraminiferal Assilina-bearing
foraminiferal marls (CEM formation of Ray et al.,
1984, Naredi Formation of Biswas and Raju, 1973)
that enclosed a virtually monospecific assemblage
of Assilina developed in a highly stressed
environment. Oxygen results indicated that the
environment received fresh water in addition to
sea water itself, in some coastal lagoons. Highly
depleted 13C values were interpreted as pointing
to the cutting off of those restricted lagoons due to
fluctuations of sea level, trapping of a large amount
of organic matter in that water and subsequent
decaying of them giving off biogenic methane.
Decomposition of this methane by sulphate
reducing bacteria produced the observed carbon-
isotope composition of the bioclastic marls.
Authors also concluded that the area was part of a
global warm humid belt with rich vegetation
(attested also by lower to middle Eocene coal-
lignite-carbonaceous shale occurrences in different
parts of India), abundant fresh water run off and
high carbonate productivity in sea.
Oxygen isotope analysis of samples of
foraminiferal limestones from two different
Eocene-Oligocene Boundary (EOB) sections in
Kachchh and correlation of the results with DSDP
(Deep Sea Drilling Project) sites, revealed a rapid
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past
cooling of ~6oC marking the Terminal Eocene
cooling event (Sarangi, et al., 1998). Effects of this
same event are also obtained from oxygen isotopic
analyses of larger foraminifers from a few onshore
sections in western India (Saraswati, Ramesh and
Navada 1993). Results of those latter analyses
reveal that late Palaeocene-Early Eocene surface
water temperature was 32°C, dropping sharply to
26°C in the late Middle Eocene and further to a
mean temperature of 22°C in the Early Oligocene.
High resolution oxygen and carbon isotope
analyses on marine fossils may cast light on
palaeomonsoonal patterns too. An example may
be obtained from Sarkar et al., (2000) who studied
planktonic foraminifera of five cores from the
northern Indian Ocean. Higher 18O amplitude of
Arabian sea cores indicated that the sea was in
general 1 per cent to 2 per cent more saline during
the last glacial maximum (LGM, 18ka) than at
present. 13C and CaCO3 calculations show reduced
upwelling and productivity (in Arabian Sea
productivity decreased by ~ 65 per cent during
LGM relative to Holocene). This might have been
due to weaker summer monsoon during that period
probably caused by excess evaporation over the
northern Arabian Sea and increased oceanic
precipitation in the equatorial Indian Ocean.
There are two hypotheses regarding the cause
of density band formation on the surface of coral. In
one, it is concluded that the sea surface temperature
(SST) controls the density band formation in corals,
while the other attributes the seasonally varying
growth rate to be the cause of the annual banding. It
is also noted that the same species of corals form
high or low density bands in summer/winter in
different geographical locations. Availability of
sunlight and nutrients, rather than SST, may control
the growth rate of bands, a conclusion also
corroborated by stable isotope studies.
Stable carbon and oxygen isotope analyses of
planktonic foraminifera (Globigerinoides ruber)
from a deep sea sediment core in the Andaman Sea
show 18O amplitude of 2.1 per cent which suggests
increased salinity and/or decreased temperature in
the glacial surface waters of that region. A pulse of
365
18O enrichment during the last deglaciation is
attributed to a Younger Dryas (a near glacial cooling
event that occurred between 11,000 and 10,000 year
B.P. and which separated the two steps of deglacial
warming, cooling and/or to a sudden decrease of
fresh water influx from the Irrawady and Salwean
rivers into the Andaman sea.
These studies also help in palaeoecology. For
instance, Sarkar Bhattacharya and Mohabey (1991)
worked on well-preserved clutches of dinosaur
eggshells and skeletal remains in the Upper
Cretaceous Lameta limestones of the Kheda
district, Gujarat. Isotopic low values of 18O (~ –
8.0 per cent) and 13C (~ – 8.5 per cent) of those
beds were indicative of fluvial or marshy lacustrine
environment of deposition. The 13C values of
dinosaur eggshells from those beds were about
–10 per cent and suggested that the reptiles were
surviving on plants of C 3 type (small palms,
confiers, dicot shrubs, etc.). The eggshell 18O
values varied widely from – 8.0 per cent to + 6.9
per cent. This range suggested that the dinosaurs
used to drink water from a variety of freshwater
sources—from rivers originating from high altitude
precipitation to small, confined evaporative pools
and indicated that the prevailing climate was of
semiarid type. Abundant higher plants grew around
those freshwater bodies, providing an ideal
ecological niche for the dinosaurs.
Stable isotope studies of foraminifers have also
been used to know how conditions at the ocean
surface or floor or in the thermocline vary with
time. This may lead to develop understanding the
changes in thermocline, in temperature gradient,
oxygen minimum layer, organic productivity,
oceanic circulation changes, seaway developments,
etc. during Cenozoic in Indian ocean.
D 13C and D 18O values of recent shallow water
low (Thecideida), middle (Terebratulida, low-
midium) and high latitude brachiopods have been
determined. It is concluded that such values for
corresponding fossil groups when compared with
the values of their recent counterparts, may indicate
the nature of diagenesis, as well as important
oceanographic or chemostratigraphic proxies.
366 Part Four: Some Synthesis

Factsheet 20.6 summarizes the major factors
and effects in regard to the distribution of animals
and plants in Neogene, which can only be arrived
at when we know and synthesize palaeogeographic,
palaeobiogeographic and palaeoclimatic details,
acquired from different studies.
A similar conclusion is reached by Erwin
(1999), where he concludes in respect of biospheric
perturbations that variations in biotic diversity
ranging from global evolutionary radiations and
mass extinctions to regional fluctuations or changes
in local community assemblages reflect vicissitudes
of environmental circumstance. This, he concludes,
is evident even in the history of marine life
structured by Neoproterozoic-Early Cambrian
radiation, Ordovician radiation, diversity plateau
or equilibrium from Late Ordovician to Permian,
punctuated by end-Ordovician, Late Devonian and
end-Permian mass extinctions. A synthesis of
palaeogeographic palaeo-biogeographic palaeo-
climatic data coupled with other geological and
geochemical evidences can lead to the development
of testable models of the scenario.
Another interesting example of close
interrelation and interaction of these data is
provided in Factsheet 20.7

FACTSHEET 20.6
Summary of Events Leading to Pleistocene Ice Age

Ice age begins

1.6 m.yrs. Ice sheets form in northern hemisphere
Gulf Stream intensified
5.3 m.yrs. .......
Isthmus of Panama closes
6.0 m.yrs. Rapid cooling takes place
Present abyssal ocean circulation established
Iceland-Faroe Island Ridge and Atlantic water wells up around
15 m.yrs. Antarctica; more moisture and copious snow-pack.
Antarctic ice cap develops
30-25 m.yrs. Antarctica much colder but still no ice cap
Circum-antarctic current established. Barriers in the Western Pacific and closing of Tethys sea in
the Middle East and Near East breaks up circum-equatorial circulation.
35-30 m.yrs. Partial circulation around Antarctica; cold current between already separated Antarctica and
Australia deflected to the south, pushing back the eastern warm current eastward.
First glaciers on Antarctica
38 m.yrs. Deep water circulation speeds up
Rapid cooling of surface water in the south and of deep water everywhere
Terminal Eocene Cooling Event
<50 m.yrs. Slight cooling takes place
No glacier or icecap on Antarctica
> 50 m.yrs. Deep water much warmer than at present
Uniform climate and a warm ocean with abundant carbonate
Free flowing circum-equatorial circulation of warm water
 Chapter 20 Fossils and Climate, Geography, Biogeography, and Ecology of the Past 367

FACTSHEET 20.7
Summary of Palaeobiogeographic and Related Changes During Neogene

Factors controlling distribution

1. Continental collisions allowing terrestrial faunal exchange
(a) Africa-Eurasia collision
(b) India-Eurasia collision
(c) Australia-Eurasia collision
2. Formation of the Panama Isthmus allowing faunal exchange between the two Americas.
3. Change in seaways favour or hinder exchange
(a) Development of marine connections, e.g. between Australia and Antarctica.
(b) Sealing off of marine connections, e.g. Mediterranean Sea, Panama Isthmus, etc.
4. Evolution of new migratory route, for example with closure of the Mediterranean-Indo-Pacific seaway
5. Development of new precipitation zones (such as in Peninsular India due to the Himalaya),
6. Climatic deterioration and concomittant aridity and/ or cooling
Effects brought about on the organic world
1. Less pronounced in the marine realm, though even there sea-mammals (whales, dolphins, etc.) radiated
rapidly.
2. Cooler and drier climate helped diversification and spread of grasses and herbs on land.
3. Development of a chequered environment particularly at the margins of forests
4. Terrestrial mammals assumed their modern character
5. Modern reptiles, amphibians and birds underwent diversification.
6. Mammals radiated in different niches
7. Apes radiated during Miocene though dwindling thereafter, mainly in forests.
8. Marginal environment between forests and grassland, Australopithecus, the oldest known hominid genus
evolved about 4-5 million years ago
9. The modern species of man, Homo sapiens sapiens, took the scene in Middle Pleistocene onwards.
 Part five
Appendices
Appendix

1
Fossil Lagerstätten

Fossil Lagerstätten through the geological column
(Factsheet 2.11) have been introduced in Chapter 2.
It has also been indicated there that knowledge of
such major Fossil Lagerstätten, may cast new lights
on the history of the organic world. Some summary
information have been provided in the Factsheets
2.12–2.14, while Factsheet 2.10 show their types.
The present chapter, to be treated as an
appendix to that discussion, gives more details of
the 14 Fossil Lagerstätten from Precambrian to
Pleistocene. No further discussion will be added,
since the information are largely self-explanatory.
It should only be added that these exceptional
occurrences of fossils, not just provide examples
of what were the constituents of the organic world
in the past, but also bring out how the ecosystem
evolved, how different kinds of living and feeding
habits emerged during the geological past and,
prehaps more interestingly, how factors and
processes of the inorganic world, including
sedimentational aspects of them, interacted with the
organic world, to produce different kinds of
exceptionally well-preserved, in quality or quantity
or both, fossil records. These remain some of latest
avenues along which palaeontology is advancing
and may move further.

FACTSHEET A1.1
Ediacara Biota

Late Precambrian (Upper Proterozoic); from Ediacara Hills near Adelaide in West Australia
Marine rocks: shallow water (Glaessener 1961), but later suggested deeper water (Gehling 1991); deposit
mostly autochthonous, at a few localities allochthonous and drifted into the site of deposition.
Biota mainly of benthos, a few may be pelagic, some may be epibionts; detritivore, suspensivore, even
planktivore. Macrophagous predators typically absent.
Ediacara biota represents biological organization, different in grade from that of the Phanerozoic times. It,
thus, represents the organisms that lived on this earth before mineralized hard parts evolved and predation as
feeding habit (a typical advanced heterotrophic feeding habit) appeared or at least became widespread.
Some typical examples
Cyclomedusa commonest and most widespread; affinity uncertain, may be a holdfast of a colonial organism,
for example, a cnidaria. Ediacaria, concentric discoid; affinity uncertain. Arkarua, probably the earliest known
(Cont...)
371
372 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.1 (Cont...)

Ediacara Biota

echinoderm. Different others are interpreted as floating medusoid, or vagrant or sessile benthic, or holdfast of
other organisms, or segmented worm-like forms.
Seilacher removed the Ediacaran organisms from the Metazoa and placed them in the Vendozoa, mainly on
constructional morphology. They are generally held as part of Animalia kingdom at an early phase of development
of metazoan body plans.
Even the primitive organic world as represented by the Ediacara biota was not singularly portrayed at the
Ediacara Hills only, though not always in the same sedimentologic, and may be, the same taphonomic mode.
At Ediacara, Australia: fossils in shale partings in sandstone beds in quiet water.
At Ediacara, Newfoundland, Canada: fossils at the base of volcanic ash beds.
At Ediacara, Namibia, Africa: fossils in sandstones and shales formed in agitated water.
Also found in Russia, Ukraine, Norway, UK, China, USA and Mexico.
Doubtfully from Iran, Morocco, Ireland, India, etc.
Fate of Edicara biota
Generally said, the biota included primitive annelids, arthropods and cnidarians unlikely to be ancestors of
later forms.
A few may have continued as group (e.g. jelly fish of scyphozoa); mostly extinct.
The biota is succeeded by the Tommotian (the earliest Cambrian) SSF (Small Shelly Fauna) though the relation
between the two is debated. The Cambrian explosion of shelled organisms took place next. Burgess Shale
fauna stands as the representative of the latter.

FACTSHEET A1.2
Burgess Shale

Middle Cambrian: from British Columbia, Canada.

Marine rocks; Mainly benthos living in, on or just above the muddy seabed at the foot of submarine cliff, in a
tropical zone, within 100 metre depth (photosynthesizing algae); both epibionts (sessile c.30 per cent, vagrant
c. 40 per cent) and endobionts (c.10 per cent), a few nektons; sessile suspensivores/filter feeders (sponges),
vagrant detritivores (arthropoda deposit collectors and molluss swallowers), scavengers (lobopods) and predators
(e.g. giant Anomalocaris), etc.
Biota indicated that nearly all animal phyla had developed; more or less complete contemporary ecological
dynamics, i.e. diverse interacting relationships of organisms of the sea floor (with predators, scavengers,
filterers, collectors, swallowers) are represented.
Biota included:
Hallucigenia, an onychophora, a lobopodian marine velvet worm.
Marrella, an arthropoda, > 15,000 specimens, unique in its occurrence.
Anomalocaris, an arthropoda/problematica, a monster predator.
Opabinia, a problematica, with five eyes and a proboscis.
Plus sponges, pennatulacean cnidaria, different kinds of worms including polychaetes and others of uncertain
affinity, etc.
(Cont...)
 Appendix 1: Fossil Lagerstätten 373

FACTSHEET A1.2 (Cont...)

Burgess Shale

Oldest chordate Pikaia from the Middle Cambrian Burgess Shale indicating that chordates appeared during
the Cambrian Explosion; small jawless agnathan fish discovered (1999) from Lower Cambrian of southern
China pushes the age further downwards.
The two most important ones among 40 equivalent occurrences of the Burgess Shale are Sirius Passet of northern
Greenland (3000 specimens, of soft-bodied Lower Cambrian fauna, slightly older than the Burgess Shale) and
Chengjiang of southern China (with many Burgess Shale organisms and many other newer ones in a deposit
slightly older than the Burgess Shale, Lower Cambrian; most important is an agnathan fish.
Recently reported from coeval successions of earliest Cambrian of China, Siberia and more recently from North
America, three-dimensionally phosphatized, spherical fossils, interpreted as metazoan eggs and embryos on the
basis of taphonomic features and cleavage patterns, suggest a wide geographic distribution of these taxa. Local
environmental and taphonomic conditions might have been ultimately crucial in preserving this phosphatic
window into the record of early animal evolution (Pyle, Narbonne, Nowlan, Xiao and James, NP 2006)

FACTSHEET A1.3
Soom Shale

Ordovician: from South Africa

Marine shallow cold quiet water not far from ice front on a muddy mainly lifeless bottom; Soom Shale
gradationally overlies tillites.
Nektons dominant; chitnozoans that may represent egg-masses of orthocone cephalopods, brachiopods, trilobites,
eurypterids, orthocone cephalopods and particularly, exceptionally preserved conodont animals with a report
of cartilage supporting large eyes suggesting chordate nature of conodonts.
Eurypterids, conodonts and orthocones among the nektons were presumably predators and/or scavengers,
brachiopods, cornulitids filter-feeder, naraoiid trilobites akin to Burgess Shale forms scavenger or deposit-
feeders, occasional lingulate brachiopods and ostracodes among benthics; coprolites, crushed brachiopod
shells, broken conodonts suggest the presence of large predator, yet undiscovered.
A unique character in preservation in Soom Shale is direct replacement of organic materials by clay minerals,
latter adsorped before replacement.

