AD Palaeontology

An Introduction
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AN INmC)DOCUON ro
PALAEONTOLOGY
. AN INTRODUCTION TO
PALAEONTOWGY
By
AMAL DASGUPTA, M.Sc., Ph.D.
Ex-Reader in Geology
Asutosh College
Kolkata
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•. . ) THE. WORLD PRESS PRIVATE LIMITED

KOLKATA · 1, • 2012 *
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PREFACE TO ·THE .SECOND EDITION

The text of ·this new ' edition is somewhat ·extended compared to that of the previous one.
However, in ·rnost of the case, the addition is kept minimum except Part V (Microfossils) where 1
foraminiferas, one of the most important group of microfossils , are discussed in more details
(including their skeletal morphologic features., their .ecologic :significance and stratigraphic
applications).- The application of microfossils · in petroleum :exploration work is also given here
in brief. This will make this book useful for post-graduate studenis ,where microfossils, especially
foraminifera occupy a substantial part of the palaeontology syJlabu~_.:This· is·also to be mentioned··.
that there were some mistakes left unnoticed in . the' previous edition. I- try to rectify them as
, far as possible. I also like to say this book, is written as a gener~l -. texr.:book on palaeontology
touching its major branches. It is written for the general students ·rather than ·for the specialists
or research workers. · ... , .- ·.
· The author offers his thanks ·to Sri -Subinial :Ghosh, :·Proprietor· of.j -Jindusthan Mineral. and
Natural History Specimen . Supply Co~ for .his :f6ssils 'ishow11 -: in cover photograph. ·
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PREFACE TO THE FIRST EDITION
During my thirty four years of teaching in undergraduate classes it appears to me that paucity
of literatures on palaeontology not only creates hardship to students in following up the class
room teaching but also they frequently lose attraction to this subject which otherwise constitutes
one of the most interesting branch of earth sciences. Of course, there are some excellent
traditional text books in palaeontology written by some eminent writers. Most of these books
deal with part of palaeontology and some of them are too upgraded and too much elaborated
to be managed by undergraduate students of Indian universities. Very often these foreign books
are not available in the market. A tood text book dealing with all the different aspects of
palaeontology suitable for undergraduate students is no doubt lacking till now. This has inspired
me to write this book. In this book I have tried to consolidate the scattered informations available
from different published books, journals and manuals which I have collected so far in my long
teaching days.
In the present book, the author intends to discuss the different aspects of palaeontology in
six parts. Part I deals with the different principles related to palaeontology. Hard part
morphology of some important groups of fossils invertebrates together with their subdivisions,
ecology and geological history have been discussed in Part II. A list of invertebrate fossils of
India is also given here. However, details of soft part morphology of different fossil invertebrates
is kept out of preview. Part III is devoted to vertebrate fossils that include their broad
subdivisions, skeletal morphology, different aspects of evolution and geological history of
vertebrates. Evolution of the horse, elephant and man has been discussed in two separate
chapters. The record of vertebrate fossils of India is also given in a separate chapter. Plant
fossils constitute the Part TV of the book where morphology, subdivisions, geological history
of plants and also an account of Indian plant fossils are given. Part V, comprising a single chapter
includes microfossils, their general account and skeletal morphology of some important groups.
The concluding or the Part VT covers some aspects of practical palaeontology where methods
of collection, preparation and description of fossils have been discussed in brief. For the benefit
of beginners, morp_hological descriptions with sketches of some invertebrate, plant and vertebrate
(molar tooth) fossil-genera are also given here.
Lastl_y I ~xpress ~y t~anks to my wife Chhanda, my colleagues, friends and students who
alwa~s msplfed me 111 _wnting this work. T am also thankful to all the predecessors in this field
whos~ books and published papers have be cited here and have also helped me in my classroom
teachmg.
_I would be an extremely happy man in case my beloved students would be benefited after
~omg th~ough the book and would be able to upgrade their ideas. My work would be successful
· ou t th e su b.~ec t pa Iaeon to Jogy.
m case 1t can create greater interest among students ab
Amal Dasgupta
I st January, 2005
Department of Geology
Asutosh College, Kolkata
CONTENTS
INTRODUCTION
PART I : PRINCIPLES OF PALAEONTOLOGY 1-94
~apter 1 : A General Account of Fossils
1.1 Fossils-What are they?
1.2 Kind of Fossils
1.3 Conditions of Preservation
1.4 Modes of Preservation
1.5 Causes of Imperfection of Fossil Record
1.6 Taphonomic Alteration 9
1.7 Use of Fossil IO
Chapter 2 : Systematic Palaeontology 12-21
2.1 · Classification and Nomenclature 12
2.2 Phylogenetic Classification )2
2.3 Phenetic (Typomorphic) Classification 13
2.4 Taxonomic Categories 13
2.5 Concept of a Species 14
2.6 Naming a Species : Bionomial System of Nomenclature )4
2.7 Law of Priority-Homonyms and Synonyms 16
2.8 Type Specimens , 16
2.9 Classification of Org~nic "Kingdom 17
2.1 O Outline of Classification of Plant Kingdom 17
2.11 Systematic Position of Man in Animal Kingdom 20
\
Chapter 3 : Grade and Growth of Animals 22-29

3.1 Grade and Body Plan : Grouping of Animals into Phyla 22
3.2 Study of Ontogeny 25
3.3 Type of Growth of Organic Skeleton 25
3.4 Growth rate of animal 27
3.5 Isometric and Anisometric Growth 27
3.6 Causes of Anisometric Growth 29
Chapter 4 : Spatial Distribution of Organisms 30-37

4.1 Study of Biogeography 30
4.2 Causes of Climatic Variation 30
4.3 Geographic Distribution of Organisms-Dispersal 30 /
4.4 Some Related Terms on 'Dispersal' 31

4.5 Factors Controlling Dispersal 31
/
4.6 Barriers to Dispersal 32
(ix)
Palae(~eo)WP(t)-2
(x)
PALAEONTOLOGY
47
· ~p~s of Climates and Related Vegetations
32
4.8 B1ot1c Distribution
34
4.9 Latitudinal changes and Taxonomic Diversity 34
4.10 Climates of the Past 34
Chapter 5 : Stratigraphic Palaeontology 38-56
5.1 Stratigraphic Subdivisions and Units 38
5.2 Law of Faunal Succession :
The Basic Principle of Stratigraphic Palaeontology 39
5.3 Principle of Uniformitarianism and Fossil 40
5.4 Time Distribution of Organisms 40
5.5 Major Lines of Evolution of Plant and Animal Group 43
5.6 History of Life Through Ages 45
5.7 Use of Fossils in Stratigraphy 51
Chapter 6 : Palaeoecology 57-70
6.1 Study of Palaeoecology and its Limitations- 57
6.2 Procedure for Palaeoecologic Interpretation 57
6.3 Environments on the Present Earth Surface 59
6.4 Terminologies Related to Ecology 60
6.5 Factors Controlling the Occurrence and Abundance of an
Organism in its Environment 63
6.6 Methods of Ecologic Study and Interpretation 65
Organic Evolution 71-94
7. l Sign of Life 71
7 .2 Origin of Life and Its Diversity 71
7 .3 Idea of Organic Evolution 71
7.4 Basic Concept of Organic Evolution 72
7 .5 Theories of Organic Evolution 72
7 .6 Genes and Heredity-Mendelism 74
7.7 Sexual Dimorphism 76 .
7 .8 Evidences of Organic Evolution 76
7.9 Neo-Darwinism 79
7.10 Modern Views on Evolution 79
7 .11 Processes of Evolution 82
7 .12 Pattern of Evolution within a Fossil Lineage 87
7.13 Pattern of Evolution among the Lineages 87
7.14 Rate of Evolution 88
~5 Course of Evolution and related Phenomena 89
7.16 Periodicity of Evolution 91
. ~: INVERTEBRATE FOSSILS 95-245
~apter 8 : Porifera (Sponges) 97-100