FACTSHEET A1.4
Hot Spring Miracle in Rhynie Chert

Devonian of Aberdeenshire, Scotland

Early terrestrial biota preserved in silicified mode.
Diverse plants, including seven true land plants, five of which are tracheophytes, true vascular plants with
both sporophytes (asexual generation) and gametophytes (sexual); resemble living primitive plant Psilotum.
Water-vascular tissues, fossil spores in the process of germination preserved.
Diverse land animals, centipedes, arthropods (including oldest mites), arachnids, springtails, myriapedes ;
carnivores preponderant, some detritivores (which ingest plants partly decomposed by bacteria, as in soil
ecosystem) and no herbivores (which use symbiotic bacteria and fungi to help them digest plant tissues that
appear later in Carboniferous). Lungs in arachnids preserved.
374 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.5
Hunsrückschiefer/ Hunsrück Slate

Devonian; from Koblenz, Germany

Biota include diverse fish (four of the five main groups), mostly agnathans and placoderms and the earliest
lung-fish (sarcopterygian); starfish and ophiuroids among echinoderms; soft bodied polychaete worms,
intermediate between Burgess Shale and Mazon Creek fauna; arthropods (trilobites, crustaceans and chelicerates)
with internal organs preserved; siliceous sponges, cnidarians (rugose, tabulate corals, scyphozoans, ctenophores,
conulariids); molluscs including gastropods, bivalves, orthoconic nautiloids and goniatites among cephalopods;
brachiopods and bryozoans; calcareous algae; fragments of terrestrial vascular plants; fish coprolites, epifaunal
tracks, infaunal traces including burrows.
Occurrence in muddy (marine) sediments affected by low-grade metamorphism.
Though in part nektonic, the biota comprised mainly a benthonic community, with photosynthetic red algae
and well-developed eyes of some fish and arthropods suggesting a photic zone community and with diverse
trace fossils suggesting thriving infauna living in oxygenated bottom waters.
At the same time, preservation of soft tissues suggests the absence of scavengers, indicating environment
turned rapidly anoxic and became inhospitable.
This biota was buried by rapid sedimentation in episodes or phases caused by turbidity currents which, in turn,
was produced by tropical storms.
Remarkably both mineralized and soft parts are pyritized, though in a few restricted taxa (only in limbs, eyes
and intestines of arthropods and tentacles of cephalopods) and in a few thin beds of short duration within a
thick sequence.
Rare instance of pyritization of soft tissues: factors and processes
Low organic content plus high concentration of dissolved iron.
Sulphate-reducing anaerobic bacteria produce sulphide from organic matter.
Iron turns it to iron monosulphide and aerobic bacteria oxidize it to pyrite.
Low organic content help iron precipitate in the carcass and not in the sediment.
Initial reduction, then oxidation taking place near the anaerobic-aerobic interface in upper layers of sediments.

FACTSHEETA1.6
The Mazon Creek Biota

Late Paleozoic: Mazon Creek, a tributary of the Illinois River, USA.

Two sub-biotas (downwards in decreasing abundance)
Nearly complete records of late Palaeozoic shallow marine, freshwater and terrestrial life with 300 species of
animals representing 11 phyla and 200 species of plants.
Terrestrial Marine
and freshwater
83% plants 7.8% coprolites 42% jellyfishes like Essexela, etc. 29% plants (drifted)
Freshwater bivalves Shrimps Burrows and trails Marine bivalves
Other molluscs Millipedes Coprolites Other molluscs
Horseshoe crabs Fish scales Worms Marine shrimps
(Cont...)
 Appendix 1: Fossil Lagerstätten 375

FACTSHEET A1.6 (Cont...)

The Mazon Creek Biota

Arachnids Insects Crustaceans ‘Tully Monster’

Fish Centipedes Fish Insects
Millipedes, centipedes, horseshoe crabs, arachnids,
Amphibians
Together they made a variety of habitats, viz. a swamp forest with tree-sized club-mosses and horsetails and
undergrowths (the Colchester Coal); upland setting, with ferns, seed-ferns and horsetail debris (Francis Creek
flora); terrestrial fauna (arachnids, insects, etc.) lived among these plants; a delta with freshwater, brackish and
restricted marine environments associated with it, where reduced salinity and muddy water prevented typical
marine animals such as brachiopods, corals, crinoids.
Mazon Creek fossils occur in siderite concretions
Fossils occur in siderite concretions in three-dimensional specimens slightly flattened at rims; as external mould
with a carbon film in plants, crystals of pyrite, etc. at mould surfaces of which the commonest mineral is kaolinite.
Concretions (in which fossils are found) formed very soon after the death and burial of the organisms, as is
evident from : fossils showing (a) little decay, (b) failed bivalve escape structure (bivalves preserved on the edge
of a nodule at the end of its death trail); (c) Lingula, an infaunal brachiopod buried in life position, (d) seed-fern,
pinnules at right angles to bedding.
Concretions formed before any appreciable compaction as indicated by organisms preserved three-dimensionally,
at least in the cores of the nodules, while the matrix around is highly compressed.
Nodules contain 80 per cent siderite cement implying 80 per cent of water by volume in the sediment before
compaction.
Siderite forms in the following course: in the presence of decaying organic matter sulphate-reducing anaerobic
bacteria release sulphur; iron normally reacts with sulphur to form pyrite; once sulphate is used up, methanogenic
bacteria help produce siderite.
Mazon Creek condition must have had, thus, abundant iron, meagre sulphate.

FACTSHEET A1.7
Rapid Sedimentation and Tidal Cycle :
Mazon Creek a Case to Show Why Palaeontologist Needs to Know Sediments Well

Mazon Creek sediments show paired clay-silt laminae, pairs widen or narrow in a cyclical fashion
A single tidal cycle has two thin clay bands, one wide silt band (widest corresponding to spring tides) and one
narrow (narrowest to neap tides)
Thin clay bands represent stillstands (i.e. flood slack/ebb slack; water is in the process of turning; no flow).
Thicker silt bands during ebb tides (outgoing tide allowed a high water flow in the basin, i.e. the creek, from
land and so more sediments)
Thinner silt bands during flood tides (incoming tide resisting inflow of water in the basin).
In Mazon Creek case 15-16 tides in a cycle from springs to next springs, which is equal to half lunar month
Complete lunar cycle has two springs and two neaps
Coral growth ring cyclicity sug gest 30 days in Carboniferous in a lunar month.
So tidal cycle at Mazon Creek is of diurnal type.
(Cont...)
376 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.7 (Cont...)

Rapid Sedimentation and Tidal Cycle :
Mazon Creek a Case to Show Why Palaeontologist Needs to Know Sediments Well

Rate of sedimentation
Each 15 day cycle measures 19-85 mm.; deposition rate 0.5-2.0 m. per year of compacted sediments; so the
entire shale sequence deposited in 10 to 50 years.
So a rapid sedimentation is concluded from the evidences of:
(i) Fossils (little decay, failed bivalve escape structure, Lingula buried in life position; seed-fern, pinnules at
right angles to bedding);
(ii) Sediments (formation of concretions very soon after the death and burial of the organisms, preservation of
organisms in three dimensions, at least in the cores of the nodules, with matrix around highly compressed,
suggesting that the concretions formed before any appreciable compaction); and
(iii) The measure of tidal cycle.

FACTSHEET A1.8
Grés à voltzia: A Lagerstätten after the End-Permian Severest Mass Extinction

Triassic: France
Semi-arid terrestrial and brackish water, low-diversity communities.
Biota includes land plants, medusoid cnidarian, brachiopods, annelids, molluscs (marine), arthropods (most
abundant) such as horseshoe crabs, crustaceans, myriapods, insects including their larvae, fish, amphibians
and rare reptiles.
Biota attests Permo-Trias transition across this level that involved:
1. disappearance on land of Carboniferous lush green tropical forests of lycopods and pteridosperms with
amphibians, insects and arachnids as faunal associates ;
2. extinction of trilobites, euryptoids/eurypteroids and graptolites in seas;
3. replacement of brachiopod shelly fauna by that of bivalves, and
4. appearance of new types of corals in seas, gymnosperms and reptiles on land.
Biota preserved in three facies have:
(i) thick lenses of fine-grained sandstones, point bars in sinuous channels and containing land-plant debris
and amphibian bone fragments;
(ii) green/red silt/clay laminite lenses formed from finer materials settling in brackish ponds with beautifully
preserved aquatic and terrestrial biota, and
(iii) beds of calcareous sandstone, products of storm controlled marine incursions with meagre marine fauna
to be overlain by marine transgressive rocks.
Biota along with sediments suggest deltaic environment: sandstones represent point bars in sinuous channels;
clay deposited in brackish ponds; calc-sandstones suggest storm-induced incursion of sea water; red beds and
xerophyte plants suggest semi-aridity; low-lying deltaic environment not too arid, probably seasonal (desiccation
cracks, reptile footprints, salt pseudomorphs and land-plant in life position at the top of each clay lens suggest
complete drying out of pools filled during wet season).
Biota passes through aquatic to terrestrial vertical transition within each clay lens.
(Cont...)
 Appendix 1: Fossil Lagerstätten 377

FACTSHEET A1.8 (Cont...)

Grés à voltzia: A Lagerstätten after the End-Permian Severest Mass Extinction

Unique feature of production of microbial mat and phosphatization:

Drying up of pools evident from desiccation cracks, reptile footprints, salt pseudomorphs and plants in life
position:
Death of aquatic fauna from drying up of pools indicated by abundance of estherids adopted to completing life
cycle in temporary water bodies.
Regular high evaporation rates lead to deoxygenation, mass mortality of aquatic fauna and rapid proliferation
of microbial films.
Microbial films formed on and above carcasses, thereby:
1. prevent scavengers’ activities, and
2. produce low O2 environment with abundant organic matter in which decomposition is largely inhibited and
phosphates released from organic matter are confined and shut off from reuse by other organisms; acidic
condition from decay, release free Ca+ ions to form apatite.
Microbial film-covered carcasses are buried: some squashed by compaction; those phosphatized are prevented
from flattening.

FACTSHEET A1.9
Holzmaden Shale (Posidonienschiefer)

Exemplifies that community and fossil assemblage may be different. Community of an epicontinental
marine basin of depth of 100-600 m, subtropical ~ 30° N.
Lower Jurassic ; Baden-Württemberg, South Germany
Constituents: Biota includes abundant and sometimes completely preserved marine reptiles and fish, some
pterosaurs (flying reptiles) and dinosaurs and marine invertebrates dominated by coleoid cephalopods (squids
and belemnoids).
Host units: Marine black bituminous shale with a number of units; shale-oil bearing unit has best preserved
fossils; bituminous shale contains upto 15 per cent organic matter, causes prolonged combustion and shale-oil
as by product.
Quality of preservation is evident from the presence of ink sacs and tentacles in belemnoids as also skin and
muscles in ichthyosaurs preserved; limestone above shale-oil unit has uncompressed fish; crocodiles come
from another.
Basin and stratinomic condition: The area belonged to epicontinental South German basin, one of the several
such between submarine highs and islands of that time; diffused pyrites and organic matter suggest that the
basin was oxygen starved, H2S rich, stagnant; thin widespread laminae suggest still water; rare benthics and
bioturbation except a few echinoids, crustaceans and burrowing bivalves suggest that bottom conditions were
hostile to life; microbials falling to the seabed could not completely decay carcasses for want of oxygen, to be
buried in anoxic mud devoid of scavengers; similar to present-day Black Sea. Stagnation conditions were
occasionally interrupted by storm-events having brief oxygenating effects; algal mat just above the sediment-
water interface would have inhibited grazers, scavengers, burrowers.

(Cont...)
378 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.9 (Cont...)

Holzmaden Shale (Posidonienschiefer)

Community Trophic chain included

Marine: Planktonic and nektonic in well-aerated Primary consumers: Filter feeders (crinoids/bivalves)
surface waters: pseudoplanktonic (crinoids, bivalves and few deposit feeders (gastropods/echinoids/
and inarticulate brachiopods attached to logs/living ophiuroids)
ammonites), nektoplanktonic (bivalve Bositra Primary predators: Bony fish and cephalopods.
(cf. Posidonia); and true nektons (ammonites and Secondary predators: Ichthyosaurs, crocodiles,
smaller fish, squids and belemnoids among sharks, plesiosaurs, the larger of the latter at the top.
invertebrates; large fish and marine reptiles, viz.
crocodiles near shore, ichthyosaurs and giant
plesiosaurs in open sea); benthonics rare, bioturbation
too (grazers, burrowers: a few echinoids, crustaceans
and burrowing bivalves).
Fossil Assemblage
Marine community members
and
Terrestrial allochthonous remains (gymnosperms represented by logs or dinosaur remains) washed into the
basin as disarticulated remains, particularly during storms and pterosaurs that predated upon fish and when
overrun by storms, drowned and were preserved articulated.

FACTSHEET A1.10
Morrison Formation

With a long story to tell on palaeoecology and taphonomy

Upper Jurassic, Western USA
Concentration Lagerstätten of late terrestrial reptiles of Mesozoic and early mammals, rare in occurrence,
yet in which dinosaur bones are so common, as to have been used as building materials, at least few years back
; also found in China and Tanzania.
Constituents: Dinosaurs both saurischian (carnivorous theropods to herbivorous sauropods) and ornithischian,
plus traces (ornithopod egg shells, coprolites, trackways), other rare reptiles (lizards, crocodiles, turtles,
lizard-like sphenodonts), pterosaurs (including pterodactyloids); mammals (rodent-like symmetrodonts,
multituberculates, etc.); fish (sacropterygian-lung fish, actinopterygian (ray-finned), primitive teleostans, etc.);
freshwater molluscs, ostracods, etc.; plants.
Host environment: Many fossil-rich beds are poorly sorted fluvio-lacustrine sediments, probably formed
by flash floods over a large area. Environment varied and included wet swamps (with coal deposits) in the
north, desertic in south; fluvio-lacustrine in mid-west (Colorado, Utah, Wyoming).
Food chain: Vegetation—small and large herbivores — carnivores (packhunters killing large sauropods).
Palaeoecology and taphonomy:
Scenario starts with retreat of the mid-Jurassic Sundance Sea; so Morrison Formation overlies mid-Jurassic
marine rocks.
(Cont...)
 Appendix 1: Fossil Lagerstätten 379

FACTSHEET A1.10 (Cont...)

Morrison Formation

This leaves wide, open plains traversed by meandering rivers and studded with lakes.
There was also mountainous water-divide to give rise to rain-shadow areas, which in turn caused aridity-
semi-aridity with seasonal rainfall.
Flora (horsetails, ferns, cycads, ginkgos, gymnosperms) suggests short period of a more humid, tropical climate.
The region became the home of herds of herbivorous dinosaurs to live on vegetation; smaller ones (e.g.
stegosaurs) lived on lowland ferns, horsetails, etc.; large long-necked sauropods on conifers, ginkgos, etc.
Carnivores followed roaming herbivores; packhunter carnivores killed large sauropods.
Freshwater invertebrates, frogs, lizards, etc. turtles, crocodiles and fish completed the fauna of perennial
streams and lakes in the plains with flying pterosaurs around them.
Mammals in caves and trees, probably nocturnal to avoid large predators.
Cyclic drought, imprinted on aridity-semi-aridity (as in Kenya today), concentrated animals around remnant
water bodies where they died of dehydration.
Periodic flash floods swept the disarticulated bones to be buried in channel fill of streams.
Mass mortality was of non-catastrophic type which took place over a long time span of starvation helping
characteristically more disarticulation and where age, health, gender and social (herd) ranking of individual
mattered, juveniles, females and senile adults dominating in number in the fossil assemblage.
Good preservation is favoured by aridity.

FACTSHEET A1.11
Solnhofen Limestone

Uppermost Jurassic: Bavaria in Southern Germany

What makes Solnhofen Limestone so important?
1. Conservation Fossil-Lagerstätten of allochthonous biota buried and protected, and thus excellently preserved
through a close interaction of (i)life-inhibiting basin condition, (ii)sedimentation (slow, stagnant water
deposition), (iii)sediments (impervious calcareous sediments), (iv)biostratinomic (burial)-taphonomic (diagenetic)
conditions
2. One of the most well-known Lagerstätten, particularly for Archaeopteryx, the earliest known bird record.
3. Exquisite preservation of soft tissues in a finely laminated, micritic, platy (plattenkalk) lithographic
(the rock was used for lithography) limestone occurring in laterally continuous beds in relatively restricted
basins; these micrites also preserve tracks and trails.
4. Excellently preserved rich biota of 600 species, mostly allochthonous, include delicate remains of (a) Vascular
and non-vascular plants; (b) A whole range of marine invertebrates (including squids with tentacles and
dragonflies showing fragile wings); (c) Fish and marine reptiles, rare dinosaurs, flying reptiles complete
with wing membranes; and (d) The only example of Archaeopteryx, the earliest known birds with its feathers.
I. Depositional setting
This formation was a deposit of inhospitable restricted lagoon within the then shelf sea of the region; basins
were generally marine; there was landmass to the north and coral reefs to the south and east, separating the
lagoon from the Tethys sea.

(Cont...)
380 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.11 (Cont...)

Solnhofen Limestone

II. The community was stratified by water conditions

Restricted basins had stagnant waters with salinity density stratifications, under semi-aridity that
could have caused much evaporation. Stratification from top towards bottom and corresponding
biota type were:
(a) Surface waters aerated and with normal salinity sustained planktonics and nektonics (as supported by
their coprolite remains).
(b) Sponge-microbial mounds high enough to reach oxygenated surface waters provided place for some
benthonics and nektonics to survive there.
(c) Hypersaline heavy bottom waters, also anoxic and poisoned by algal blooms, generally hostile for life
and inhibited benthonic, including scavenging and burrowing organisms; only salt-tolerant
cyanobacteria grew and lived on the bottom.
III. Periodic ‘catastrophic’ events determined burial and sedimentational aspects
Stratification was disturbed during violent periodic monsoonal storms; that periodically mixed up surface
and bottom waters causing death of surface-dwellers, as also marine organisms of the open sea and reef
community, from sudden fluctuation of salinity and oxygen; so fossil record suggests mass mortality and
sudden death (e.g. fish killed while eating or fish remains in pterosaur stomach).
Storms also caught flying pterosaurs and Archaeopteryx in strong winds and drowned them; flying insects
and plant fragments were blown across the lagoon and sank. Fish remains in pterosaur stomach shows why
the latter flow over the lagoon. Some pterosaurs had teeth suggesting their insectivore habit. Lesser number
of reptilian fossils, of lizards or dinosaurs, than number of flying forms was because of the fact that latter
could reach lagoons, former did not or could not.
Storms also stirred up fine carbonate particles (micritic ooze) from around the reefs. These sediments were
washed into the lagoon to rapidly bury the corpses on the floor. This produced the host lithofacies of finely
laminated, micritic, platy (plattenkalk) lithographic (the rock was used for lithography) limestone occurring
in laterally continuous beds in relatively restricted basins.
An alternative model suggests micrite to have been produced by cyanobacteria of the lagoon.
However, produced this micritic limestone, fine-grained and cemented early, presented an impervious
burial or preservational medium.
IV. Good preservation came from
Stagnant, anoxic, hypersaline waters retarded microbial decay of dead on the bottom. Some animals
(e.g. horseshoe crabs and crustaceans preserved at the ends of their trails) survived for short periods and
then died.
Rapid burial in fine grained micrite (impervious, early cemented) preserved soft tissues (wings of insects,
feathers of birds and tentacles of squids) or organic material (ink-sacs of cephalopods, original feather of
Archaeopteryx), occasionally replaced by calcium phosphate (muscles of fish and cephalopods).
Cyanobacterial material (leaving hollow spheres of coccoid cyanobacteria in the sediment) may have helped
in good preservation by encapsulating and protecting corpses and by binding the micrite together preserving
tracks and trails.
Impervious micritic limestone, fine-grained and cemented early, presented a host that inhibited diagenetic
destruction.
 Appendix 1: Fossil Lagerstätten 381

FACTSHEET A1.12
Santana and Crato Formations

Introduction
Cretaceous N-E Brazil
A vertically paired record of two limestone formations from Brazil, Santana (younger) and Crato Formations
separated by a third evaporite in between, assumes importance in different respects. They:
1. add to our knowledge of Cretaceous innovative evolution that ushered in life of modern affinity;
2. provide fossils with soft parts preserved in two modes, in one (Santana Formation) inside limestone
concretions, in the other (Crato Formation) on beddings of platy limestones; and
3. compare with other Fossil Lagerstätten (Santana Formation with Mazon Creek in nodular preservation,
Crato with Solenhofen Limestone in platy, micritic limestone host) though with distinct differences.
Biota in the background of Cretaceous innovation
Constituent biota inthe background of Cretaceous innovative evolution:
During Cretaceous, Pangaea was already broken-up. In result, many land bridges were lost; the Atlantic ocean
was opened and the Tethys extended westward. These disturbed migratory routes of land fauna and dispersal
routes for spores-pollens of land-plants. Intense volcanism during this period in different parts of the world
(e.g. Deccan Trap in India) caused toxic emanations including carbon dioxide. Increase of this gas in the
atmosphere brought the greenhouse effect, a major cause to produce the warmest climate in history. All this
must have provided the stage for significant evolutionary changes in reptiles on land (dinosaurs), in the sea
(ichthyosaurs and plesiosaurs) and in the air (pterosaurs), as also the fishes in water, in mammals on land and
in land plants with advent of flowering plants angiosperms and flourish of insects linked with their pollination.
The two Brazil formations present an early glimpse of this evolutionary innovation.
Santana Biota dominated by fish, Crato biota by insect
In this background, Santana biota is dominated by pelagic organisms, mainly fish (>20 taxa, actinopterygian
ray-fined fish most common; two coelacanth sarcopterygians and a condrichthyan i.e. a cartilaginous shark),
pelagic marine invertebrates being notably absent. Semi-aquatic reptiles (turtles, crocodiles) are there; truly
marine ichthyosaurs are absent. Epibenthic molluscs are known, echinoids are rare, but corals, brachiopods
and crinoids are absent; pterodactyloid pterosaurs are well-preserved sometimes with wing-membranes; a few
theropod dinosaurs, also sometimes with fossilized skins, muscle fibres, intestinal tracts and air sac are there.
Crato biota includes the most diverse Cretaceous insect assemblage in the world, including aquatic, semi-
aquatic and terrestrial groups, essentially modern in affinity (true flies, beetles, wasps, bees, cockroaches,
termites, crickets, locusts, grasshoppers, leafhoppers, bugs and water bugs, dragon flies, etc.). Other terrestrial
arthropods include scorpions, spiders, centipedes, rare decapod crustaceans. Besides there are fragments and
shoots of conifers (gymnosperms) and flowers, seeds, fruits, leaves and roots of many early angiosperms
among plants, one very commonly occurring fish species of a sister group of catfish; a flying reptile pterosaur
(wing span of 4 m/13 ft.) and another complete with its soft tissue head crest. Less common, though important
are a juvenile frog, lizards, turtles (with soft tissue) and a fossil bird (with feathers). Micro-ripples on bedding
planes represent cyanobacterial mat.
Environment, burial and preservation
CRATO FORMATION platy micritic limestone.
Shallow stagnant freshwater lake deposit with increasing salinity from aridity (also evident from the evaporite
formation overlying the Crato Formation).
Salinity-stratified water column and/or oxygen deficient stagnant bottom waters hosted only allochthonous
remains, without benthonics.
(Cont...)
382 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.12 (Cont...)

Santana and Crato Formations

Deltas of rivers at lake margins allowed some autochthonous organisms (abundant individuals of a fish species
and some crustaceans).
The common fish species suggests mass-mortality from water mixing leading to sudden increase in salinity.
The presence of cyanobacterial mat intact and, thus, the absence of grazers as well as other benthics and
bioturbating organisms also support oxygen-deficiency and high salinity. Occasional plants, feathers and tetrapod
remain (frog, lizards, etc.) and very common insects suggest drifting in from rivers or blowing in by wind from
the land around that had a thick vegetation, where lived a variety of insects and spiders; these were the food for
tetrapods and scorpions. Above them flew the birds and pterosaurs, to die and drown in strong winds.
Preservation of soft tissues may be due to rapid fossilization.
Then SANTANA FORMATION represented a shallow embayment passing into a coastal region with periodic
marine incursions to cause mixing of waters and influx of huge amount of fish from the open sea, which
subsequently died in a mass-mortality event due to sudden upward migration of increased salinity from bottom
waters. Soft tissues of fish fossils in concretions are preserved in calcium phosphate (cryptocrystalline francolite).
Some bacteria feeding on (proteins in) the carcass could produce phosphate to initiate fossilization (known
from recent examples). Concretions could form around organic remains in oxygen deficient, low pH (acidic)
conditions, with a local increase in pH in the micro-environment around the decaying body to help lime
precipitate. Putative cyanobacterial mat on the sea floor could have contributed to the process. Rapid nucleation
after burial of a carbonate concretion around the phosphatized fish could preserve fossils in 3D.
Three types of concretions in which fossils occur in Santana Formation
Concretion Shape Oval, small Large, platy Large, thick
Concretion Outline Not following fossil Following fossil outline Not following fossil
Environment Clear, oxygenated, Muddy/sandy, anoxic, Muddy, anoxic, bottom
near-shore water slightly away from shore of deeper open water

FACTSHEET A1.13
Grube Messel

Lower/lowermost part of Middle Eocene (Lutetian): near Rhine, Frankfurt-am-Mein and Darmstadt in
Germany
Record of lake biota in basins within Rhine Rift Valley with syndepositional faulting and volcanic activity and
commonly rare, forest biota from around with different kinds of habit.
Both show excellent preservation of soft parts and instances of how fossils can be used to interpret different
kinds of habit of ancient organisms and their relation to the environment.
Relatively younger age helps comparison with extant instances
Biota dominated by placental mammals and rarer, though interesting marsupials.
Placentals both
(a) indigenous primitive insectivores, early hedgehogs and ungulates, and
(b) immigrant (invading from elsewhere) and varied modern mammals.
Bat fossils very common, occur in hundreds, all insectivore, preserved excellently with skin, wing membrane,
muscle, fur, gut contents; different types of wings suggest :
(Cont...)
 Appendix 1: Fossil Lagerstätten 383

FACTSHEET A1.13 (Cont...)

Grube Messel

(a) lower level, open-space hunters,

(b) high fliers among trees and foliage,
(c) ground-level skimmers, more common as fossils, as they are more vulnerable to poisonous
(d) volcanic gases emanated at times
Primates (4 genera) include relatives of African lemurs, early pangolins. In these too, gut contents reveal food
habit.
Rodents include one squirrel and two mouse-like genera.
Ungulates include both extinct, ancestral forms and early representatives of modern ones.
Perissodactyls represented by horses (70 specimens: of foals to adults : of ˜ 14 to 24 inches height: four front
legs hooves, three hind leg-hooves: leaf-eating browsers (from gut contents).
Artiodactyls include cattle, deer, pigs, camels, giraffe and hippopotamus–all early representatives of their extant
examples plus a few primitive ones.
Birds modern and varied, of terrestrial and forest type; some showing excellent fine plumage and include owls,
swifts, woodpeckers, rollers, an ostrich-like ratites, and a single specimen of a flamingo, a waterbird.
Fish diverse and modern:
Advanced bony fishes (osteichthyans: neopterygians), one specimen of a 2 ft long eel genus (of the type which
lives for the most of its life in freshwater, while it spawns and is born in the sea) and including predators.
Amphibians like salamander, frogs and toads are few, but the latter, at least, are well-preserved (some showing
skin and muscles; some tadpoles are also there).
Reptiles are represented by semi-aquatic to aquatic turtles and crocodiles, often with complete skeletons and
even in juvenile form; terrestrial lizards and snakes are rarer, but are again often with complete skeletons.
Arthropods diverse and abundant.
Beetles (63 per cent preserved because of their strong wing cases and as they are of various types) (larvae of a
water-beetle genus that require highly oxygenated water as at water falls may have been washed into the basin);
Ants, bees and wasps are next frequent, of varied types and include fossils of giant queens (160 mm in wingspan);
Bugs, cockroaches, crickets, flies, butterflies and moths and other ground and plant dwelling insects
(arge winged insects are rarer because their carcasses would have floated on the water surface, rather than sunk
to the basin floor);
Arachnoids like spiders, mites, ticks, scorpions are only handful.
Opossums (marsupial mammal) are of two types:
(i) Tree-climber (with a long, prehensile tail and of smaller size),
(ii) Probably a ground dweller (large, short tailed).
In the flora abundant leaf, fruit, seed, spores and pollens suggest a diverse and bush forest with variation from
open water, swamp, bank-side, damp forest and drier areas as also shallow, open, oxygenated water near the
site of deposition.
Subtropical angiosperms dominate include grass-like/damp or wet conditions/ those of water lily family
(suggest shallow, open, oxygenated water conditions near the site of deposition); gymnosperms rarer (some
swamp- cypresses).
Record of soft part preservation by anoxia formed as interaction of environmental, sedimentological and
biotic factors and presenting details of biology and physiology including living, reproductive and feeding
habits.
(Cont...)
384 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.13 (Cont...)

Grube Messel

HOW: Environment and taphonomy

An extensive river system existed in the Rhine Graben. This was separated into a number of continually deepening
lakes with fault movement. Distribution and alignment of fish and some water-dwelling insects suggest flow
in water.
Some of the continually deepening lakes (e.g. Messel Lake) were crators that erupted ash, had steep and
deep- sides, and poisonous volcanic gases that caused anoxia at depths, anoxia trapped not only water-dwellers,
but also low fliers and terrestrial animals slipping into it. Fault-bound lake basins have coarse gravel/sand
lenses at the bottom and occasionally above, which represent high-energy slumps from renewed erosion (screes
at the margin due to renewed fault activity). The rest are oil-shales with clay, 15 per cent kerogen
(the oil) and 40 per cent water; volcanic ash may have contributed to the formation of shales, rich biota to the
organic matter.
Expressive algal growth led to an excess of decaying remains and, thus, using up all oxygen made the bottom-
water anoxic.
Anoxia caused partial or no decay of the organic matter of the large number of carcasses.
WHAT: Examples of good preservation
Bat fossils with skin, wing membrane, muscle, fur, gut contents; plant remains in gastrointestinal tracts of
mammals like grape seeds and varied leaves inside small horse pointing to its varied diet; complete skeletons
of turtles and crocodiles (even in juvenile form) and of terrestrial lizards and snakes; skin and muscles in frogs
and toads; some tadpoles; giant ant queens (160 mm in wingspan); evidences like leaves with fungal spots,
insect eggs on leaves, larval chewing marks, pollens found under the wing cases of beetles implying their
pollinating activity.

FACTSHEET A1.14
Baltic Amber

Mid-Palaeogene to Holocene from around Baltic Sea in northern Europe

Baltic Amber is obtained from
1. Quarrying the blue earth (mid-Palaeogene).
2. The tillite (Pleistocene)
3. The beach (Holocene)
How amber is concentrated in the three deposits
Amber forest in Streams wash out amber Pleistocene glaciation Amber from Blue Earth
Samland in Russia lumps into a glauconite eroded the Blue Earth or tillites is washed out
(between Poland clay, the Blue Earth, outcrop to redeposit and dumped in the beach of
and Lithuania) found in three layers in a it in tillites (2). North Poland, Germany and
around the marine sandstone-shale Denmark (3).
Baltic Sea. unit of mid-Eocene-early
Oligocene (1).

(Cont...)
 Appendix 1: Fossil Lagerstätten 385

FACTSHEET A1.14 (Cont...)