97
8.1 Systematic Position
97
8.2 Morphology
. ·.-
CONTENTS (xi)
8.3 Classification JOO

8.4 Skeletons 100
8.5 Geological History 100
: Cnidaria 101-119
9. I General Characters IOI
9.2 Broad Subdivisions IOI
9.3 Morphological Features of Coral Exoskeleton 103
9.4 Classification 112
9 .5 Diagnostic Characters of the Four Orders I 12
9.6 Reproduction and Growth 113
9.7 Evolution I I3
9.8 Coral Reef - 115
9.9 Ecology 115
9.10 Geological History O' I 16
.H - 9.11 Stratigraphic Use of Corals 116
·, ~ _;:"J 9.12 Some Coral-Like Fossils of Uncertain Affinity 117
~ a ~ t e r c , : Brachiopoda 120-136
· 10.1
Introduction · 120
t\I
I 0.2
Structure and Composition of Shell . 120
10.3
External Morphologic Features 125
l 0.4
Internal Morphologic Features 129
10.5
Classification 131
l 0.6
Ecology 133
~ Stratigraphic Importance. 135
' / ~ Geological History 135
~apter 11 : Bryozoa 137-139
11.1 General Features 137
11.2 Morphology 137
11 .3 Ecology 137
11 .4 Geological History 139
Mollusca 140-147
12.1 Introduction 140
12.2 Subdivisions 140
12.3 Some Aspects of Growth Related to Shell Morphology and
Shell Shape 143
12.4 Origin and Phylogeny of Mollusca 144
12.5 Effect of Predation on Life-habit of Molluscs 147
Pelecypoda (Bivalvia) 148-163
13.1 Generat Features 148
13.2 External Features 148
\ '
13.3 Internal Features 151
(xii)
PALAEONTOLOGY
13 .4 External Surface Ornamentation
155
13.5 Dimensions and Orientation
157
13.6 Classification
158
13.7 Ecology
159
~· / ;l
te~ 13 ·8
Geological History and Stratigraphic Value 162
C
~te'(_Y : Gastropoda 164-174
14.1 General Features 164
14.2 Shell Forms and Composition 164
14.3 A. Major Morphologic Features Associated with
Shape of the Shell 164
B. Coiling Patterns and Associated Features 169
C. Features Associated with Spire 171
D. Features Associated with Body Whorl 171
E. Surface Ornamentations 172
14.4 Classifications 172
14.5 Ecology 172
14.6 Morphological Analysis of Gastropod Shell Forms 173
/ ~ _.14.7 Evolution ~nd Geological History 174
\?3pter AS): Cephalopoda 175-194
0 15 .1 General Features 175
15.2 Morphological Features of Cephalopod Shell 175
15.3 Morphology of Belemnoids 183
15.5 Function of Some Features Found_in Cephalopod Shell 186
\.)>.§'. Morphological Specialization in Response to Life Habit 189
~ Evolution of Ammonoidea 189
JK.8 Probable Causes of Extinction of Ammonoids 193
~ Geological History 194
-"'L_ 15.10 Stratigraphic Use 194
vpu:r ~ : Arthropoda 195-210
16.1 Introduction 195
16.2 Subdivisions 195
16.3 General Features of Arthropods 198
16.4 Trilobites-General Characters 198
. 16.4.1 Morphological Features of Trilobites 199
16.4.2 Classifications 205
16.4.3 Ecdysis and Ontogeny 207
16.4.4 Ecology 208
16.4.5 Trace Fossils Related to Trilobites 208
1~.6 Stratigraphic Uses
208
~.4.7 Ancestry and Geological History
210
CONTENTS (xiii)
Echinodermata 211-236
~17
17 .1 General Characters 2I I
17 .2 Classification 216
17 .3 Echinoids 216
17 .3.1 Morphology of Fossil Echinods 216
17 .3.2 Functional Morphology 226
~ . 3 Classification 228
17.3.4 Ecology and Mode of Life 229
~ . 5 Origin, Evolution and Geological History 233
17.3.6 Stratigraphic Importance 235
17 .4 Outline of Morphology of Crinoids 235
17 .5 Outline of Morphology of Blastoids 236
: Graptolites 237-240
18. l General Morphologic Features 237
18.2 Ecology 239
18.3 Biqlogical Affinity 239
~ Geological History and Evolution 239
. 19 : Record of Invertebrates Fossils from Phanerozoic
Rocks of India . 241-246
19.1 Occurrence of Invertebrate Fossils in India 241
19.2 Palaeozoic Fossils 241
19.3 Mesozoic' Fossils 241
19.4 Cenozoic Fossils 245
PART III : VERTEBRATE Fossn.s 247-386

Chapter 20 : Major Subdivisions of Vertebrates 249-255
20.1 Vertebrate Fossils 249
20.2 General Plan of Vertebrate Skeleton 249
20.3 Broad Subdivisions of Chordates 251
20.4 Some Diagnostic Characters of Major Groups of Vertebrates 253
Chapter 21 : Outline of Morphology of Some Skeletal Elements
of Vertebrates 256-283
21.1 Skull 256
21.2 Jaw Suspension 258
21.3 Diagnostic Features of Skulls of Tetrapods 259
21.4 Vertebrae and Vertebral Column 261
21.5 Some Characteristic Features of Vertebrae of
Different' Groups of Vertebrates 263
21.6 Vertebral Column and FisH~taii 267
-· 21.7
21.8
Ribs
Sternum
267
267
PALAEONTOLOGY
(xiv)
21.9 Bony Girdles and Limb Bones 269

21.10 Stages of Evolution of Limb Bones 271
271
21.11 Teeth
277
21.12 Accessory Skeletal Elements
Some Aspects of Evolution of Vertebrates 284-292
Chapter 22
' 284
22.1' Invertebrate to Vertebrates
286
22.2 Evolution of Jaws
286
22.3 From Water to Land
287
22.4 Rise of Reptiles
290
22.5 From Land to Air
Rise of Mammals 292
22.6
Geological History of Vertebrates 293-323
Chapter 23
23.1 Jawless Vertebrates 293
\.,
23.2 Advent of Fishes 295

23.3 Advan~ed Fishes 295
23.4 Amphibias-The First Terrestrial Vertebrates 300
23.5 Reptiles-The Rulers of Mesozoic 304
23.6 History of Birds 311
23.7 Mammals-Rulers of the Present 312
Chapter 24 . Dinosaurs 324-337
24.l Introduction 324
24.3 Ancestry 327
24.4 Some Important Sites of Dinosaur Fossils 328
24.5 General Characters, Lifeways and Adaptations 328
24.6 Geological History 333
24.7 Causes of Extinction 334
Chapter 25 .. Evolution of Horse and Elephant 338-349
25.1 Evolution of Horse 338
25.2 Evolution of Elephant 345
Chapter 26 . Primates and Ancestry of Man 350-375
26.J Insectivora-Primate Transition 350
26.2 Basic Lines of Adaptation for an Arboreal Life 350
26.3 Position of Man in the Order Primate 351
26.4 Kins of Man
353
26.5 Ancestry of Man
355
26.6 Stages of Evolution of Man
.
Chapter 27 • Record of Vertebrate Fossils of India
361
376-386
27.1 Major Occurrences
376
27.2 Importance of Vertebrate Fossils in Gondwana Statigraphy 376
CONTENTS (xv)
27 .3 Importance of Dinosaur Fossils of India 379