Baltic Amber

Modes in which organisms are commonly included in plant-resin

l Insects

– living on the bark

– living in the cracks
– living in the mosses
– flying in the forests
l Predators (e.g. spiders) are drawn by the insects
l All these may be trapped in the sticky resin secreted by the tree.
BALTIC AMBER includes 216 species of plants: a diverse assemblage of TEMPERATE,
MEDITERRANEAN, SUBTROPICAL, even TROPICAL TYPE
05 bacteria 01 slime mould 18 fungi 02 lichens
18 liverworts 47 mosses 02 ferns 52 gymnosperms
101 angiosperms (2/3 flowers, fruits or seeds, remainder leaves and twigs).
Animals include
Most common are more advanced winged insects
Certain kind of flies having aquatic larvae;
Diverse cockroaches, crickets and grasshoppers,
Carvings, termites, bugs, book-lice that live under bark and on dead organisms (found in the binding paste of
old books);
Large, predatory, flying insects such as mantis;
Beetles of varied types, flies with a single pair of wings, etc.;
Ant, bees and wasps and a host of others; and
Arachnids (spiders, >90 per cent of fossil spiders known from amber), scorpions, etc.
Also present
primitive hexapods (insects);
microscopic roundworms (one such parasitic form is preserved emerging from its host);
crustaceans; predatory centipedes, detritivore millipedes; rare land snails; and lastly
bird feathers, a mammalian footprint and some hairs.
Biota, particularly the flora, indicate:
1. change in climate over the long period during which amber formed;
2. different altitudes and positions of mountainous terrain; and
3. the presence of isolated ‘tropical’ islands in ‘subtropical and temperate’ regions as in Florida today.
In essence the ‘Baltic Amber Forest’ was temperate to tropical and wet
NB: Change from plant resin to amber involves loss of volatile oils, polymerization and hardening to form a soft, non-
pliable substance called COPAL that dissolves in organic solvents. Further polymerization produces AMBER which
is harder and does not dissolve in any organic solvent.
386 Appendix 1: Fossil Lagerstätten

FACTSHEET A1.15
Rancho La Brea

Number Puzzle
RANCHO LA BREA, situated in the city of Los Angeles and enclosing Pleistocene Ice Age fossils,
stands out on three major points:
I. Record of > 600 species including 160 plants + 440 animals, breaking up into
50 mammals + 135 birds represented by
> 1,000,000 fossils and ~ 100,000 fossils
II. Carnivores outnumber herbivores forming an inverse trophic pyramid
90 per cent carnivores as against 10 per cent herbivores
70 per cent carnivorous birds as against 30 per cent herbivorous birds
It is explained in the following manner:
1. one herbivore body trapped in asphalt;
2. a pack of carnivores attracted and trapped; and
3. scavengers try to feed upon the whole trapped lot and are themselves trapped.
III. Young (juveniles), aged and maimed individuals more frequent.
(May also be a sampling bias).
Rancho La Brea provides one of the world’s richest samples of Pleistocene Ice Age fossils in a record about
10,000 to 40,000 years old and, thus, throw light on this glaciation-extinction event in North America.
Pleistocene mass extinction wiped out mammoths, mastodons, horses, tapirs, camels, ground sloths and their
predators like sabre-toothed tigers (about 73 per cent of large mammals in North America)
NB: Indiscriminate hunting by the then humans is held as a probable, may be even major cause of extinction.
Biostratinomy:
Marine Tertiary shale-sandstone-oil sand interbeddings occur below Quaternary alluvium.
From around 40,000 years ago, crude oil migrated along the limbs of the fold upwards to the crest into the
overlying horizontal beds of fluvial alluvium. Lighter fraction of petroleum, natural asphalt formed sticky
pods of at the surface (asphalt is the naturally occurring bituminous fraction, etc. of petroleum; tar is not
asphalt).These asphalts, viscous in summer, formed natural traps for organisms, which was preserved when
asphalt cooled and solidified in winter and was then covered by fluvial deposits. New traps formed each
summer in annual cycles.
Rancho La Brea is, thus, a Concentration Lagerstätte which preserved huge number of fossils. It does not
preserve soft parts, but rapid burial and asphalt-impregnation preserve even organic constituents and details of
bones.
Constituents of the biota:
Fractured skull (murdered and dumped/ a ritual) and part-skeleton of a human female (about 5 ft. tall; 20-25
years of age; of a C14 age of 9000 ybp).
More than 1600 individuals of 4 species of Canis, viz. Dire wolf (large head, strong jaws, massive teeth and
huge packs suggest them to be the major predator), domestic dogs (one associated with the female human) and
two others.
Felidae represented by sabre-toothed tiger (as big as the African lion; the large, curved canine, really fragile
might have been used for opening the soft underbelly of the dead victim than for stabbing and killing), the
American lion, puma and jaguar.
(Cont...)
 Appendix 1: Fossil Lagerstätten 387

FACTSHEET A1.15 (Cont...)

Rancho La Brea

Mammoth (Mammuthus imperatus; ˜ 13 ft tall; 5000 kg in weight, grass-eater) and mastodon (6 ft tall; leaves
and twig-eater) among proboscideans.
Ground sleuth (molars suggest grass- eating: ˜ 6ft tall); variety of large carnivores like different kinds of bear;
diverse large herbivores like bison, horses, tapirs, camels, llamas, deer. Among smaller mammals, carnivores
like racoons, badgers, etc. insectivorous shrews and moles; rodents like mice, rats, squirrels; lagomorphs like
hares and rabbits, bats.
Largest fossil record of birds in the world; predators or scavengers, e.g. vultures trapped when feeding on
carcasses; waterbirds (herons, ducks, geese, etc. probably allured by shining asphalt surface to land on it) ;
eagles, hawks and falcons and storks, turkeys, owls and various song birds.
Lizards (seven types), one pond turtle, five amphibians (frogs, toads, salamanders), three species of fish.
Freshwater molluscs (5 bivalves and 15 gastropods), 11 terrestrial (pulmonate) gastropods, insects such as
grasshoppers, crickets, termites, bugs, beetles, flies, ants, wasps, scorpions, millipedes and several spiders.
Wood, leaves, pine cones, seeds, pollens and diatoms among floral elements.
The biota represent a virtually complete terrestrial ecosystem in a cool, glacial climate of terrain replete
with freshwater ponds and streams and rich vegetation of mountainous, deep wet valleys, river banks and
plain types; plains supported horse, camels, mammoths and other ungulates, attracting carnivores such as
dogs, cats and bears; smaller mammals were similar to those of recent times suggesting closely similar climate.
Appendix
2
Lab Exercises
A2.1 Collecting Samples and
Preparing Them
No doubt, theory and practice are intertwined.
Hence, answer to any question in field or laboratory
would require some or other theoretical
understanding. How we should collect plant
impression fossils from a shale will have a different
set of answers from those in reply to how an echinoid
test may be collected and extracted from hard
limestone. The two kinds of fossils differ in their
composition, mineralogy and taphonomy. We must
have a background idea to get the best results for
each individual case. Thumb rules are not always
paying. Yet, a few general points are added here.
Fossil hunt in field, like mineral hunt, may be
frustrating to a student till he comes across a good
occurrence. The rewarding relief may add extra
impetus to the hunter; he just tends to pounce upon
the treasure of nature. A sense of restraint is
necessary at this stage.
Collection of fossils must be based on the
‘Principle of Three E-s’: Economy, Efficiency and
Ethics. Any fieldwork consumes time and money.
Bringing load of collections implies freight charges
and, thus, more costs. So, a collector needs to be
economic; too big a sample means unnecessary
expenses; too small a sample also means that. A
388
sample found too small, back in the laboratories,
may demand a fresh collection and, thus, a fresh
fieldwork and fresh drainage of fund. Choosing
the right size relates to efficiency of collection.
When the team of vertebrate palaeontologists from
the Indian Statistical Institute of Calcutta located
the two phytosaur reptile fossils lying side by side,
they needed to dig the earth, drive wooden planks
around the mass of the earth housing the fossils
and, thus, enclose that mass within a wooden box
of several cubic feet, lift the box by crane onto a
fairly large vehicle and then bring it to their
laboratory. Similarly, when they found dis-
articulated bones of the large sauropod dinosaur,
they had to carry several tonnes of the earth, the
rock and the fossils embedded in it. But for work
on microfossils or for isotope studies for palaeo-
climate reconstruction, a few cloth or polythene
sachets may prove to be adequate. So, efficiency
of collection will depend on the issues on table.
A third point is as vital as the two discussed.
A collector needs to be self-retraint. He must warn
himself to never make a single stroke of hammer
and chisel at any fossil if one does not need it; one
should not destroy it if one is sure that one cannot
take it out of the rock; better try a second time
with more resources and means. The fossil, if and
when destroyed by the collector, might have

otherwise proved to be very vital and interesting
for any future scientific work. It is true that
urbanization, exploration-quarrying for industria-
lization or simple greed of consumerism have
thoughtlessly taken toll of huge amount of natural
treasures. But a palaeontologist is a scientist who
can never play to the tune and destroying a natural
treasure for simple pleasure or whims.
Preparing fossils in field or in laboratory for
further work, hinges basically on disaggregating
the fossils from their rock-matrix. Here again each
specific case has its own problems. The host rock
may be soft and shaly or clayey, in which case it
can be made softer with soaking in water. That
solves the problem; fossils can be separated
thereafter. A little amount of chemicals (alkalis)
may be needed in some cases. When the host rock
is well-bedded and fissile, careful cleaving along
the beddings may yield beautiful fossils preserved
on the bedding. Many plant fossils come out this
way. Calcareous fossils preserved in shales, even
marls are not difficult to extract. The problem
mounts when the host rocks and the enclosed
fossils are of the same composition, for instance,
calcareous fossils in hard well-cemented
limestones. Use of hydrogen peroxide or other
reagents or instruments like ultrasonic vibrators
may help; but often the solution may lie in patiently
removing the matrix by soft strokes of hammer or,
failing everything, in looking for microfossils in
thin sections under microscope. All said and done,
it must be borne in mind that this is the toughest,
yet unavoidable part of any research work, on
account of the boredom and fruitlessness involved
till the end is reached. But it pays fully in return,
when the finished prepared samples shine out in
full and clean shape.
Details of different methods and techniques
of collection and preparation of samples are
provided in a number of publications on the topic
and individual research papers. Kummel and Raup
(1965) is a useful handbook for the purpose.
Appendix 2: Lab Exercises 389
A2.2 Describing and
Identifying Collected
Samples
It has already been indicated that identification and
naming of fossils are done on the basis of
comparison with previously described similar
fossils; descriptions are found in monographs and
other publications. In countries where tradition of
documentation and preservation of documents in
libraries, museums or such other institutions are
not upto the mark, this process of consultation faces
a problem to reckon with. Microfilms and
computerised data may serve as alternative in
recent days.
For students and their routine exercises in the
laboratory, however, such elaborate process
remains out of scope. And hence, in such exercises
emphasis is generally placed on objective, faithful
description and comparison among those
descriptions. Here a few examples are provided to
help objectively describe fossils of some major
invertebrate groups. In each case, a format for
description is given; it is followed by two or three
examples of how to write a description of a genus
(in its simplest form, adequate for classwork).
Next, there are examples of how to make
comparisons at generic level. For necessary
definitions and other details relevant and
corresponding sections in the text will have to be
consulted. It must be borne in mind that the
descriptions provided are examples; concrete
individual specimens may differ slightly or largely
on this or that character.
A2.2.1 Anthozoa (corals): Format for
Description
Solitary or colonial: Shape of corallites in
solitary corals and shape and arrangement of
corallites in the colony. Wall: epi-,para-or
synapticulotheca. Internal features:
390 Appendix 2: Lab Exercises

1. Septa: long/short; major/minor; straight/ Wentzelella: Fossil colonial form with a

wavy, etc. perforate/ spinose/ solid.
2. Tabulae: complete/incomplete; convex up/
concave/ flat.
3. Dissepiments: present/ absent; dense or thin;
circular or elongate.
4. Axial structure: solid/ spongy.
A2.2.2 Anthozoa (corals):
Description and examples
Montlivaltia: Fossil of a simple/solitary coral;
corallite short conical, turbinate; wall absent; septa
externally visible, numerous, radially symmetrical,
both long and short (major and minor), straight;
dissepiments and axial structure absent; tabulae
cannot be found[1] synapticulae present; small
horizontal bars between septa.
compound corallum; individual corallites prismatic,
polygonal in section; adjacent corallites have walls
(epithecal) fused; cerioid arrangement; septa
numerous and largely radial in disposition, both
long and short, slight wavy; dissepiments
prominent, elongate, occur in a broad peripheral
zone; axial structure is a composite structure formed
of two sets of tabellae; tabulae cannot be found.[1]
A2.3 Brachiopods: Format for
Description
Shell: Number of valves, their size and
symmetry; terms for the valves. Shape of shell on
(i) commissure and (ii) lateral profile : length,
breadth and thickness.

FACTSHEET A2.1
Anthozoa (Corals); Compare and Contrast; Example
Genus Halysites Favosites Syringopora

Similarities
Habit Colonial Colonial Colonial
Arrangement Parallel Parallel Parallel
Septa Absent Absent Absent
Dissepiments Absent Absent Absent
Axial Structure Absent Absent Absent
Dissimilarities Chain coral Honeycomb coral Organpipe coral
Corallites Tubular Prismatic Tubular
Cross-section Elliptical Polygonal Circular
Arrangement In a linear seriesa In massive aggregatesb In loose bundlec
Tabulae Not found in usual specimens; may be found in thin sections which show
Complete flat horizontal Incomplete, convex up Complete concave up
a
Each corallite longitudinally in contact with two adjacent corallites at the end of the major diameter of its cross-section
b
Corallites are in contact, with walls fused (with mural pores, rarely to be found)
c
Corallites run parallel with space left in between; they are connected by short, horizontal cross-tubes

1Tabulae cannot be found from outside and so are not visible without a vertical / longitudinal scetion. In that
case, while describing a specimen, the point remains undecided whether tabulae are present or not. They
should better be described as ‘not found’. A feature can be described as absent, only when it is not found
even in views or sections where it is expected. Thus, in Cystiphyllum, even the calical view does not show
any septa; had they been present, they must be seen there. Hence, we can infer ‘septa absent’ in Cystiphyllum.

Posterior margin: Beak and umbo;
curvature (convex or concave), prominence (highly
curved, moderate or flat), etc. of umbo.
Hinge: Strophic or nonstrophic; hingeline
long or short, straight or curved; interarea whether
present, if so, whether on one valve or both; their
relative prominence.
Pedicle opening: Whether in one (pedicle)
valve or shared by both valves; delthyrium,
notothyrium, pedicle foramen: present/absent;
if present, shape, etc.
Punctae: present /absent.
Surface ornaments: (i) concentric growth
lines, generally grooves, (ii) radial ribs and
(iii) median sulcus and ridge; spines, etc.
A2.4 Brachiopods: Example of
Description
Productus: Fossil of a bivalved, strongly
inequivalved, equilateral (i.e. bilaterally sym-
metrical across the hingeline), closed brachiopod
shell; pedicle valve (P-valve) highly convex and
much larger than a concave, smaller brachial valve
(B-valve); shell transverse trapezium shaped in
commissure, concavo-convex in lateral profile,
L<W >Th.
Shell with a strophic hinge; hinge line straight,
as long as the width of the shell; no hinge area;
pedicle (P-) umbo strongly convex/ highly incurved,
brachial (B-) umbo concave; no pedicle opening.
Surface with strong ribs, both concentric and
radial, the former more prominent near the
umbones; strong spinal nodes on ribs of P-valve
suggest fixosessile or quasi-infaunal habit, without
a pedicle.
Pentamerus: Bivalved, inequivalved,
equilateral closed fossil brachiopod shell; shell oval
in commissure, slightly longitudinal (L>W>Th);
biconvex profile.
2 Unlike brachiopod shells, bivalve shells are usually found disarticulated, with individual valves preserved
separately; the reason is discussed in section 10.8.1. In such cases, the description may be modified as “single
disarticulated left/right (as the case may be) valve of a bivalved, equivalved, inequilateral bivalve shell.” It
should also be mentioned whether the specimen is a fossil or a recent specimen.
Appendix 2: Lab Exercises 391
Shell non-strophic, with a short, curved hinge
line, no hinge area, highly incurved umbones and
B-umbo overlapped by/ tucked below P-umbo; no
pedicle opening.
Surface smooth, with a few weak concentric
growth lines; anterior margin trilobate from the
presence of a shallow median sulcus on P-valve;
valves anteriorly gaping.
A2.5 Bivalves: Format for
Description
Shell: number of valves, their size and symmetry;
terms for the valves; right and left valves.
Shape, dimensions and orientation;
geometry of the commissure; anterior-posterior;
length, height and thickness.
Dorsal margin: umbones prosogyral/
opisthogyral/orthogyral; hinge plate vertical or
inclined in the opposite direction from that of the
main valve; hinge area; hinge line straight or
curved. Lunule (anterior) or escutcheon (posterior):
present/not; if yes, character.
Ligament at dorsal margin external
opisthodetic/prosodetic or internal amphideticin
triangular pit, i.e. resilifer or spoon-shaped
chondrophore.
Dentition: Edentate, Taxodont, Dysodont,
Isodont, Schizodont, Heterodont, Pachydont,
Desmodont.
Adductor muscle scar: Monomyarian/
dimyarian, isomyarian/anisomyarian. Pallial line
entire, shell integripalliate/with pallial sinus, shell
sinupalliate; pedal scar.
Ornaments: Radial and concentric. But
there is no median.
A2.6 Bivalves: Example of
Description
Arca: Shell equivalved[2], inequilateral; bilateral
392 Appendix 2: Lab Exercises

FACTSHEET A2.2
Brachiopods; Compare and Contrast, Examples

Syringothyris Rafinesquina Strophomena Leptaena Streptorhynchus

Similarities: Bivalved, inequivalved/Strophic hinge; umbones non-overlapping/Shell largely transverse (W>L)/Hinge

line straight as long as the width/Delthyrium closed by plates (deltidial)/Surface ornaments similar on both valves
Dissimilarities:
Commissure
Triangular Semi-elliptical Trapezium Trapezium Triangular
Width : Length
W>>L W~L W>L W>L W>L
Hinge area
Very high Low, on P-valve Low, on both valves Prominent
Profile
Biconvex Concavo-convex Resupinatea Plano-convexb Convexo-concave
Umbones
Both convex P-umbo convex P-umbo convex Both convex P-umbo concave
B-umbo concave/flat B-umbo concave B-umbo convex
Pedicle opening
On P-valve On P-valve On both valves On P-valve
Surface ornaments
Strong ribs Very fine ribs Very fine ribs and Thin ribs Thin ribs
Sulcus on P-valve Concentric folds (rugae)
a P-valve convex at umbo, concave thereafter; B-valve concoave at umbo, then convex
b P-valve with a trail near anterior margin, which is an abrupt knee-shaped bend that brings anterior growth of the valve at
right angles to the main valve.