27.4 Siwalik Mammals 379
PART IV PLANT Fossn.s 387-430

,_ Chapter 28 Introduction to Palaeobotany 389-399
28. l Limitations in Using Plant Fossils 389
28.2 Outline of Morphology of Plants 391
Chapter 29 Major Subdivisions and Geological History of Plants 400-416
29. l Major Subdivisions of Plants 400
29.2 Algal Structures : Stromatolites 401
29.3 Psilopsids 40]
29.4 Lycopsids 403
29 .5 Sphenopsids 405
29 .6 Pteropsids 405
29.7 Geological History of Land 'Floras 412
,., Chapter 30 Record of Plant Fossils of India 417-430
30.1 Broad Subdivisions of Indian Floras 417
30.2 Precambrian Flora 417
30.3 Post-Cambrian-Pre-Gondwana Flora
(Cambrian to Carboniferous) 418
30.4 Gondwana Elora_(Premo-Carboniferous-Lower Cretaceous) 418
30.5 Post-Gondwana Flora (Upper Cretaceous-Cenozoic) 426
PARTV : MICROFOSSILS 431-478

Chapter 31 : Introduction to Micropalaeontology 433-478
31.1 Historical Development of Micropalaeontology 433
31.2 Microfossils : A General Account 434
31.3 Advantage of Study 434
31.4 Limitations of Study 435
31.6 Microscopic Animal Fossils 437
31.6.1 Foraminifera 437
3 J.6.2 Radiolaria 461
31 .6.3 Ostracods 462
31.6.4 Estheriid 465
31.7 Microscopic Parts of Animals-Scolecodonts 465
31.8 Fossils of Uncertain Affinity--Conodonts 465
31.9 Microscopic Plants 469
31.9.1 Dinoflagellates 469
31.9.2 Coccoliths (Calcareous nannoplanktons) 469
31.9.3 Acritarchs 470
(xvi) PALAEONTOLOGY
31.9.4 Diatoms 470

31.9.5 Chara 471
31.10 Microscopic Parts of Plants-Spore-Pollen 471
31.11 Application of Microfossils in Petroleum Exploration 477
PART VI STUDY OF Fossn.s 479-546
Chapter 32 : Practical Works on Fossils 481-546

I. Collection and Preparation of Fossils and Method of their Description
32.1 Collection of Fossils in the Field 481
32.2 Separation of Fossils from Rocks and their Cleaning 483
32.3 Description of Fossil Specimens 484
II. Description and Identification of Some Fossil Genera
32.4 Phylum : Protozoa , 489
32.5 Phylum : Cnidaria 495
32.6 Phylum : Arthropoda 501
32.7 Phylum : Brachiopoda 503
32.8 Phylum : Mollusca 509
32.9 Phylum : Echinodermata 527
32.10 Description of Plants Fossils 535
32.11 Molar Tooth of Some Mammals 543
BIBLIOGRAPHY (i)-(xi)
INDEX (i)-(xxxi)
TABLE CONTENTS
Table No. Contents Page No.

I. Important compositional materials of organic skeletons 5
2. A simplified scheme of the classification of animals 18
3. Climates and types of vegetation 33
4. Leaf morphology and climate 3,S
5. Sediments and climate 37
6. Geological time units and corresponding chronostratigraphic units 39
7. Geological time and major groups of organisms 52
8. Periods of mass extinction and the affected fauna 94
9. Distinction between TetracoraJs and Hexacorals 11 I
IO. Distinction between Brachiopod and Pelecypod shell 158
I I. Distinction between Gastropod and Cephalopod shell 185
12. Distinction between Nautiloid and Ammonoid shell 185
13. Distinction between regular and irre·g ular Echirtoids 225
14. Major skeletal elements of skull of different Vertebrates 281
15. Outline of phylogeny of Horse 343
16. Outline of phylogeny of Elephant 349
17A. Sites of Australopithecine and Early Hominid fossils 365
17B. Succession and important fossils of Olduvai Gorge 367
18. Sites of fossils of Homo sapiens 373
19. List of some Gondwana Vertebrate fossils of India 377
20. List of some Post-Gondwana-Pre-Siwalik Vertebrate fossils of India 380
21. List of some Siwalik Mammals of India 383
22. Distribution of Lower Gondwana flora in India 421
23. Distribution of some Indian Plant fossils in Gondwanaland 423
24. Distribution of Upper Gondwana flora 425
25. A comparison between Lower and Upper Gondwana flora 426
26. Neogene flora of India 429
27. Foraminiferal assemblage in successive depth at Latitudes 455
28. Zonation of Marine Tertiary Rocks of India 460
29. Distinction between Conodonts and Scolecodonts 469
30. A Comparision between Spore and Pollen 476
Palae(Geo)WP(t)-3
(xvii)
r -~ _ . ---- -
FIGURE CONTENTS
Page No.
r;,· N Contents .
rigure o. . . mmon sediments (Chemical
1.1 pH - eH Boundaries controlhng formaft1onKof co bein & Garret, 1952) 4
/Biochemical) with fossil-potential (a ter rum 8
1.2 Various modes of fossil preservation. d . and after fossilization 8
1.3 Different types of taphonomic alteratmn ur~~g Pl ,, 24
3.1 Divisions of animals on their "Grade" and Body an
24
3.2 Two types of growth in animals
28
3.3 Change of growth rate . 28
3.4 Isometric and Anisometric growth of animal
42
5.1 Different types of stratigraphic units f I t
A broad phylogenetic tree showing evolution o.f diff~rent group: o p an
42
5.2
Broad phylogenetic relationship among the maJor ~mmal group 44
5.3
48
5.4 Some elements of Ediacara faun_a
Different kinds of biostratigraphic zones 48
5.5
6.1 Major types of marine environment and related marine habitats
represented by a hypsographic curve 58
6.2 Food pyramid and food chain 66
6.3 Nature of spatial distribution of a species-population 66
7.1 Pattern of evolution within different lineages 84
7.2 Pattern of evolution among different groups 86
8.1 Morphological features of Porifera (sponge) 98
8.2 Different types of spicules 99
9.1 Wall structures and inner features of Anthozoan po~yp (coral) 102
9.2 Longitudinal section of three major groups of Cnidarian polyp 104
9.3 Basic morphologic features of an ideal simple coral skeleton 104
9.4 Shape of Corallites 106
9.5 Different types of colonial coral 106
9.6 Morphology of septum
108
9.7 Axial structure, dissepiments and tabulae
110
9.8 Types of asexual reproduction of coral
110
9.9 Three successive stages of growth of coral reef
114
9.10 Geological range of major groups Cnidaria ~d related forms
9.11 114
Some coral-like fossils of uncertain affinity '
10.1 118
Attachment and internal organization of a typical articulate Brachiopod
10.2 Inarticulate Brachiopods . 121
10.3 Shell microstructure of Brachiopod 121
J0.4 122
External morphology of Brachiopod shell
10.5 123
External morphology of Brachiopod shell
124
(xviii)
C& k f
FIGURE CONTENTS (xix)
10.6 Internal morphology of Brachiopod shell 128

10.7 Muscles and their function in Brachiopod shell 128
10.8 Life habits of some Brachiopods 132
10.9 Geological range and relative abundance of ·some major groups of
Brachiopods 134
11.1 Morphology of Bryozoans )38
12.1 Basic structural plans of different living Mollusc classes with reference to
Hypothetical archimollusc 141
12.2 Growth and related shell morphology of Mollusca 142
12.3 Geological range of major groups (orders) of Pelecypod, Gastropod and
Cephalopod 146
13.1 Basic morphological features and orientation of Bivalvia shell 149
13.2 General morphological features of Pelecypod (Bivalvia) shell 150
13.3 M~rphological features of Pelecypod (Bivalvia) shell 152
13.4 Types of dentition in Pelecypod shells 154
13.5 Muscle -scar and mechanism of opening and closing the valves by
action of adductor muscle and Ligament shown in cross sections 156
13.6 Symmetry, orientation and dimensions of Brachiopod a~d Pelecypod shell 156
13.7 Life habits of some marine Pelecypods 160
14.1 General features of Gastropods 165
14.2 Morphology of Gastropod shells 166
14.3 Morphology of Gastropod shells 168
14.4 Morphological analysis of shell-forms (after Linsley, 1977) 170
15.l Chief viseral organs and .wall _microstructure of an ideal living
Cephalopod (Nautilus) 176
15.2 Morphology of Cephalopod shell 177
15.3 Morphology of Cepha]opod shell 178
15.4 Morphology of Cephalopod shell 182
15.5 Morphology of Belemnoid shell 184
15.6 Evolutionary pattern of Ammonoid in Mesozoic
(after Moore in Treatise, Part-L) 188
15.7 Evolution of Ammonoids 190
16.1 Some representatives of Arthropods 196
J 16.2 Skeletal structures and appendages 197
16.3 Basic morphology of Trilobites-dorsal views 197
16.4 Morphology of Cephalon 200
16.5 Morphology of Glabella,and Eye 202
16.6 Morphology of thorax 204
16.7 Morphology of Pygidium (dorsal view) 206
16.8 Ontogenic stages of Trilobites 206
16.9 Geological range of Trilobites (orders) 209
17.1 Some members of living Echinoderms 212
17.2 Broad similarity in internal morphology of different groups of living
Echinoderms (Cross section passing through mouth and anus) 213
(xx)
PALAEONTOLOGy
17.3 Morphological features of regular and irr,egular Echinoid test 214
17.4 Morphological features of Echinoid test 215
17.5 Morphological features on Echinoid test 218
17.6 Ornamental features on Echinoid test 220
17.7 Some special morphologic features of Some Echinoid tests 224
17.8 Water vascular system and associate features 227
17.9 Different modes of life of Echinoids 230
17.10 Stratigraphic range and relative abundance of the common groups of
Echinoids 232
17.11 Structures of Crinoids 234
17.12 Morphology of Blastoids 234
18.1 Morphological features of Graptolites 238
20.1 Major skeletal elements of aquatic and terrestrial vertebrates 250
21.l Three stages of development of Chondrocranium 257
21.2 Different types of jaw suspenion 257
21.3 Major skeletal elements of skull of Vertebrates 260
21.4 Morphology of Vertebra 262
21.S Pattern of vertebrae of Tetrapods 264
21.6 Sternums and pectoral girdles of Tetrapods 265
21.7 Bones of forelimb of Vertebrates 266
21.8 Pelvic girdles and hind limbs of Vertebrates 268
21.9 Caudal fins of fish and foot postures of Mammals 270
21.10 . Evolution of limb bones from fish-fin 272
21.11 Morphology of teeth of Vertebrates ·274
21.12 Morphology of cheek tooth 275
21.13 Different types of dermal scales in fish 276
21.14 Dermal skeletons of some Vertebrates 278
22.1 Diagramatic representation of origin of Chordates based on idea of
berrill ( 1955) 285
22.2 Evolution of jaw form branchial arch in Placoderm fish 285
22.3 Change of skull pattern and evolution of major groups of Vertebrates 288
22.4 Transverse section of an amniote egg of Reptile 288
23.1 Extinct and living jawless Vertebrates 294
23.2 Representatives of major groups of Fishes 294
23.3 Evolution of different fish-groups 296
23.4 Extinct and surviving Amphibias 301
23.5 Some extinct Reptiles and their basic skull-structure 302
23.6 Divergence of Reptiles in Mesozoic 310
23.7 Divergence of Eutherian Mammals in Cenozoic 310
23.8 Pattern of molar tooth of Early Mammals 314
23.9 Some ancestral groups within Mammals 316
23.10 Diagramatic sketch showing phylogenetic relations, geological ranges
and relative abundance of major groups of Mammals 318
(xxi)
FIGURE CONTENTS
23. 11 A simple phylogenetic tree of Vertebrates showing evolution of different

surviving groups 321
23 .12 Geological range and relative abundance of major groups of Vertebrates 322
24.1 Structure of pelvis of Dinosaurs 325
24.2 Different groups of Dinosaurs 325
24.3 Some earlier forms of Dinosaurs and their probable ancestor 326
24.4 Different types of crestal structures of head in Dinosaurs 330
24.5 Phylogeny of Dinosaurs 330
25.1 Adaptive modifications of structure of horse's foot 339
25.2 Change of some skeletal features in the evolution of horse from
Hyracotherium to Equus 339
25.3 Evolution of different groups of Elephants 344
26.1 Family members of primates showing variation of skull and jaw 352
26.2 Structural changes from Ape to Man 354
26.3 Change of skull pattern in Hominid evolution showing probable relation
among the groups 357
26.4 Human fossils and cultural evolution 358
26.5 Stages of evolution of Man 360
26.6 An outline of phylogeny of Man 368
28.1 General morphology of plants 390
28.2 Internal morphology of stem 393
28.3 Morphology of leaf 394
28.4 Types of venation in leaf 396
28.5 Types of leaf and phyllotaxy 397
28.6 Flower, fruit and seed 398
29.1 Stromatolites 402
29.2 Some Fossil-plants and their living counter-parts 404
29.3 Some Fossil-plants and their living counter-parts 406
29.4 Living Conifer and Ginkgo 408
31.1 Foraminifera and its life cycle (Dimorphisms) 438
31.2 Micro-structures of test-wall 440
31.3 Mode of addition of chambers wall 440
31.4 Type of test and coiling 441
31.5 Type of test (Multilocular) and coiling 442
31.6 Ontogenic development of chambers 442
31.7 Shape of tests 446
31.8 Shape of chambers 446
31.9 Aperture of Foraminifera 448
31. 10 Ornamental features and septal filaments of Foraminifera 450
31.11 Pillars in thin section (Vertical sectional views)
450
31.12 Change in proportion of recent planktonic, calcareous benthic and
arenaceous benth~c Foraminifera with increasing water depths from
shore across continental shelf down to continental slope
455
1
(xxii) PALAEONTOLocy
31.13 Radioloria 462