Terebratula Athyris Atrypa Rhynchonella

Similarities: Bivalved, inequivalved/ Non-strophic hinge/ Umbones convex; P-umbo overlaps B-umbo/ Hinge line
short, curved; no hinge area/Surface ornaments similar on both valves
Dissimilarities:
Commissure
Elliptical Elliptical Circular Rhombic
Width : Length
L>W W>L W~L W>L
Profile
Biconvex Biconvex Convexo-plane Biconvexa
Pedicle opening
Large, circular foramen Small, circular foramen Absent, atrophied Circular foramen
Surface ornaments
Weak, concentric Distinct, concentric Both radial and Strong radial ribs
growth rings growth rings concentric ribs
Anterior margin
Straight Shallow sinus Very weak sinus W-shaped from strong
sinus on P-valve
a Presence of a strong sinus on P-valve renders the shell an apparent convexo-concave shape.
 Appendix 2: Lab Exercises 393

FACTSHEET A2.3
Bivalves: Compare and Contrast; Examples
(In each case, a single disarticulated valve is described here as an example)

I.
Cyrena Venus
Similarities
Commissure Trigonal (L~H)/Shell nearly equilateral/Isomyarian/Heterodont
Curved hinge line, no area/Ligament external, opisthodetic/Only concentric growth rings
Dissimilarities
Dentition Both cardinal and lateral Only cardinal teeth and sockets,
teeth and sockets no laterals
Pallial line Pallial line entire Pallial sinus angular
Integripalliate Sinupalliate
Ventral margin Smooth Internally dentate
Lunule and escutcheon Absent Both present

II.
Unio Mya
Similarities
Subelliptical, long (L>>H)/Distinctly inequilateral/Anisomyarian/Curved hinge line, no area
Dissimilarities
Ligament External, opisthodetic Internal, amphidetic in chondrophore
Pallial line Entire With a very broad sinus
Adductor scar Both quadrate, but not Anterior scar long elliptical,
similar in size posterior quadrate
Dentition Schizodont with some Edentate
rugged teeth
Pedal Scar Present Absent

III.
Ostrea Pecten Spondylus
Similarities Nearly equilateral commissure/Monomyarian
Dissimilarities
Commissure Triangular Orbiculara Triangular
Hinge line Slightly curved, short Straight, with ears at ends (in both)
Dentition Edentate Dysodontb Isodontc
Hinge Area Small area No area Distinct area
Ligament External,on grooves Internal triangular pit or resilifer (in both)
Shell form Foliaceousd Byssate e Foliaceousf
Surface Concentric growth rings Strong radial ribs Concentric growth rings
and weak radial ribs
Adductor scar Deep Weak Weak
a
(Tear-drop shaped)/
b
With two teeth/socket weak radial
c
With two equal teeth strong, hooked teeth, may be bifid and two broad-based socket
d
Cemented to bottom by left valve (mark of cementation near umbo on the outer side)
e
Temporarily attached; byssal notch below anterior ear/
f
Attached by spines (on right valve)
394 Appendix 2: Lab Exercises
symmetry plane running in between the two valves,
right and left, and containing the hinge; shell
(and the valves as well) with larger portion
posterior to beak. Commissure trapezium-shaped,
shell moderately convex[3]; L >H >Th.
Shell (or valve) with a long, straight hinge line
and a distinct triangular area marked by parallel
grooves[4] for external, opisthodetic ligament;
umbo prosogyral, strongly curved[5]; taxodont
dentition, with numerous small, more or less
similar teeth and sockets, alternately disposed
nearly at right angles to the hinge line (which is
the upper edge of the inclined hinge plate).
Shell (or valve) anisomyarian with quadrate
scars; posterior scar much bigger, joined by an
entire pallial line to the anterior scar (shell
integripalliate).
Surface with both concentric grooves (marking
growth rings) and radial ribs, latter more
prominent. Ventral margin corrugated.[6]
Venus: Single disarticulated left (say, for
instance) valve of a bivalved, equivalved,
inequilateral bivalve shell; valve inequilateral with
larger portion posterior to beak. Commissure curved
trigonal, valve moderately convex; L ~H >Th.
Valve with a short, curved hinge line; no hinge
area; a narrow platform posterior to beak for
external, opisthodetic ligament; umbo prosogyral,
slightly curved[5] heterodont dentition, with two
distinct cardinal teeth (one of them bifid) below
the beak and radiating downwards and outwards
and three sockets on the vertical hinge plate, but
no lateral teeth. Lunule[7] and escutcheon[8] on the
dorsal margin[9].
Valve nearly isomyarian with ovate scars;
valve sinupalliate with a distinct angular pallial
sinus.
3 In some species, it may be highly convex.
4 Ligamental grooves may otherwise be inverted V-shaped.
5 Curvature of umbo may vary from strong to weak, prosogyral to opisthogyral (in rarer cases).
6 Surface ornaments may vary in details.
7 Lunule is a short, heart-shaped depression in front of the beak on the hinge line.
8 Escutcheon is a long, partially depressed region behind the beak on the hinge line.
9 In disarticulated single valve, only half of the both will be seen.
Surface with concentric grooves (marking
growth rings). Ventral margin internally dentate.
A2.7 Gastropods: Format for
Description
Shape of shells, type of coiling, dimensions,
symmetry, orientation, etc.: Conispiral/
planispiral/pseudoplanispiral; bilateral symmetry:
present or not; coiling clockwise/right-handed/
dextral or anticlockwise/left-handed/sinistral
(both viewed from apex); coiling evolute/advolute/
involute/convolute;
Directions: anterior/oral/distal- posterior/
apical/proximal; dimensions (a) spiral angle
(b) relative length (or height) of spire and body-
whorl, (c) curvature of the shell or whorl surface
(convex or flat, rarely concave)
Aperture and associated features:
Geometry of apertural opening—circular, semi-
circular, straight, narrow, rectangular, curved slit-
like, crescentic, etc.
Outer and inner lips of aperture—various
combinations, such as smooth and simple, inner
dentate, outer smooth, inner with folds
(columellar), both lips dentate and outer lip
infolded, outer with a slit.
Siphonal canals–absent/present, if present,
long/short, drawn out/abrupt, straight/twisted, only
anterior/anterior, posterior both.
Internal structures: Columella; smooth/
twisted; Umbilicus: shell anomphalous, phanero-
mphalous, cryptomphalous.
Surface ornaments: Apertural, generally
grooves/spiral, ribs or rows of spines or radial or
longitudinal, grooves or varices.

A2.8 Gastropods: Description
and Example
Fusus: Univalved, asymmetrical, conispiral
(recent or fossil, as the case may be) gastropod
shell. Coiling involute (later whorls slightly
overlapping earlier ones and visible from outside).
Shell spindle shaped, fusiform, with acute spire
(~30o) and anteriorly drawn out body whorl, almost
equal in height/length; whorl surface convex with
fairly deep whorl sutures.
Aperture right-handed, subelliptical with a
long anterior canal; siphonostomatous; inner lip
with weak columellar folds, outer lip smooth.
Shell anomphalous (without umbilicus);
surface with spiral grooves and broad radial/
axial ribs.
Trochus: Univalved, asymmetrical, conispiral
(recent or fossil, as the case may be) gastropod shell.
Coiling involute. Shell conical, trochiform, with
acute spire (about 45°) and flat distal base; body
whorl almost equal in height/length to spire;
whorl surface flat with weak whorl sutures.
Aperture right-handed, rhombic with entire
margin (without canal); holostomatois; both lips
smooth.
Shell phaneromphalous, with a small umbilicus;
surface with spiral and apertural grooves.
A2.9 Cephalopods: Format for
Description
Shape of shells, type of coiling, dimensions,
symmetry, orientation, etc.:
Bilateral symmetry: present or not;
Shape: Simple cones (longicone/brevicone/
orthocone/cyrtocone)/planispiral (sphaerocone/
cadicone/planulate/oxycone/serpenticone)/
coniplanispiral/aberrant shell shapes partially coiled;
Diameter and thickness
Aperture and associated features:
Geometry of aperture; height and width:
depressed (W>H)/ compressed (W<H); Weak or
strong impressed zones;
Appendix 2: Lab Exercises 395
Umbilicus large/small; deep/ shallow .
Internal structures or their external
expression:
Suture: Goniatitic/Ceratitic/Ammonitic/
Pseudoceratitic;
Siphuncle
Surface ornaments:
Spiral nodes or tubercles; Radial ribs, simple,
bi-/trifurcating, point of bi-/trifurcation; apertural
grooves; peripheral keel/sulcus, number and nature
A2.10 Echinoids: Description and
Format
1. Shape, symmetry, orientation and
dimension of test/tests: symmetry radial or
bilateral; shape includes geometry of the
ambitus and the curvature (concave/convex/
flat) of oral and aboral surfaces; fixing
orientation involves marking anterior-posterior
that defines the length and aboral-oral
(dorsoventral) that defines the height.
2. Corona:
(a) Ambs: Similarities-dissimilarities;
structural characteristics, viz. simple,
subpetaloid, petaloid; appearance: raised,
depressed or flat in relation to interambs;
pore-zones and pores: number, shape,
position and conjugation; width of pore-
zone, etc.
(b) Interambs: Similarities-dissimilarities;
shape and size.
(c) Tubercles: Varieties and their nature;
fascioles and their types and positions.
3. Peristomial and periproctal systems: Their
position, alignment and outline geometry,
floscelle, lantern, food groove, etc. in
peristome. Normally, these two systems are not
preserved in fossils; they are represented by
corresponding openings. Mouth and anus are
really smaller openings included within the
larger openings of peristome and periproct,
respectively.
4. Apical/oculogenital system: Position, shape;
number of plates, their position; madreporite.
396 Appendix 2: Lab Exercises

FACTSHEET A2.4
Gastropods: Compare and Contrast, Examples
I.

Turritella Cerithium Fusus Murex

Similarities Coiling: Conispiral; partially overlapped earlier whorls visible from outside/Shell sort of
conical in shape, acute spire (~30o)/Aperture subcircular/subelliptical.
Dissimilarities
Shape Conical Spindle-shaped
Turretted Turretted Fusiform Fusiform/Muriform
Umbilicus Holostomatous Siphonostomatous in all three
Canal Absent Short twisted anterior Long anterior Long anterior
Inner lip Smooth With columellar folds Columellar folds Smooth
Ornament Spiral ribs Spiral nodes Spiral grooves Spiral grooves
Apertural Grooves in both Broad axial ribs Three varices
II.

Nerita Natica Cypraea Bellerophon

Similarities Shell globose or subglobose in shape, whorl surface convex/largely or completely overlapped
earlier whorls slightly or none visible from outside
Dissimilarities
Coiling Asymmetrical conispiral in the three Bilateral, planispiral
Shape Naticoid Naticoid Ovoid (cypraeiform) Globose (D=Th)
Overlap Earlier whorls partly visible Only last whorl visible
Aperture Semicircular Semicircular Curved slit-like Crescentic
Inner lip Dentate Smooth Dentate for length Smooth
Outer lip Smooth Smooth Dentate and infolded Occasional slit
Umbilicus Large, filled Large, open None None
(cryptomphalous) (phaneromphalous) (anomphalous)
Canals Absent Absent Both anterior and posterior Occasional slit
abrupt and twisted
Surface Spiral Apertural Smooth Smooth
grooves grooves
III.

Conus Physa Voluta

Similarities Between spire and body whorl spire shorter
Dissimilarities
Shape Obconical Conoidal (biconical with Volutiform
curved enveloping surface)
Spire Obtuse Acute Acute
Whorl surface Flat Convex Flat
Aperture Narrow, rectangular Elliptical Oval
Canal Anterior, abrupt None Anterior abrupt and wide
Inner lip Weak columellar folds Smooth Strong columellar folds
Outer lip Smooth Smooth Smooth and flunged outwards
Umbilicus None Present small None
Ornaments Apertural grooves Apertural grooves Apertural grooves and spiral nodes
 Appendix 2: Lab Exercises 397

FACTSHEET A2.5
Cephalopods: Description, etc. and Examples

1. Most cephalopods are preserved as internal molds, in which the shell material is generally not retained.
Such preservation is expected to show sutures on the outer surface of the mold, but that occurs never as a
rule. Coarser surface ornaments like ribs and tubercles are reflected on the mold surface, albeit finer
structures like apertural grooves, that mark the growth lines are found only if there is some portion of the
shell substance.
2. Ammonoids and belemnoids are extinct groups. Particularly with the former, systematics is ever changing
with more and more work. Hence, the descriptions provided here are absolutely elementary, meant only to
introduce the generic variations in the group.)
Orthoceras and Baculites
Both the genera have straight shells. But Orthoceras has a straight cone (orthocone; rather cylindroconical on
account of its slowly tapering form that never reaches a pointed apex) with a circular cross-section. Baculites
has a baculiticone, in which the straight cone has an oval/elliptical cross-section. Orthoceras has simple straight/
smooth sutures parallel to each other, from simple concave upwards septa; Baculites has intricately frilled
ammonitic suture. Surface of Orthoceras is without any feature, hence smooth; Baculites has faint chevron
ribs across the cone. Orthoceras has a central siphuncle, whose position may be confirmed if there is any
septum visible from outside. Baculites has a marginal siphuncle. Previously Orthoceras was assigned an age
of Ordovician to Triassic; presently it is held as an Ordovician genus. Baculites is a heteromorph ammonoid of
Upper Cretaceous age.
Perisphinctes, Macrocephalites and Scaphites
The three genera are ammonites with bilaterally symmetrical planispiral shells, complex ammonitic suture
and highly ornamented surface, in which numerous fine radial ribs bi-trifurcate. But they differ in other
details.
In Perisphinctes, ribs bifurcate only once and near the periphery or venter; there is no node or tubercle at the
point of bifurcation. In Macrocephalites, they bi-trifurcate midway between umbilicus and venter, rather
nearer the former; there are no tubercles here either. In Scaphites they branch out at different points near
umbilicus, midway or venter, often with a tubercle at the point of division; Scaphites has tubercles arranged
in two spiral rows.
The genera differ also in shape, etc. Perisphinctes has a planulate, serpenticone, evolute shell with a number of
whorls visible; its aperture (often broken in fossils and represented as whorl section) is circular with weak
impressed zones; umbilicus is shallow, yet broad. Macrocephalites has a cadicone to sphaericone, nearly involute
shell with virtually the last whorl only visible; its aperture is subcircular; but since the coiling is compact, it has
two strong impressed zones and an umbilicus is moderately deep and large. The two are regarded as important
Jurassic forms. Scaphites is an Upper Cretaceous marker; it is a heteromorph, in which the earlier portion of the
shell is coiled involute (only last whorl visible), but later parts are uncoiled and separated from the former. Thus,
being made fragile, the shell does not generally retain the later uncoiled part. The early portion is a cadicone; the
aperture is rather squarish from a flatter venter with two strong impressed zones; umbilicus is deep and small.
Ceratites, Acanthoceras and Amaltheus
The three genera are artificially grouped here on the basis of their having coarse radial ribs as ornaments.
Ceratites has a thick, planulate, symmetrical planispiral shell, evolute with earlier whorls largely visible; D >
Th. Acanthoceras also has a thick, planulate, symmetrical planispiral shell, but less evolute with earlier whorls

(Cont...)
398 Appendix 2: Lab Exercises

FACTSHEET A2.5 (Cont...)