': '
31.14 Ostracods, Estherids and Scolecodonts 464
31.15 Conodonts 466
31.16 Some Plant Microfossils (Chara, Diatoms, Dinoflagellate and Coccolith) 468
31.17 Life cycle of Plant and formation of spores and pollen 472
31.18 Morphology of spores and pollen 474
32.1 Field equipments related to fossil collection 482
32.2 Nature of symmetry within different groups of Animals 485
32.3 Different types of shapes found within animal skeleton 486
32.4 Larger Foraminifera 491
32.5 Larger Foraminifera 492
32.6 Smaller Foraminifera 494
32.7 Tabulate Corals 497
32.8 Rugose (Tetracorals) Corals 498
32.9 Hexacorals 499
32.10 Trilobites 500
32.11 Brachiopoda 502
32.15 Pelecypoda (Bivalvia)
510
32.16 Pelecypoda
32.17
512
Pelecypoda
32.18 Gastropoda
514
32.19 Gastropoda 516
32.20 Gastropoda 518
32.21 Cephalopoda 520
32.24 Echinoidea 526
32.27 Crinoidea and Blastoidea 533
32.28 Plant Fossils 534
32.32 Molar Tooth of Mammals 540
32.33 Molar Tooth of Mammals 544
545
INTRODUCTION
Geology is the science of the earth. More precisely it deals with the different aspects of the
past earth. Thus geologists have to study the origin and mode of development of the earth, its
internal structure and composition, the different dynamic processes now operating on and below
the earth's surface, organic remains within the rocks of the earth and so on. All these studies lead
us to a precise knowledge about the physical, chemical and biological history of the past earth
since its formation and to reconstruct this history is the ultimate aim of a geologist. Study of all
these aspects of the primitive earth is however based on the two principal materials : rocks and
fossils. The study of different aspects of rock leads us to interpretation regarding history of physical
and chemical development and fossils are carrying informations about developmental history of
the past organisms of the earth.
Fossils thus constitute an important branch of geology called 'Palaeontology' (paleo : past;
ontology : life history), literally meaning lie history of the past organisms of the earth. Fossils
can be studied from different angles and accordingly, palaeontology has several subdivisions viz.:
lnverlebrate palaeontology (dealing with invertebrate fossils), Verlebrate palaeontology (study
of vertebrate fossils), Palaeobotany (study of plant fossils), Micropalaeontology (study of
microscopic fossils) etc. Again study of plant or animal fossils can be done along different lines
such as : study of morphology and classification (Systematic palaeontology), habits and habitats
of fossils organisms (Palaeoecology), distribution of fossil organisms in time and space
(Stratigraphic palaeontology) and also the evolutionary history of organisms.
Study of fossils involves several steps : (i) collection of fossils in different field sites
(ii) morphologic study leading to identification and recognition of systematic position of the
different fossils (iii) study of ecology from their modes of fossilization, morphologic features
. and from the associated fossils and rocks (iv) study of stratigraphic distribution of the fossils and
finding out the ages of fossils and associated rocks.
Study of a large number of fossils of different plants/animals from a wide area may lead us to
build a regional history of the past organisms. Correlation of numerous such data obtained from
different parts of the world will help to reconstruct the organic history of the past earth as a whole.
The fossil record is only a small sample of past life. Palaeontologists may give several
interpretations based on this sample which are probably correct but never certain. Most .o f these
interpretations are based on 'principle of actualism' which itself has its own limitations. Thus
any study of fossils or use of palaentological data must be based on clear understanding of strength
and weakness of the record. Interpretations from a fossil record in many cases, become limited
for several reasons such as : incompleteness of the record; complexity in its preservational history,
presence of such fossils which have no living counterparts to compare and so on. Yet, palaeontology
has lot of importance in the geological science. William smith discovered about 150 years ago
that fossil sequence may indicate relative age of rocks. Comparison of fossils with their living
counterparts may lead to interpretation of past climate, envin;>nment and even to reconstruction
of past geographic elements of the earth. Moreover, the understanding of modem biological system
remains incomplete until its historical development is known from the study of fossils.
(x.xii1)
j
PART - I
PRINCIPLES OF PALAEONTOLOGY
Chapter 1
A GENERAL ACCOUNT OF FOSSILS
1.1 FOSSILS-WHAT ARE THEY?

The word fossil in latin means 'anything dug up' or extracted from the earth. The terms
Fossilia nativa and Fossilia petrificata were used to describe minerals and organic remain s
respectively. A precise definition of fossil may be as follows : Fossil is an actual remain or
indirect evidence of a prehistoric life preserved within a rock under natural conditions. Therefore,
before considering an object as a fossil one should be careful. Firstly, the object in concern
must have some relation to an organism. This relation may be direct or indirect. No structure
produced by an inorganic process should be called a fossil. Secondly, the related organism must
be prehistoric in nature. By the term prehistoric we mean anything prior to the beginning of
history of modern man who appeared at about 40,000 years ago. So anything related to modern
man should not be treated as a fossil. Thus remains of the Indus civilization or the Nile
civilization are not fossils; rather they are ancient historical remains or archaeological remains;
but the remains of Neanderthals and the stone tools used by them are fossils as these are
prehistoric remains. Thirdly, the fossil must be preserved by natural processes . An organic
material preserved artificially is not a fossil. Thus 'mummy' of Egypt is by no means a foss il.
As organisms are mostly inhabiting on the earth surface, their fossils are mostly preserved within
sedimentary rocks forming on the earth surface. Volcanic rocks, also forming on the earth surface
may sometimes contain well-preserved fossils.
1.2 KIND OF FOSSILS

According to type of organisms fossils may be called plant fossils or animal fossils. Both
of them may be represented by complete organisms or fragmented parts of them. In another
way fossils may be grouped considering their size viz., microfossils and megafossils. Larger
animals and plants found in unbroken or broken conditions constitute mega or macrofossils.
Microfossils are those which may or may not be visible in naked eye but the microscopic
observation is a must for their proper recognition. They may represent very small-sized animals
or plants or microscopic parts of larger animals or plants. Foraminifera and radiolaria are
protozoan animals often preserved as microfossils. Ostracodes and estherians are arthropod-
microfossils. Diatom is a microscopic algae. Spores and pollen, microscopic reproductive units
of plants are also common plant-microfossils. On the other hand, echinoid spines, plates and
some microelements of unknown affinity like co11odo11ts are also treated as microfossils. Study
of microfossils has gained much importance at present as study of microfossils has several
advantages over megafossils. A small rock sample may yield numerous microfossils so that
they can be investigated as a population. They also show a minimum degree of deformation
and morphologi~ changes at the time of fossilization for their small size. The branch of
palaeontology which deals with various aspects of microfossils is called Micropalaeontology.
Paly11ology is a branch of micropalaeontology dealing with fossil spores and pollen only.
2 PALAEONTOLOGY
1.3 CONDITIONS OF PRESERVATION

All the organisms once existing on the earth surface cannot be fossilized after their death .
Preservation of an organism as a fossil within a rock is merely a matter of chance. ~ore
precisely, it can be said that a series of favourable conditions are necessary for transforming a
dead organism to a well preserved fossil. These favo.urable conditions are as follows :
1.3.1 Presence of hard skeletal matters · . .

The most favourable condition for an organism for becoming fossil after its d.ea~h ts the
presence of some sort of hard materials in the form of exoskeleton or endoskeleton m its body.
The soft organic tissues are usually found decomposed and destroyed after the death o~ the
organism by the activity of bacteria in the presence of oxygen in atmosphere and are unlikely
to be preserved. Hard tissues or skeletal matters .may escape such destructive processes for a
considerable time and this may favour its · fossilization provided subsequently other conditions
be fulfilled. For this reason, soft bodied animals like most of the protozoans, worms and annelids
are rarely preserved as fossils and so also the soft parts of other organisms are hardly found as
fossils.
1.3.2 Detachment from atmosphere

Oxygen in atmosphere i,s one of the strong decomposing agent. Thus after the death of an
organism, the dead body needs to be detached from atmospheric contact by some natural
processes as early as possible. Otherwise it will be easily decomposed by bacteria in presence
of oxygen and hence lost. Soft parts will go first foJlowed by hard· parts of the body. A temporary
covering on this dead body by sand, silt (say in desert) or water in river, lake and sea may be
helpful to protect the body from its immediate destruction·. From this viewpoint, marine and
other aquatic animals have already some plus points over terrestrial animals, as the former after
their death usually settle to the bottom of sea, river or lake bed and are unlikely to come in
contact directly with open atmosphere. Thus they have always a greater chance of fossilization
compared to terrestrial organisms which usually after their death remain in open atmosphere
and are likely to be destroyed before preservation.
\
1.3.3 Sedimentation and permanent covering

The. chance of preservation. . of a dead organism under a temporary cover m ay b e increase· d
considerably only when 1t will be covered permanently by some material th d.
· ·. · · h. · . . s, e se 1ments,
prec1p1tating wit in nver, lake and sea. As this rate of sedimentation 1· s f· · .
. . , ar more quicker m
sea, marine orgamsms face the most suitable condition to be fossil. d · h" .
. . · ize wit m sediment and
fossils of such animals are much greater in number than terrestrial . d h · .
. H h , . . , an ot er aquatic group of
organisms. owever, t e rate of sed1mentat10n and size of sed· t t· .
· p· · . · unen o ten result m selective
preservation. me grained sediments like clay silt chemic·lll . · · ·
. · ' ' ' Y piecipitated limestone act as
exce 11 ent med1ums. Again, slow rate of sedimentation is b tt i . . .
condition leading to one time rapid sedimentation m·1 e. er or prellservat1on. Catastrophic
. d d . . .
loca 11ze area pro ucing a highly fossiliferous rock sot ·
'y c,mse exce ent preservations . t .
. . d . ·· a a
, 11e1I mes ca 11 e cocqm a. 11
1.3.4 Chemical e11viro11ment
Preservation often becomes selective depend·ing th f
.. . . on e nature o chemic·1t e · f
the depos1t1onal basin. In an acidic environment ( H < S) . , . ' nv1ronment o
. P 7 · most of the calcareous skeletons
., - ------ --- - -
A GENERAL ACCOUNT OF FOSSILS
of organisms are lost by dissolution. Similar is the fate of silicious skeletons in an alkaline
condition where pH > 7.8. This is the reason for general absence of fossil organisms with
silicious skeletons within limestone and absence of calcareous fossils within silicious sediments.
The types of chemical/biochemical environments controlling types of chemically precipitated
sediments and their organic content is shown in fig. 1-1.
In general, within the photic zone of sea, the excess of photosynthetic plants compared to
animals keeps CO 2 content of the water at lower level where pH value ranges between
7.8-8.3. Such an alkaline environment can supply more CaC01 than its power of dissolution.
But at depth below the photic zone, general absence of photosynthetic plants and presence of
other organisms performing respiration, cause increase of CO 2 content in sea water that lowers
its pH value making the environment more addic. Such a condition can exist even at shallow
depth of the· sea in the higher latitude. The decrease of sea water temperature with increase of
its depth or increase of latitudinal value also causes increase of its rate of solubility of CaC0 3 .
Here the water has more power of CaC01 dissolution than its supply. The level of sea water at
which the rate of CaCn'.\ dissolution becomes almost equals to its supply is called Calcium
Carbonate Compe11satio11 Depth (CCCD). CCCD level is quite variable at different seas
depending upon their geographic locations. In the Pacific Ocean this depth normally lies between
4000m.-5000m. at lower latitude but lies only at 400m.-500m. depth in higher latitudes. It
is quite obvious that organisms with calcareous skeletons are unable to thrive here and most of
the dead skeletons made up of CaC0 3 get dissolved under such environment. On the other hand
non-calcareous organisms are likely to be preserved here. For this reason, in a rock association
of calcareous and silicious fossils is an uncommon phenomenon.
1.3.S Effect of diage11etic processes

Several diagenetic processes may operate within a newly deposited sediment to convert it
into a suitable hard rock. Leaching by acidic ground-water may remove calcareous skeletons,
completely or partially. Occasionally, some inorganic minerals in solution within ground-water
may replace the organic skeletal materials partially or completely making the skeleton harder
causing its excellent preservation. Silica in solution frequently replaces CaC01 of calcareous
skeleton. Recrystallization often results hardening of organic shell helping its preservation but
excessive recrystallization may destroy most of the organic structures.
1.3.6 Less deformation and metamorphism of rocks

Excess deformation and/or metamorphism arising from orogenic movement, igneous
intrusious and other natural causes may destroy an otherwise well-preserved fossil within
sediment partially or completely. Himalayan Phanerozoic rocks are in general fossiliferous b t
there are areas of excessive deformation where fossil are scanty or almost absent. Re k' u
I . f f ·1·f d' I wor mg
or recyc mg o. a oss1 1 erous se 1ment may a so result in reduction or destruction of alre· d
preserved fossils. a Y
1.4 MODES OF PRESERVATION (Fig. 1-2)

Modes of preservation of fossils may be of three types fossils of , f · . . .
parts and indirect fossils. ' '· so t parts, fossils ot hard
~
Acidic alkaline
p'= 7 p" = 7.8

~I UC::::: ::.C::::: I f I f 1
11G
C
.N
l 8
.I!
zI ~i· · I I >~, -« I I ~~ 1
00
j I
M
0
II
1,
~
~
tr]
a
FIG. 1-1 : pH - eH BOUNDARIES CONTROLLING FORMATION OF COMMON SEDIMENTS <'::
""'1
(CHEMICAL/BIOCHEMICAL) WITH FOSSIL-POTENTIAL (AFTER KRUMBEIN & a
t""'
GARREL, 1952) a
C"}
~
. -· ,: . r·
.,. • • 'fl
A GENERAL ACCOUNT OF FOSSILS 5

t.4.1 Soft parts as fossils
In rare occasions soft parts of organisms or a soft bodied animal may be preserved in nature
under a very special condition. The entombment of insects in fo ssil resins or ambers (found
within Silurian rocks of China) or frozen mammoth and Rhinoceros within Pleistocene Siberian
ice are some examples. This is a very rare type of preservation which may be possible under
some conditions which prevent action of bacteria and scavengers (such as sudden entrapment
within resin or ice). Outer woolly skin, soft muscular inner parts, blood or even the last food
taken by the animals are found intact and undecomposed within the foss ils of Siberian ice that
has acted as a natural refrigerator.
1.4.2 Hard part fossils

The most of the common type of fossils found in nature are the exoskeletons or endoskeletons
of animals or hard tissues of plants. They may be divided into two groups viz., unaltered fo ss ils
and altered fossils.
(i) Unaltered hard parts

In this case, the original structure and/or composition of the hard parts remain unchanged
even after their fossilization. Many animals have hard parts composed· of inorganic matters
and accordingly, the hard-part fossils may be calcitic or aragonitic (most common within
invertebrate exoskeletons such as corals, brachiopods, molluscs and echinoderms),
phosphatic (vertebrate bones) and silicious (such as skeletons of radiolarias, sponges and
also some planktic algae, like diatoms). Organic hard parts are composed of chitin (found
in many arthropods), keratin, cellulose, bark etc. The last three are generally found within
a plant body. The common type of fossil hard parts and their relative abundance are shown
in Table-I.
TABLE-1
Important compositional materials of organic skeletons
~aa r t Carbonate Phosphate Silica Chitin Cellulose
X X
Algae
X
Higher Plant
X X
Protozoa
X X
Porifera
Cnidaria X
X X
Bryozoa
Brachiopoda X X
Mollusca X
Annelida X X
Arthropoda X X X
Echinoderma X
Chordata X . ~
PALAEONTOLOGY
6
(ii) Altered hard parts .