Cephalopods: Description, etc. and Examples

partially visible; D>Th. Amaltheus is a bilaterally symmetrical planispirally coiled cephalopod with an oxyconic
(thin disc-shaped) shell, with D>>Th.
In Ceratites, the aperture is squarish from a flat venter with weak impressed zones; umbilicus broad and shallow;
suture is ceratitic. In Acanthoceras, the aperture is squarish from a flat venter with weak impressed zones;
umbilicus broad and shallow; suture is ammonitic, whereas Amaltheus has a lenticular aperture with height >
width, hence compressed. Its shell is involute and, hence, has strong impressed zones. Suture is ammonitic;
umbilicus broad and shallow. .
In Ceratites, the coarse ribs radiate on two sides; they are slightly curved near venter with a weak node at the
point of curvature on each rib; ribs do not cross the venter, hence the venter is smooth.
The surface of Acanthoceras is with radial coarse ribs, a few merging towards umbilicus. Three spiral rows of
weak nodes, one near umbilicus and two near the venter nodes occurring on the ribs. Amaltheus surface has
broad, fulcate ribs, rarely bifurcating. They do not cross venter which is occupied by a ropy keel.
Ceratites is a Traissic genus, Amaltheus Jurassic and Acanthoceras Cretaceous.
Nautilus
It is a well-known genus, whose definition, like Orthoceras, has changed; earlier known as a genus ranging from
Carboniferous to Recent, it is now restricted to a Cenozoic form of Olgiocene to Recent age.
It is a planispiral form, involute (with only last whorl visible), sphaericone with diameter nearly equal to thickness.
It has a crescentic aperture, generally depressed (height < width) with a deep, but small umbilicus. The surface
is smooth (when the shell is preserved, fine apertural grooves may be seen); on internal molds wavy nautilitic
suture may be found.
Belemnites is a stratigraphically important coleoid of Jurassic to Cretaceous age. It has three parts in its
shell:
1. A phragmocone, which compares with the phragmocone i.e., the shell of other cephalopods. It is a chambered
cone with concave upwards (anteriorly) septa. It is made of aragonite and is, hence, unstable. It was the main
house of the animal.
2. Dorsal side of phragmocone is extended anteriorly into a long, flat, tongue-shaped projection, called pro-
ostracum. Its function is not well-known and may have been used as a protection.
3. The third part called guard is the most important as fossils. It is a bullet-shaped, solid body formed of
calcite fibres radially arranged and concentrically grown, though the fibres converge not at the centre, but
somewhat towards the ventral margin. Being calcite, the guard is stable, making the guard a common fossil.
It has a conical cavity at its anterior end; the tip of the fragile phragmocone fits into the cavity, called
alveolus. Hence, normally, we find, part of a guard with alveolus and a remnant of phragmocone material
preserved in it.
This configuration is inferred from a few well-preserved specimens and by comparison with recent representatives
of coleoids, viz. squids, etc. The guard seems to have acted as a measure to counter-balance the weight of the
animal in the phragmocone and thus to keep it horizontal, while swimming, much in the way explained in
sections 12.11 and 12.12. Being made of uncrystallized calcite, belemnite guard is very suitable for
palaeotemperature studies on oxygen isotopic ratios; in fact, belemnite shells of a Gretaceous marine PeeDee
Formation have been used as standards.

A2.11 Echinoids: Description, etc.
and Examples
Section 13.14 contains a brief morphology of a
few common echinoid genera. They may be used
here as examples. In addition a few other genera
are mentioned here either with a full description
(as a model) or compared with genera already
mentioned.
Stygmatopygus: Bilaterally symmetrical,
irregular echinoid; test dome-shaped with a higly
convex aboral surface, flat oral; ambitus nearly
oval/elliptical; length> breadth>height; maximum
height slightly behind the centre of aboral surface.
Ambs differentiated in size, though similar in
subpetaloid nature and similar pore character in
all ambs; ambs flush with interambs; posterolateral
petal smallest in length and width, unpaired largest;
pore-zone narrow in all ambs; pores uniserial, inner
circular and outer slit-like in petals, lost in the
remaining parts of the ambs. Interambs wide;
interambs and inter-pore-zone of ambs studded
with granules, smaller on aboral surface, larger
on oral.
Peristome subcentral, with a well-developed
floscelle; periproct supramarginal, elongate bottle-
shaped, longitudinal in disposition.
Apical system eccentric, slight anterior.
Echinolampas differs from Stygmatopygus on
the following characters: Echinolampas has
posterolateral ambs largest, unpaired smallest;
periproct inframarginal, anal opening being
elliptical, transverse; oral surface may be concave
or even convex; floscelle present, but not as well
developed as in Stygmatopygus.
Hemiaster, a well-known spatangoid form
(i.e. with heart-shaped ambitus, etc.) differs from
Schizaster, mentioned in section 13.14, in the
following respects: Posterolateral petal smallest in
both the genera; ratio of posterolateral:
anterolateral petals is 1:3 in Schizaster, 2:3 in
Hemiaster. Maximum height is at the posterior end
in Schizaster, slightly behind centre in Hemiaster.
Appendix 2: Lab Exercises 399
Subpetaloid unpaired amb is deeply incised in
Schizaster, not so much incised in Hemiaster.
Temnopleurus is a radially symmetrical regular
echinoid. Unlike, Cidaris (see section 13.14), it has
uniformly expanding straight ambs with compound
plates and multiserial pores.
A2.12 Trilobites: Examples
Well-preserved trilobite specimens are extremely
rare and so laboratories may not show enough
confidence to let beginners handle them freely for
description and other studies. Nevertheless, there
should be some idea about how to look
systematically at these fossils so important in
Palaeozoic.
Calymene, Phacops, Redlichia, etc. are Indian
forms, students are familiar with them at least by
their names. Here they are introduced with brief
descriptions.
Calymene is a posteriorly narrowing oval
shaped in the outline of the carapace; cephalon
is semicircular; glabella distinctly narrowing
towards front, with a few incomplete lobes;
eyes small, crescentic (holochroal); facial suture
gonatoparian; genal angles rounded; thorax
with some 11-13 segments, rounded at ends;
pygidium rounded and smaller (carapace
micropygous).
Carapace of Phacops is broadly elliptical in
outline; cephalon semicircular; glabella broadens
anteriorly and extends beyond the cephalon
margin; eyes very large and schizochroal; facial
suture proparian; genal angles rounded; thoracic
segments about 11, rounded at ends; pygidium
much smaller (micropygous).
Redlichia is elliptical in outline; cephalon
semicircular; glabella narrow towards front and
lobed; eyes long; facial suture opisthoparian; genal
angles drawn out into long spines; thoracic
segments and pygidium also with spines; carapace
micropygous.
For plant form-genera see Factsheet 19.17.
400 Appendix 2: Lab Exercises
A2.13 Mammalian Molar Teeth
In spite of commiting in section 18.1 that there
will be no detailing of vertebrate anatomy or
morphology or classification in this book, the
author takes liberty to introduce a few words on
mammalian molar teeth. In justification of this
breach, these are the following reasons:
1. Mammalian teeth are often very common and
useful fossils for the group. Unlike reptilian
teeth, mammalian teeth are differentiated in
morphology and function along the jaws. Thus,
there are four types of teeth, viz. incisor,
canine, premolar and molar. Of them, the last
one is particularly worthy, because of its
following attributes.
(a) Broadly, mammalian molar teeth are made
of three major kinds of substances: a soft
dentine [that makes the main volume of
teeth and has cells ‘rarely encapsulated
within mineralized tissues’ (Benton 2005)],
covered by hard enamel and a cement of
lesser abundance and importance and
filling some space between the
components of teeth. The hard enamel
outer covering added to these relatively
small sized parts of the body (in regard to
the size of the body of these animals)
renders preservability to the teeth.
(b) Molar teeth vary in their morphology and
that variation is closely linked with the
food habit. Since different larger groups
(orders and families) have typical food
habits, their characteristic molar teeth help
in the identification.
So, mammalian molar teeth being small,
externally hard, functionally significant
and morphologically distinct have turned
out as good fossils for these large land
animals.
2. To understand the morphology of teeth, the
following basic facts are needed as background
material:
(a) Molar teeth occur at the posterior end of
the dental battery along the jaw; the mouth
opens towards the anterior of the jaw and
teeth.
(b) Teeth of upper and lower jaws differ in
morphology. This is primarily because,
while grinding or chewing (mastication)
the components of the teeth (cusps and
cusplets: explained later) must fit with
each other or occlude to grip and crush
the food material firmly between them.
(c) Set in a jaw, a tooth has an outer side
towards the lip or mouth; hence it is called
outer/labial/buccal side. The opposite side
towards tongue is inner or/lingual side.
(d) Roots hold a tooth firmly in the jaw; the
rest is the crown. The crown surface,
upper surface for a lower jaw tooth and a
lower surface for an upper jaw tooth is
the grinding surface.
Thus, the three-dimensional body of a tooth has
six sides or directions: anterior, posterior, labial,
lingual, crown/grinding surface and roots. To
determine whether a tooth is of right or left jaw, we
need to fix up the first four sides of the above six.
3. Mammalian molars (particularly in placentals,
marsupials and their extinct relatives ; for
monotremes and relatives, the scheme is
slightly different) have a ‘tritubercular’
(from three primary tubercular components)
or rather ‘tribosphenic’ (meaning rubbling
wedge) arrangement. The three primary
tubercles or cones (also called cusps; a cusp
is conical when unworn; smaller cusps or
cusplets or conules, between major ones make
the tooth solid) form an inwardly pointing
triangle in a upper jaw molar fitted, rather
occluded in a basin like talonid in the lower
molar, between an outwardly pointing triangle
of three cones and a posterior extension of
talonid. The primary cones in the upper jaw
are protocone (anterior, lingual/inner),
paracone (anterior, labial) and metacone