In most of the cases however fossilization processes cause partial or complete change
of the composition a~d structu~ of hard parts thus producing altered fossils .. In a disti/lized
fossil, some portion of original skeletal matters may be lost through leach1~g. or solution
by chemically active fluids making the fossil porous and lighter than th~ _on~mal one. In
other cases, when an original skeletal body itself is porous, the foss1hzat10n .leads to
infilling of those pores by mineral matters making the fossil skeleton more massive, hard
and heavy. These are called permilieralized fossils. More common is the process where
materials of original organic skeleton are removed with simultaneous addition of new
mineral matters. Sometimes, this replacement takes place slowly volume to volume or
even molecule to molecule. When such a process operates, in most of the cases, the
original organic structure of the skeleton may remain intact with only change of
composition (petrified fossil). This type of replacement fossil commonly found within
plants where organic matters of plant tissues are replaced by some inorganic minerals
through solution without destroying the tissue structure of plant. Silica is a common
replacing mineral and silicious petrified stem-fossils are very common in nature. Other
common minerals causing petrification are hematite, pyrite, calcite etc.
Carbonization is also a process commonly found in plants somewhat analooous to
distillation: where all _other organic matters within a plant body are removed ex;ept the
carbon. Th~s remov~l 1s due to incre~sed temperature and pressure together with activity
of anaerobic bacteria after deep bunal of plant-materials. For the reason of enrichment
of carbon most of the plant fossils are looked black. Formation of coal from plants
however represents an extreme stage of carbonization process when the or anic struct
of plants are completely lost. g ures
In some cases, it is found that a mineral of the original skeleton be

at de th "th · f may come unstable
p w1 mcrease _o pressure-temperature when it recrystallize t ~
mineral. These are recrystallized fossils. Aragonite is most) recs o ~rm a more ~table
the latter is a more stable form of the mineral co d fy ~stalhzed to calcite as
.. mpose o calcium carbonat F h'
reason,
I. aragomt1c
. skeletons are very rare with1'n & ·1, Id
,ossa s o er than· Cen . e. or F" is
t
ca cite grams of a skeleton may be recrystallized to coars . ozo1c age. me
not cause any compositional change and are oft II . d . e calc1.te. Such processes do
recryt~llization under metamorphic condition ~; ca. e ,socli.e,mcal alteratio11. Excess
orgamc structures. Y cause partial or total destruction of
1.4.3 Indirect fossil

Sometimes what we get as a fossil .
but may give some indirect evidenc~s o7~~ :~!. represent the actual organism or its skeleton
leaves a~e very often found as imprints i.e .. the iste~ce. Th~se are called i11direct fossils. Plant
soft sediment surface. These ire called impri,11:~~~;J:.af is lost leaving its impression on the
A mould fossil may be formed wh en an o .. · '
totally by solution or leaching leav·1ng rga.mc body (skeleton) within sed·an·, ent . d
d" h' · a void of 't h 1s remove
. f mg t is vmd be fine enough • det··a1·1 s of ext
surroun I s s ape at its place It' th
1 • ' • e se d"I ment
Th 1s orms an external moald. If th . . erna features of the ·h II .
e organic shell encloses a h II s e may be preserved.
O ow space as for in;;;t!lnr;, th;,
space between the pair of valves of pelcypods, brachiopods or the space within univalved
gastropods and cephalopods, it may be filled up by fine sediments or minerals. Such infillings
the voids within skeletons exhibiting internal morphological features, are often called i11temal
moulds. In general, infillings of moulds are called cast-fossils.
More important are those indirect fossils which not only give evidence of existence of some
organisms but also indicate their some sort of life activities. These are called trace fossils or
ichnofossils or lebe11spurrens. Trace fossils are of various types. Some of them represent some
sort of movement or locomotion of animals on the upper surface of a fine grained sediment-
bed. These include, tracks of insects, trails of worms, foot pri11ts of higher vertebrates
(dinosaurs) etc. Other common trace fossils are burrows of worms or arthropods, bori11gs,
tunnels, or shafts (made by some invertebrates), resting impressio11s, feeding traces, breeding
materials (eggs), coprolites (excretory materials of higher vertebrates such as dinosaurs)
gastroliths, faecal pallets (excreta of invertebrates) etc. Trace fossils are usually classified on
the behavioural pattern of the organism concern, such as : repiclmia (crawling impression),
cubichnia (resting impression), pascich11ia (surface feeder trace), fodi11iclznia (feeding trace),
domiclmia (dwelling structures) and fugiclmia (escape structure) etc. Trace fossils at present
are considered as powerful tools for understanding past sedimentary environments and behaviour
(ecology) of the animals which lived in them. One reason which makes these fossils more useful
in ecologic study is that such fossils are always preserved in their original sites (in situ fossils) .
Some common types of trace fossil are shown in fig . 1-2.
Study of modes of preservation of fossils is getting a considerable importance at present,
especiaJJy for understanding the nature of depositional environment and diagenetic processes
as they affect in various ways on the nature of preservation of fossils. For example, in an acidic
condition silica has a lower solubility than calcium carbonate and in such a condition calcium
carbonate of a calcareous skeleton is likely to go in solution and replaced by silica. In a reducing
environment formation of pyrite is very common which can replace the skeleta1 matters of many
invertebrate fossils like ammonities and brachiopods.
1.5 CAUSES OF IMPERFECTION OF FOSSIL RECORD

Fossil record may becomes imperfect or incomplete owing to the following reasons.
(a) Some animals may be without any hard part and consequently their fossi1s are rare or
absent. Many ancient protozoan animals, worms, annelids failed to leave the traces of
their body as fossils for want of hard skeletons.
(b) Many organisms especially terrestrial animals and plants are decomposed and destroyed
after death for want of suitable covering.
(c) Fossils even after preservation may be lost by various diagenetic processes.
(d) Even afte~ settling to bottom of the sea the dead animals/plants may be eaten up by
various deposit-feeders and scavengers living on the sea-floor.
(e) Fossils may b~ destroyed, fragmented and lost during post-dcpositi~nal deformation and/
or metamorphism.
Palac(Gco)WP-2
PALAEONTOLOG Y
C)
l,
b
b
(b) DISTILUZED FOSSIL
b
C PERMINERALIZED FOSSIL
Cast {External Cast (Internal)
(c) MOULD FOSSIL Showing Inner (organic shell materials completely
I
showing external replaced by mineral matter)
featw'cS Features
d
d d
HELMINTHOIDS SKOLITHOS
CRUZIANA C
(trailing impression (pennanent dwellings
(aawling impression (tunnel of worms\ ofwonn) of worms/insects)
of trilobites) (d) TRACE FOSSil..
FIG. 1 - 2 : VARIOUS MODES OF FOSSIL PRESERVATION
L UNALTERED FOSSil.. b. ALTERED FOSSil..S c. INDIRECT FOSSil..S d. TRACE FOSSILS
+U ~!i
*~
ylindri"cal
0
Spherical
c:~::>
_ , . Lenticular
after before
stem
flattened after
~~ N~tt
Shape change after fossilization : fossiJization
Dcfonned and loss of symmetry Articulated Disarticulated valves
....... .
after fossilization bivalved shell after fossilization
,...... (a)
.. ~--
.~,1
before fossilization
Unusortcd shells of different siu