(posterior, labial). Those of the lower jaw are
protoconid (anterior, labial), paraconid
(anterior, lingual) and metaconid (posterior,
lingual). Talonid of lower molars have, in
addition, entoconid (lingual), hypoconid
(labial) and hypoconulid (median, posterior).
Additional cones and conules tend to make the
upper molars assume a squarish cross-section; thus,
protoconule and hypocone on the lingual side to
the anterior and posterior of protocone
respectively, make the upper molar of equids
(e.g. Hipparion or Equus) squarish. Lower molars,
on the other hand, are generally elongate
rectangular/elliptical/oval from their talonids.
In more primitive forms and in omnivorous
mammals like primates including man, or pig-
family (Sus, Listriodon), the unworn teeth (during
lifetime of an individual, i.e. ontogeny, teeth suffer
wear with mastication) retain the separate, conical
shape of their cones. Thus, the crown surface is
studded with a few conical rises.
However, with evolution and for herbivorous
animals that require greater extent of mastication,
molar teeth underwent changes. The first notable
change is the formation of many smaller conules
in between the cones: to start with, the latter
retaining their distinct separate entity. But with
more complexity, adjacent cones and conules are
fused together with their enamels coalescing. The
two extremes may be found respectively in
specialized grazers and forest dwellers that feed
upon large trees, their branches and leaves. The
first, including the equids, develop a complex
enamel pattern with their major cones joined to
adjacent ones by conules.
The second extreme is found in proboscids,
where a linear series of numerous cones and
conules are fused together (to a varying degree in
different genera) to form a ridge or loph (a loph
which is a ridge when unworn is actually the
expression for a dental plate on the crown surface).
In worn out teeth, lophs are represented by a
continuous elongate closure of enamel with
wrinkles suggesting the original cones/conules.
Appendix 2: Lab Exercises 401
These lophs, rather dental plates are introduced in
a tooth at the posterior end, often as smaller half-
formed loph called half-loph, which then grows
during ontogeny to a full-formed loph. The entire
aggregation of these strong dental plates makes
the tooth robust and strong.
Depending on these features molar teeth are
termed bunodont (cones separate and distinct:
Homo), selenodont (cones are crescentic and
distinct; Hippopotamus), lophodont (cones fused
into distinct lophs: Elephas, Stegeodon); other
different combinations are selenolophodont,
bunoselenodont, etc.
4. A few other necessary terms are :
(a) Singulum is a thickening or fold on lateral
sides of teeth that mark the root from the
crown portion.
(b) Hypsodont is a tooth with crown portion
is longer/higher than root; such a tooth is
necessary for grazers in which teeth suffer
much wear and tear as the animal grinds
soft grasses with relatively harder soil
material mixed with them.
(c) Brachydont is a tooth with crown shorter
than root; soft or succulent plant or leaf-
eating browsers can do with this type
of teeth.
(d) Upper jaw teeth often bear some strong
ridges called styles on the labial surface,
not to be found in lower jaw teeth or on
ligual surface of upper jaw teeth.
On this background knowledge, a few
examples are provided here. They are not
exhaustive, but are meant to introduce to students
how molar teeth description may be handled.
Equus: Upper jaw molars are long, slender
prismatic in shape with four flat walls and a
squarish cross-section; crown is much higher than
root; hence tooth strongly hypsodont, suggesting
advanced grazing habit; roots four in number, may
be mutually joined.
Crown surface in worn out teeth is flat and
sloping inwards; enamel pattern is highly complex
with five major cones, of which protocone marks
402 Appendix 2: Lab Exercises
lingual and anterior; protocone is attached to the
adjacent cones by small necks of enamels.
Outer or labial surface with three distinct styles
separated by broad valleys. Inner, lingual surface
has two grooves corresponding to the limits of the
protocone.
Hipparion, the only other equid whose fossils
are obtained from India differs from Equus on
being shorter and, hence, stouter prismatic, and
having a protocone which forms a closed ring,
detached from adjacent cones.
Rhinoceros upper molar is also squarish in
section, but short prismatic in shape. Its crown is
shorter than root, suggesting brachydont nature and
browsing habit. Labial surface has broad styles;
lingual one has curvatures corresponding to the
transverse lophs.
Crown surface in a worn out tooth shows three
lophs, one along the labial margin (ectoloph) and
two tongue-shaped transverse lophs ending in
broad rounded curves on the lingual side (anterior
protoloph and posterior metaloph); the two latter
are separated by a sharp valley.
The angle subtended between the ectoloph and
any transverse loph points to anterior; since
protocone cannot be distinguished readily, this
criterion may be used to fix right or left jaw
position of the tooth.
Stegodon is generally seen in lower molar
which has an elongate subelliptical section and a
columnar shape; tooth brachydont; labial and
lingual surfaces unmarked.
Crown surface in unworn tooth shows six and
a half upwardly arched well-formed lophs made
of numerous cones and conules fused together; they
are separated by sharp v-shaped valleys; hence, the
tooth is typically lophodont.
Elephas has similar characters; however, it has
nine lophs on the crown surface. Gomphotherium
or Trilophodon is a primitive proboscid in which
the lophodont character is not as well-developed
as in Stegodon or Elephas. Here there are three
lophs each made of two major cones, one at each
end and a number of conules set in between, all
still retaining their original conical shape. Conules
are also there in the valleys.
Appendix
3
Indian Stratigraphy
for Palaeontologists
Fossils are preserved in stratigraphic successions.
Hence, knowledge about Indian fossils may not
be complete without an idea about the Indian
stratigraphic successions from which they were
collected. That itself is a vast subject, which cannot
be negotiated in details in the span of a chapter.
Thus, a brief overview of the Indian Phanerozoic
stratigraphy is added here as the substitute.
Tectonics and climate are considered two
important controlling factors in the development
of regional stratigraphy. Tectonic determines the
regional setting in which basins form and evolve,
whereas climate controls, mainly, the character of
sediments and biota.With plate tectonic gaining
ground, it has been possible to offer an explanation
of changing global dispositions of continents and
oceans, as well as location and development of
sedimentary basins of different tectonic regimes.
Regional stratigraphy of any part of the world may,
thus, be understood best on the background of plate
tectonics.
Indo-Pak subcontinent, though developed
geographically in a broadly unified manner, has
different geologic-stratigraphic history for its
different parts. The three broad physiographic
403
divisions are significant enough to understand the
stratigraphic history of this subcontinent. Thus, the
main southern Indian land mass, geographically the
Peninsular India, has a geological history different
from that of the other two parts-the Extra-Peninsula,
the northernmost and the Indo-Gangetic alluvial
plain that lies in between the Peninsula and the
Extra-Peninsula. The Peninsular India is the most
ancient part of the three and developed as a cratonic
land mass right back in the Precambrians. Towards
the end of this era, the Cuddapahs, the Vindhyans
and their equivalents developed in marginal seas.
As it is well-known, the Peninsular India does not
record any stratigraphic unit of worth mentioning
for the age between Cambrian and Carboniferous
(there is a controversy whether the upper age limit
of the Vindhyans should be above the Precambrian-
Cambrian boundary). The huge sequence of the
Gondwana Supergroup(/Group) developed since
Lower Permian (Sakmarian: earlier opinion that the
Gondwana Supergroup started from Uralian/ Upper
(Up.) Carboniferous is not generally accepted now-
a-days, though, of late, Veevers and Tewari (1995)
have reiterated Up. Carboniferous age) on this
stable land mass activated during Up. Palaeozoic
404 Appendix 3: Indian Stratigraphy for Palaeontologists
to give rise to the development of faulted basins,
rather half grabens, which were continuously
affected by synsedimentational subsidence to
accommodate several thousand metre thick
sequence of mainly fluvial clastics. Till then the
Indian land mass, rather the Indian plate was a part
of the Gondwana supercontinent; this has left
imprints on the development of similar types of
coal-bearing continental deposits, a common
Gondwana flora as also a distinct amphibia-
reptilian fauna typical in the late Palaeozoic-early
Mesozoic of the southern continents (Chapter
18 and 19 for details).
Subsequent history of the Peninsula involved
rifting of the Indian plate from the Gondwanaland
supercontinent (or rather Pangaea) sometime
during Triassic and its drifting first northwards and
then northwestwards since mid-Mesozoic.
Immediate result of these events was the
development of the Peninsular India in its present
configuration along the trailing boundary of the
Indian plate. Obviously, this was marked by the
formation of marine sequences of late Mesozoic-
Cenozoic age in different basins along the newly
formed coastline of India. The latter did not form
all at once and so the basins did not originate all at
one time.The earliest example of these sequences
come from Kachchh at the northern tip of the west
coast where the marine sequence starts from
Middle Jurassic (Bathonian: age fixed on
ammonoids e.g. Macrocephalites triangularis and
M. macrocephalus). Slightly later, since Callovian,
there developed a smaller succession rather inland
in Rajasthan (Jaisalmer basin) and that too at its
northern tip. The sea transgressing these newly
formed marine basins must have come from already
existing southwestern arm of the Tethys that
extended into the Sind-Baluchistan regions of
present-day Pakistan, containing the basins of Salt
Range in addition to those of Sind and Baluchistan.
Geologic history of Kachchh reveals that marine
sedimentation continued in the western part of
mainland of Kachchh peninsula into Lower
Cretaceous when it was ultimately interspersed
with continental sedimentation of the equivalents
of the uppermost Gondwana Supergroup, viz., the
Umia Plant Beds. In the northern islands and the
eastern Wagad Highland of the Peninsula, only the
lower part of the Mesozoic succession is recorded
indicating that the sea receded from those areas
gradually westward with the upliftment of land
centering round a point at the junction of Kachchh,
Rajasthan and Cambay basins (Therad High)
(Biswas 1971). This was again followed by
subaerial volcanics of the Deccan Traps. In most
of the other coastal basins, such as Cambay basin,
Saurashtra-Kathiawar basin, Bombay Offshore
basin and Quilon basin of south-western coast,
marine sedimentation took place only after the
Deccan volcanism, though some of the basins like
the Saurashtra basin might have developed in
Cretaceous when they recorded uppermost
Gondwana unit or their equivalents like Wadhwan
Sandstone, Himmatnagar/Ahmednagar Sandstone,
etc. that contain Wealden (Lower Cretaceous) floral
elements like Weichselia or Onychiopsis.
The Kachchh sequence is particularly rich in and
so well-studied for ammonoids (macrocephalitids,
perisphinctids, etc.), brachiopods (rhynconellids,
terebratulids, etc.), corals, bivalves (including
Trigonia vetricosa and T. crassa). In Cenozoic,
sedimentation continued on west coast in offshore
basins, whereas onshore, Kachchh and Cambay
basins had less prominent Palaeogene-Neogene
successions (though highly fossiliferous with
well-developed foraminiferal limestones as also
different horizons of echinoids and corals, etc.) and
Saurashtra basin has the well-known Miliolite
Formation of Quaternary age.
Another important basin in western India lies
along the present-day Narmada valley and Satpura
regions. It formed sometime during upper
Mesozoic when continental Upper Gondwana
sediments were deposited (Lower Cretaceous).
Thereafter, a Turonian-Campanian marine
succession with almost equivalent continental
lateral facies variation is found in the Bagh and
Lameta Beds. They indicate an Upper Cretaceous
 Appendix 3: Indian Stratigraphy for Palaeontologists
marine transgression in this part which probably
came from the west. It was, however, followed by
the Deccan volcanism in the uppermost
Cretaceous.
East coast basins developed with a slightly
different history. The most prominent of them are
the Cauvery basin and its adjacent ones which
recorded Upper Gondwana sediments (Sivaganga
Fmn) to be followed by marine deposits of
Neocomian or Middle Cretaceous transgression.
This continued through later part of Cretaceous
into Palaeogene, though younger sediments above
Palaeocene are found only in subcrops. During
Neogene or Mio-Pliocene, the basin witnessed
regression which produced deposits like Cuddalore
Sandstone. Equivalents of this last named unit are
found scattered in different areas along the east
coast in Andhra Pradesh (AP), Orissa, etc. Another
significant development in these east coast regions
particularly in AP is the close interfingering of
marine and continental deposits, more commonly
known as coastal Gondwanas. These include well-
known Raghavapuram Mudstone with important
marine fossils like ammonoid, foraminifers and fish
(Clupavas neocomensis) marking its Neocomian
age, intercalated with unfossiliferous or Gondwana
plant fossil-bearing sandstones. It is obvious that
in the east coast, land-sea marginal condition
prevailed since Upper Cretaceous time.
Further eastward in Bengal basin too marine
transgression did not take place before Upper
Cretaceous which is evident from the presence of
uppermost Gondwana equivalents and Rajmahal
Trap equivalents (110 Ma; some authors consider
them as earlier phase of the Deccan Traps; age
Lower Cretaceous; effused subaerially) . Upper
Cretaceous marine units are also present in
different basins of Assam-Arakan region, which
occur in the north and north-eastern limits of
Bengal basin. Bengal basin encloses marginal
sediments grading into deeper marine equivalents
during Palaeogene. Towards the end of this time,
the basin started closing in a zipper-like manner
from northeast to southwest. As a result, since
405
upper Palaeogene, different basins in the north-
eastern India started to house continental deposits
equivalent to the Siwalik Group of Formations.
As compared to the west coast basins, the east coast
basins contain younger sediments and more
prominent development of Neogene continentals.
History of extra-Peninsular India and their
continuation in the present-day Pakistan (mainly
in Salt Range, Sind and Baluchistan) is entirely
different for obvious reasons. The whole area was
occupied by the Tethys since Cambrian time, which
was the receptacle of huge amount of sediments.
Different basins there, however, have different
history. Salt Range, one of the most important
basins, houses Cambrian deposits that end in the
Saline Series. There is a long gap from Ordovician
to Upper Carboniferous (Uralian) after which there
formed the Eurydesma-Conularia beds of
lowermost Permian age. The same fossil
Eurydesma is said to occur in the marine beds at
or near the base of the Gondwanas, particularly in
central Indian occurrences of Umaria,
Manendragarh, Anuppur, etc. Salt Range basin
experienced vigorous marine transgression in
Permian; as a result there developed Productus
limestone. In addition to the rich fossil record of
orhtid, productid, strophomenid and other
brachiopods, anthozoans, bryozoans, etc. this
limestone also presents earliest representative of
ceratitid ammonoids like Xenaspis carbonaria. The
overlying Ceratites beds present more advanced
ceratitids. Marine sedimentation in Salt Range
continued till Oligocene. Bugti Beds and overlying
Murree Formation are evidences of end of marine
sedimentation in this basin. These units, and
particularly Bugti Beds contain large land mammal
fossils which were precursors of the Siwalik
mammals (Chapter 18).
Salt Range area also includes the Potwar
Plateau, which is considered as the type area of
the Siwalik Group of Formations. Thus, from
Middle Miocene to Lower or Middle Pleistocene,
there developed a thick succession of continental
sediments of ‘molasse’ type with rich mammalian
406 Appendix 3: Indian Stratigraphy for Palaeontologists
fauna enclosed in it. This was terminated by a cold
arid phase with evidence of glaciation found in the
topmost units of the Siwaliks, namely, the Boulder
or Tawi Conglomerate. The mammalian fauna
became largely extinct or its elements migrated out
of this land to either Africa or Far East countries.
(The sudden appearance of rich mammalian fauna
in Oligocene of this subcontinent may also have a
plate-tectonic link. It was in OliogoMiocene times
that the African plate was welded with the Eurasian
plate as a result of which the rich African fauna,
including primates, spread out to the Eurasian
lands; reported pre-Oligocene Indian mammals
mostly include Lower to Middle Eocene marine
forms from Kachchh and some other areas).
Equivalents of the Siwaliks are present in the
Indian territory (see Chapter 18).
The other two important extra-Peninsula
basins, namely, Spiti and Kashmir are among the
Himalayan basins. Of these, Spiti recorded a
virtually continuous marine succession from
Cambrian to Cretaceous. It includes the following
well-known units (arranged from top to bottom)
like Giumal Sandstone and Chikkim Series (Lower
and Upper Cretaceous, respectively), Spiti Shale
(Jurassic), Lilang System (Triassic), Conglomerate,
Calc. Sandstone, Productus Shale (successively
during Permian), Po Series (Middle to Upper
Carboniferous), Lipak Series (Lower to Middle
Carboniferous), Muth Quartzite (Devonian/Siluro-
Devonian), Ordo-Silurian beds and Haimanta
System (Precambrian-Cambrian) .
Though many of these units recur in Kashmir
basin, the history and fauna of the latter are
different at many points. Thus, during late
Permian, Kashmir basin records subaerial
volcanics of the Panjal Traps with Agglomerate
Slates at their base and Gangmopteris Beds (now
called Mamal Formation) of continental origin and
Zewan Limestone of marine origin above the Traps.
The difference in fauna is most evident during
Cambrian, when the Salt Range fauna was
characterized by Redlichia noetlingi, an index
trilobite species of Middle Cambrian age found
also in Australia and other parts of east Asia. They
are associated with primitive brachiopods,
particularly Neobolus. It has also the trilobite
Ptychoparia, whose species are different from
those found in Spiti. The Spiti fauna, on the other
hand, has trilobite as the main constituent with
Redlichia noetlingi also present there; there are also
primitive brachiopods without Neobolus and a
probable Olenus, an Upper Cambrian trilobite.
Ptychoparia is represented by P. pervulgata, P.
consocialis, etc. Both these basins further contain
some pteropods like Hyolithes.
The Kashmir fauna is comparatively largely
local and endemic. The more cosmopolitan forms
that are found in Kashmir Cambrian have affinity
to Tonkin fauna of China, than the equivalent
Australian or other Far East faunas. These include
trilobites such as Tonkinella, Chuangia,
Anomocare. The basin is also marked by the
striking absence of Redlichia and primitive
brachiopods of Salt Range or Spiti types; most of
the species of Ptychoparia and Hyolithes are
different.
The difference was, however, largely lost
during Middle Palaeozoic, but was again
prominent in Carboniferous in particular. The
Kashmir basin records one of the few occurrences
of extra-Peninsular Lower Gondwana with plant
fossils and more particularly fish and amphibian
fauna (mammal Formation : see Chapter 18).
Continental condition is also evident from the
subaerial basic volcanics. The basin was again
transgressed during Upper Carboniferous-Lower
Permian and since then it recorded successions
comparable and correlatable with those of Spiti,
till Cretaceous. Extra-peninsular marine
sedimentation virtually stopped during Cretaceous
and in all likelihood it had some bearing with
initiation of the Himalayan Orogeny.
The Himalayas may be subdivided from north
to south into Lesser, Great/Central, Tethyan/
Transaxial Himalayas. All these belts virtually
continue from west to eastern end of the mountain
range. They have different stratigraphic histories
 Appendix 3: Indian Stratigraphy for Palaeontologists
and are mutually juxtaposed tectonically (Ghosh
2000). They include Neogene and Quaternary
sediments of the Lesser Himalayan basins
terminating to the north by the Main Boundary
Thrust against the pre-Tertiary rocks of the Greater
Himalayas. These are correlated to the Siwaliks.
Some of the Lesser Himalaya belts in eastern India
developed Miocene brackish water sediments. Pre-
Tertiary low grade metamorphic rocks of the
Greater Himalayas, floored by the Main Central
Thrust, tectonically underlie the central crystalline
belt. The Greater Himalaya succession essentially
consist of Palaeozoic sediments and unfossiliferous
low-grade metasediments probably of Proterozoic
age. Metamorphic and granitoid rocks of the
Central crystalline belt or parts of Lesser Himalayas
were affected by Proterozoic metamorphism and
igneous activity. Fossiliferous marine Eocene
sediments occur sporadically in intimate tectonic
juxtaposition with continental to paralic Permian
sediments as a melange zone along the Main
Boundary Thrust. The Lesser Himalayan rocks are
associated with basic and acid volcanics and
volcanoclastics which sometimes contain
infratrappeans of lower Gondwanas in Northeast
India (Abor volcanics) or in Kashmir. In the
Transaxial Himalaya, Phanerozoic metasediments
overlie the crystallines. Those successions
represent a nearly complete Phanerozoic
epicontinental fossiliferous sediments of Tethyan
affinity with a few minor breaks.
Stratigraphy and its interpretation for these
tectonized belts obviously depend on an adequate
understanding of plate tectonics. This is also
evident from the stratigraphic development of
basins in Assam-Arakan Yoma regions and in the
Bengal basin.The former includes broadly two
kinds of sedimentary associations or facies, one
of shelf and the other of deeper ocean basins. The
Bengal foreland basin located between the eastern
edge of the main Indian craton and the Indo-Burma
Range and lying to the south of the Shillong plateau
is really an asymmetric depression on a crust
sloping easterly. The basin is a remnant ocean basin
407
formed due to northwestward drift of the Indian
plate and its oblique collision with the Sino-Tibetan
and Burmese plates. The basin progressively closed
in a zipper-like manner from north east to south-
east with gradual filling up by sediments (flysch)
derived from rising mountain belts nearby. It is,
thus, marked by gradually receding sea from north
to south and consequent marine to marginal
freshwater sedimentary facies developed in the
same direction of the receding sea. The basin was
created essentially during late Cretaceous
succeeding a volcanic episode (see Biswas 1971).
It is clear from this brief, rather cryptic,
overview that:
1. Different regions of India and adjoining
countries, often referred to as Indo-Pak-
Burmese subcontinent, show different tectonic
and stratigraphic development. Not only they
differ between Peninsular and extra-Peninsular
India; they are different even within each of
these two regions. Correlation between these
regions is, thus, problematic at different points.
2. To correlate different basins in these regions,
even the presence of stratigraphically
important biota of marine origin is often not
enough. Different basins or parts may have
been palaeogeographically and palaeobio-
geographically linked to different provinces.
In extra-Peninsula, the problem is
demonstrated in the Palaeozoic part of the
successions of Spiti, Kashmir and Salt Range.
Thus, even Spiti and Kashmir show Cambrian
faunas that may have affinities with two
different faunal provinces. Similar problems
are there with Peninsular occurrences too, for
instance in regard to correlation of Middle to
Upper Cretaceous marine units of central
India, viz. the Bagh Beds and the Lameta Beds.
These have been correlated by some authors
with Madagascar and East African
successions, particularly on the basis of the
dinosaurian fauna of the Lametas, e.g.
Laplatasaurus madagascarensis, etc. On the
other hand, on the basis of echinoid and other
408 Appendix 3: Indian Stratigraphy for Palaeontologists
invertebrate fauna, Bagh Beds are correlated
with Mediterranean province (Krishnan 1968).
3. A third problem of a different kind exists, for
example, in connection with the Siwaliks. The
rich mammalian fauna of these continental
deposits includes many genera and species
present in other continents too. Of these,
particularly significant are Hippaion and Equus,
the two equid of Neogene and Quaternary,
respectively. Since, there are no other equid
genera in the Siwaliks and further, the two
are interpreted as belonging to two different
lineages from the phylogeny reconstructed
mainly on North American records, the Indian
genera are considered immigrants from North
America via a land-bridge connection across
the Bering Strait and Eastern China. Obviously,
the age of the Indian genera should be slightly
younger than their North American counterpart,
considering the time of migration.
This migration is considered as a part of a
North-American-Eurasian-African mammal
exchange, the so-called Hipparion event. This, in
its turn, developed as a result of two global
palaeogeographic changes. The first is the
Eurasian-African collision which permitted the
first major mammal exchange between these
continents in the Neogene. The second involves
the changing seaways during the Miocene
including a closure of the Mediterranean-Indo-
Pacific seaway (related to the Messinian event).
New migration routes were thereby established
leading to the so-called Hipparion event.
It may be surmised at the end, that
notwithstanding controversies in freaking age of
fossils and their host units or differences in opinion
in interpretation of successions of different basins
and regions, the brief overview above will help
provide a general and overall idea on which the
controversies may be assessed.
 Epilogue

The book is a partial representation of the subject. A lot is left to be discussed about each of the issues
and the organic groups covered. So, there will always remain the risk of facing the criticism that they
are underrepresented. Yet one has to stop somewhere and with a sizeable portion of the target readers
being the beginners, it is felt enough was enough. However, the author cannot avoid ending his discourse
without quoting two expressions on work-experiences that readers of this book may find worth sharing.
1. The Challenger Expedition was undertaken in 1872-76 for global biological collections and to
study existence of life in deepest waters. Thousands of samples, 715 new genera and 4417 new
species of marine organisms were collected. “ Never did an expedition cost so little and produce
such momentous results for human knowledge”, Commented Ray Lancaster. Alexander Agassiz,
the famous scientist worked on the monograph on echinoderms, on samples collected during the
Expedition. On finishing his assignment, he remarked:
I felt when I got through that I have never wanted to see another sea urchin and hoped they would gradually
become extinct. (Lalli and Parsons, 1997)
2. In a voyage to the Antarctic waters on board Discovery I , Alister Hardy narrated:
Expecting a host of surface life we slung a bo’sun’s chair (a board supported by ropes on each side like a
swing) close to the water… right in front of the bows themselves. Here Kemp and I took turns with a hand-
net and bucket. For sheer pleasure it was ideal; swinging in mid-air and gently rising and falling with swell
over the deep blue surface which occasionally rose to bathe and cool one’s legs; one advanced like a gliding
and soaring bird with nothing in front of one but the virgin ocean, as yet quite undisturbed by the bows
behind… . I rode in triumph, fishing out treasure after treasure as they came floating towards me on the very
gentle undulating swell. An experience never to be forgotten (Lalli and Parsons, 1997).

409
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 Index
Aboral, 213 Arthropoda, 231
Abyssal, 35, 37 Articulata, 146, 149
Abyssopelagic, 37 Articulating,
Acanthothyris, 159 furrow, 236
Accretion, 11, 210 half-ring, 236
Adaptation, 38–40, 174 Asaphid, 240
Adaptive radiation, 74, 75, 86, 113, 176, 206 Ascon stage, 265
Addition, 11 Asterozoa, 212
Adradial, 221 Athyridida, 157
Agglutinated test, 258, 263 Atrypa, 37, 392
Allochthonous, 44, 253 Atrypida, 157
Allopatric speciation, 64 Autochthonous, 44
Amaltheus, 188 Autotrophs, 34, 37
Ambitopic, 159 Axial,
Ambitus, 214 column, 135
Ambulacrum (-a, pl), 210 furrow, 237
Ammonitida, 204 ring, 236
Ammonoidea, 186, 187 structure, 11
dimorphism, 199 vortex, 135
palaeobiogeography, 199
suture, 196
Ampulla, 211 Baltic amber, 384
Anarcestida, 203 Basal, 81, 128, 262
Ancyloceratida, 197, 204, 209 Bathyal, 35
Anomphalous, 183, 394, 395 Bathypelagic/aphotic, 37
Antagonism, 35 Bathyurid, 240
Anthozoa, 71, 125, 127, 135, 137, 389, 390 Beak, 153, 163
Antibiosis, 35, 43 Benthic/benthonic, 37
Aperture, 188, 258, 262 Binomial system, 55
Apical, 179, 213 Biological
disc, 213 environment, 34
Aragonite tubes, 193 species concept, 60–62
Aristotle’s lantern, 226 Biostratigraphic zone (biozone), 67, 69
423
424 Index

Biostratigraphy, 66–68, 71 Conchiolin fibres, 193

Biostratinomy, 8, 14 Concretions, 382
Bivalve, 171, 172 Concurrence, 35
Bivalved, 140, 162 Connecting ring, 187, 194
Bivalvia, 140, 162 Cope’s rule, 205
adaptation, 171, 174 Coral,
Blastozoa, 212, 213 ahermatypic, 129
Body whorl, 179 hermatypic, 138
Boreal realm, 200 Corallite, 128
Boring, 37 Corallum, 128
Bositra, 378 Corona, 212
Bourrelets, 226 Costae, 157
Brachial, 47, 150 Costellae, 157
Brachidium, 50, 156, 159, 160 Craniiform, 156, 157, 159, 160
Brachiopods, 140, 150 Cribrate, 263
Brevicone, 188, 395 Crinozoa, 212
Bryozoa, 140 Cryptomphalous, 183
Burgess shale, 372 Cryptopora, 37
Burrowing, 37 Cryptorhynchia, 158, 160
Byssal notch, 173 Ctenophora, 125
Cyrtocone, 188

Cadicone, 188
Calceola sandalina, 138 Decomposers, 35
Calcichordates, 211 Delthyrium, 154
Calice, 128 Deltidium, 154
Calliphylloceras, 202, 203, 205 Denticles, 238
Calymenacean-dalmanitacean, 240 Dentition, 164
Canals, 211 Depressed, 188
Carapace, 233, 234 Desmodont, 168
Carpoids, 210 Dextral, 178, 188
Cenogenesis, 207, 208 Diagenesis, 14
Cenosteum, 265 Dicyclic, 227, 228
Cephalon, 233, 234 Diductor, 155
Cephalopoda, 140, 186, 203 Dikelocephalinid, 240
Ceratitida, 204 Dimorphism, 199
Chance, 60 Diploblastic, 125
Chilidium, 154 Discoidal, 236
Chondrophore, 162, 170, 173 Dissepiments, 132
Chonetes, 159 Dysodont, 168
Cladistics, 61
Clymeniida, 203
Cnidaria, 125 Earthbound mechanism, 78
Coelenterata, 125 Ecdysis, 11, 12, 232
Coiled shells, 140 Echinozoa, 212
Coleoidea, 140, 186 Echinodermata, 210
Columella, 135, 183, 190 Echinoid, 37, 40, 217, 225, 228, 229
Commensalism, 35 test, 212
Community, 34, 43, 45 Ecological
Compass, 226 niche, 34
Composita, 47 species concept, 61
 Index 425

Ecology, 32 Foramen, 154, 194, 258

Eco-morphotypes, 172 Fossilization, 13–15
Ecosystem, 34 Fossils, 3
Ectoderm, 125 aspects, 6
Edentate, 165, 168 assemblage, 33
Eleutherozoa, 212 definition, 3
Endobenthic, 37, 141 guide, 68, 69
Endocyclic, 227 index, 68, 69
Endoderm, 125, 138 kinds of, 9
Endopetalous, 225 living, 77
Enteron, 125, 126, 132, 265 micro, 245–256
Entocoel, 134 Fossula, 135
Entosepta, 134 Free cheeks (librigena), 233
Epibenthic, 37, 143 Fulcra (-um, sin), 153
Epibiosis, 35
Epipelagic, 37
Epiphyses, 226 Gastropoda, 54, 140, 142, 146, 176–185
Epitheca, 132 Gelatinous sheath, 156
Evolution, 74, 75 Geological time scale, 5
bradytelic, 77, 208 Gills, 141, 177
explosive, 75 Glabella, 233
horotelic, 77 Goniatitida, 203
iterative, 206 Gonopores, 227
macro, 74 Gregarious, 211
micro, 74, 80 Growth lines, 132
of ammonoid, 203 Grube messel, 382
of equidae, 86 Gryphea, 37
of hominidae, 91 Guard, 398
of proboscidea, 81
organic, 74, 77, 79, 80, 113, 118, 119, 121
parallel, 78 Habitats, 34, 35
tachytelic, 77 Habits, 34, 35
Exocoel, 134 Hadal, 35
Exocylic, 227 Haeckel’s law, 11
Exosepta, 134 Hallucigenia, 49
Exploitation, 35 Halysites, 135, 138
Extinction, 178 catenularia, 138
Extraterrestrial mechanism, 78 Hercosestria, 47
Heterochronous, 159
Heterochrony, 74
Facet, 236 Heterodont, 168
Favosites, 135 Heteromorphic, 186
Fascioles, 224, 225 Heteromorphy, 196
Feeders, 37 Heterotrophs, 34, 37
Fixed cheeks (fixigena), 233 Hierarchy, 55
Fixosessile, 37, 149 Hinge, 149, 153, 163
Floscelle, 217 area, 165
Food groove, 216, 226 line, 149, 153, 163
Formation Holaspis, 11, 232
Morrison, 378 Holochroal, 235
Santana and Crato, 381 Holoplanktic, 37
426 Index

Holotype, 54 Linguliformea/linguliform, 140, 160

Holzmaden shale, 377 Littoraltidal, 55
Homalozoa, 212 Lobes, 196, 197, 233
Homeomorphy, 79, 159, 206 Longicone, 188
Homo sapiens, 6 Lytoceratid, 204
Homologous structure, 48, 49
Homology, 60
Homonym, 55 Macrocephalites, 205
Homoplasy, 48, 60 Macroconchs, 199
Hot Spring Miracle in Rhynie Chert, 373 Macropygus, 238
Hunsruck slate, 374 Madreporite, 215
Hunters/hunting, 37, 142 Mandible, 232
Hyaline, 258 Mantle, 141
Hydrocarbon generation, 254, 255 cavity, 141
Hydroida, 127 Marine environment, 51
Hydrozoa, 125–127 Mazon creek biota, 374
Hyponome, 141, 194 Median sulcus, 159
Medusa, 125
Megalospheric, 258
Iguanodon, 50 Meraspis, 11, 232
Imperforate, 258 Meroplanktic, 37
Impressed zones, 190 Mesentries, 125
Inarticulata, 149 Mesoglea, 125
Individual, 34 Mesopelagic/disphotic, 37
Interambs, 212–215, 217, 221 Metasepta, 134
Interarea, 154, 159 Metazoa, 125, 126
Interpleur al furrow, 237 Microconchs, 199
Intervallum, 265 Micropalaeontology, 245
Invertebrates, 11 Micropygus, 238
Involute, 179, 195 Microspheric, 258
Isochronous, 159 Mimicry, 60
Isodont, 168, 173 Molluscan body plan, 141, 208
Isopygus, 238 Mollusca, 10, 140, 162, 176, 186
Monocyclic, 227
Monophyletic, 63
Jointed appendages, 232 Monoplacophora, 74, 141
Morphological similarities, 60
Morphology, 8
Kutchithyris, 160 Moulting, 11, 12, 232
Mouth, 125, 141, 194, 213
Multilocular, 258
Labrum, 226, 229, 230 Muscles, 154, 168
Lamellibranchia, 140 adductor, 155, 168, 169
Lappets, 199, 204, 206 adjustor, 156
Lateral Mutualism, 35
chambers, 262 Mytilus, 37, 165, 168, 169
profile, 150
Lectotype, 54
Leucon stage, 265 Nautiloidea, 186
Liberosessile, 37 Nautilus, 48, 193
Librigena, 233, 234 Nektons, 37
 Index 427

Neoteny, 75 Pentameral organisation, 210, 211

Neotype, 54 Pentamerida, 157
Neritopelagic, 37 Peramorphosis, 75
Nomenclature, 53 Perignathic girdle, 226
Notothyrium, 154 Periostracum, 156, 157
Nova Scotia, 180 Peripetalous, 225
Periproct, 213, 215, 217
Perisphinctes, 188
Oceanopelagic, 37 Peristomial system, 213, 226
Oculogenital system, 213 Perradial, 221
Olenellid, 239 Petaloid amb, 223
Ontogeny, 11 Phaneromphalous, 183
Operculum, 135 Phanerozoic periods, 67
Opisthobranchiata, 182 Phragmocone, 11, 48, 193, 194, 398
Ornaments (surface ornaments), 157, 171, 184, 188 Phyletic gradualism, 65
Orthocone, 188 Phylloceras, 188
Osculum, 265 Phylloceratida, 196, 197, 204
Ostrea, 37 Phyllodes, 226
Oxycone, 188 Phylogeny, 10, 18
Phylogenetic
relationships, 63
Pachydont, 168 species concept, 62
Paedomorphosis, 75 systematics, 62, 81
Palaeoautecology, 33, 34 Pinacoceras, 205
Palaeoecology, 32, 33, 49 Planispiral, 178
Palaeontology, 3 Planktic (nektic), 37
Palaeosynecolgy, 33, 34 Planulate, 188
Pali, 135, 268 Plastron, 226
Palingenesis, 207 Plates, 210
Palintrope, 154 adoral, 221
Pallial adpical, 221
line, 170 chilidial, 154
sinus, 141, 170–172 demi, 221
Pallium, 170 genital, 213, 227
Paragaster, 265 hinge, 165
Parallelism, 60 ocular, 213
Paraphyletic, 63 Plenipedunculate, 154, 159
Parapuzosia sepperadiatus, 205 Pleuron, 236
Parasitism, 35 Plications, 157
Paratheca, 132 Podia, 211, 215
Paratype, 54 Polymerase chain reaction, 81
Parazoa, 125, 126 Polymerid, 233
Pedal scar, 170 Polyp, 125
Pedicle, 150 Polyphyletic, 63
foramen, 154 Polyplacophora, 141
opening, 154 Polytypic species, 61
Pedunculate, 154 Population, 34
Pelagic, 37, 162 Porcellaneous, 258
Pelecypoda, 140 Porifera, 125
Pelmatozoa, 212 Posidonia, 378
428 Index

Preservation, 13, 15 Septa (sin. septum), 42, 48, 128, 132, 134, 193, 258
Primary layer, 156 Septal neck, 187, 194
Proboscidactyla, 127 Septotheca, 132
Productus, 153, 391 Serpenticone, 188
Progenesis, 75 Sessile, 37, 145, 182
Prolecanitida, 204 Setae, 268
Prosepta, 134 Sicon stage, 265
Prosogastropoda, 54 Sinistral, 178
Protaspis, 11, 232 Sinupalliate, 170, 391
Proterogenesis, 207–209 Siphon, 141, 162, 182
Protoconch, 187, 193, 203, 204 Siphuncle, 42, 141, 193
Protozoa, 125 Sirius passet, 373
Proximal, 179, 287 Skeletons, 140
Pseudoceratitic, 197 Sockets, 155, 163
Pseudoplanispiral, 178 Speciation, 60, 64, 74
Pseudoplanktic, 37 Species, 34, 54, 55, 57, 60, 61
Pseudopunctate, 157 Sphaerocone, 188
Pulmonata, 54, 146, 177 Spheroidal, 236
Punctuated equilibria, 65 Spines, 210, 238
Pygidial segment, 233, 238 Spiral angle, 179, 394
Pygidium, 232, 233, 236 Spire, 179
Pyramid (hemi), 226 Spiriferida, 157
Spiriferinida, 157
Spirula, 194, 195
Quasi-infaunal, 159, 160, 391 Spiroceras, 200
Spongiomorphida, 127
Stegosaurus, 32, 311
Radial, 40 Stephanoceras, 184, 188
Radula, 141, 194 Stratigraphy, 66
Rancho la brea, 386 Stasipatric model, 64
Rectimarginate, 158 Stasis, 74
Redlichid, 239 Strophomenid, 157, 159
Resilifer, 170 Suspension feeders, 37, 141, 143, 171
Resupinate, 150 Suture, 193, 196, 204, 209, 234, 236, 258
Rhizopedunculate, 159 Swallowers, 37
Rhizosessile, 37 Symbiosis, 35
Rhynchonelliformea/rhynchonelliform, 146, 160 Sympatric speciation, 64
Roots, 135 Symplesiomorphic, 63
Rotula, 226 Synapomorphies, 63
Rugosa, 127 Synapticulae, 132, 135
Synapticulotheca, 132, 389
Synonym, 55
Saddles, 196 Syntype, 54
Scavengers, 34, 37 Syringopora, 138, 390
Schizochroal, 235 Systematics, 8, 52
Schizodont, 168
Scleractinia, 127
Scyphozoa, 125 Tabulae, 134
Secondary layer, 156 Tabulata, 127, 134
 Index 429

Taphocoenosis, 33, 44 Trilobite, 231–241

Taleolae, 157 enrolment, 236
Talon, 135 ontogeny, 232
Taphonomy, 13, 14 tracks and trails, 239
Taxodont, 168 Trochospiral, 177
Taxon, 54 Trophic chain, 34
Taxonomic Tube feet, 224
category, 54 Tubercles, 217, 224, 225
hierarchy, 6, 10, 56 Typological species concept, 54, 58–60
Taxonomy, 6, 8, 53 Typomorphic, 54, 59, 61
Teeth (tooth), 153, 163, 226
Temporal polymorphs, 126
Tendons, 156 Umbilicus, 183, 188
Tentacles, 141 Umbo, 153, 163
Terebratula, 37 Unilocular, 258
Terebratulida, 157, 365 Univalved, 140
Terrestrial,
and aquatic, 37
environment, 51 Vagrant, 37, 182
organism, 37 Valves, 140, 149, 163
Test, 210, 258 Vertebrates, 81
Tethyan realm, 200
Thanatocoenosis, 33, 44
Theory of recapitulation, 11, 209 Wall, 132, 258
Thorax, 233 Water-vascular system, 210
Titanites, 205 Whorl section, 143
Titanosaurus, 32, 311 Williamsonia sewardiana, 15
Tolerance, 35
Topotype, 54
Transformers, 35 Zooecium, 266
Trema, 182 Zooids, 272