deposited at place of death · (b)
• ea •-:r,p •
-~·-
~
~
;.r. 4a1::••
.,.
Sorted shells after transportation
-
FIG. 1 - ~ DIFFERENT TYPES OF TAPHONOMIC ALTERATION DURING AND
AFTER FOSSILIZATION
(t) There may be loss of record of life m a sedimentary sequence showing one/more
stratigraphic breaks.
(g) Fossils may occur in such places beyond the reach of man or remain below the surface
unexposed.
1.6 TAPHONOMJC~ALTERATION (Fig. 1-3)

Individual organism or an assemblage of organisms have to undergo a series of natural
processes in the interval between their death and fossilization causing several types complications
and distortion within a fossil or in an assemblage of fossils. Several informations as regards
the morphology, ecology, environment, derived from study of fossils may be incomplete,
distorted or even erroneous due to such processes unless palaeontologists, take necessary
precautions. Thus a careful study of post-mortem history of a fossil or a fossil-assemblage is a
must. This study is called taphonomy, which mainly includes study of two groups of processes.
Study of the processes operating after death and before final burial is called biostratinomy and
the study of other processes operating after burial are called diagenetic study. The chief pre-
deposition process is transportation, causing physical, chemical and biological damages to an
organic body/skeleton. This may cause mechanical wear, partial loss through solution. It is quite
obvious that a fossil/assemblage of fossils preserved in situ may escape the hazards of
transportation. Further post-depositional and pre-burial damages may be caused by epifaunal
attachment, attack of scavengers or boring by borer animals (often called bioturbation). After
the final burial within sediment, the fossil may be further altered and damaged by several
diagenetic processes, such as compaction, compression, solution and leaching, replacement of
original skeletal matters by foreign materials, recrystallization etc. Palaeontologists have to detect
all these taphonomic alterations and overprints in a fossil or within an assemblage of fossils
before reaching to a correct interpretation as regards morphology, environment, palaeoecology,
and evolution related to the fossil or fossil assemblage. Study of taphonomy may be done from
two view points.
1.6.1 Taphonomic changes found within a fossil (Fig. l-3a)

Skeleton of a dead organism during its transportation may lose some materials through
mechanical erosion or action of chemical solution. Long transportation may result in loss of
such structures like external spines/tubercles, marginal flange which may cause an overal I
change of external shape and ornamentation of the skeleton or even fragmentation of the
skeleton. More distortions appear after deposition of the skeleton on the basin floor (sea floor) .
Before final burial the skeletal mass may be attacked and even eaten up by sediment eaters or
scavenging animals; may be used by epifaunal animals as their host. Burrowing or boring
animals may also cause some serious damage of the shell. After deep burial the skeleton has
to bear enormous pressure of the overlying sediment, and this may cause further distortion of
its overall shape and symmetry. For example a spherical/subspherical body under such condition
tends to become lenticular by the effect of vertical pressure. Unequal lateral pressure may cause
change of its original symmetry resulting in a deformed fossil. Excessive deformation leads to
frag~entati~n of the skeleton. Original skeletal matters may be lost or replaced through solution
leading to distillation, petrifaction and permineralization. There may be loss of internal organic
structure by recrystallization.
10 PALAEONTOLOGY
1.6.2 Changes foulld in fossil assemblage (Fig. 1-3b) ·

The term fossil assemblage means a group of fossils (of one or more sp~cies) found together.
Thus a fossil assemblage may be homogeneous or heterogeneous. Ve1~1cally an assemblage
may occupy one bed or extend through several beds. Horizontal extension of an assemblage
may be few meters to several kilometers. The homogeneous or heterogeneous composition of
an assemblage is the result of ecologic relationship among the species within the assemblage
and this also results in the same or different preservational history. An 'in situ' fossil assemblage
is called life assemblage or biocoenosis whereas an assemblage drifted from its original site
by transportation and deposited elsewhere is called death assemblage or thanatocoenosis. Again
a death assemblage may be grouped into three categories : indigenous (assemblage not in
original site but deposited ·under same environmental domain); exotic, (assemblage occurring
in a different but contemporaneous environment) and remain (assemblage derived from other
rocks through reworking of sediments). All interpretations as regards palaeoecology or
palaeoenvironment should be ·done from the study of in situ fossil assemblage. However, it
becomes difficult to ascertain whether an assemblage is in situ or drifted. A drifted assemblage,
as it suffered transportation to some distance, may show disarticulation of skeleton, breakage
along shell boundaries, markings of mechanical wearing, rounding of skeletons and often size-
sorting. There.is always relative abundance of different species within an original assemblage,
but for a drifted assemblage the observed relative abundance may not represent their actual
relative abundance. Dominant species within an assemblage may be drastically reduced after a
lo~g transport for its particular type of skeletal-structure and composition that failed to bear
hazards of long transportation. It is almost impossible to say the actual relative abundance of
the different species from the study of a death assemblage.
1.7 USE OF FOSSIL

FossHs may be successfully used in geological science specially in stratigra h o ·
t tl h · f · P y. rgamsms
~e ~ons an y ~ an_gmg ro~ simple to complex types with increase of geological time So
1oss1 1
d bmay provide mformat1on about
. the time when they Jived ' about th e PIace where 1t· lived
:
an a out themselves, how they hved. The chief uses of fossils in ge 0 I · I ·
briefly outlined as follows : ogica science may be
1.7.1 Fossils ~sed in biost~atigrapliic classification of sedimentary rocks

As .fossils are changmg
k · with
· time i.e. in a vertical direction due to organic
·· evo 1utmn
·
d
se 1mentary roe m which they occur may be subdivid d · '
d d' d' · e mto a number of zones
epen mg on 1sappearance of older forms and appearance of newer organisms (~ d t . 1•
see Chapter 5).. or e a1 s
1.7.2 Fossils used in correlation and age determin ti'

· & • • a on oJ,r roek s
Id ent1ca 1 1oss1ls m separate areas may 1·nd' t .
1ca e same time and hence d'
layers may be correlated. This correlation ma be . . con:espon mg rock
depending upon the nature of fossil/fo . (~ Yd . local, regional or intercontinental
ss1 1s or eta1ls see Chapter 5).
1.7.3 Fossils used as indicators of past env,·r .
., ,mate
. onmenuc 1
Observation on recent . organisms shows th at terrestrial
. plants a d · .
to some sort of environments. Occurrence f h . · . . n amma 1s ar~ restricted
l o t eir fossils m older rocks may indicate a
..
.. r . . r .
.Ji.
. . .... '" ·~ ' .. .:.. -
A GENERAL ACCOUNT OF FOSSILS IJ
similar environmental condition. Plants or animals often bear on their body some traces
of the types of environment in which they live and these structures when preserved
within fossils, would be helpful in the reconstruction of palaeoenvironment (for details
see Chapter 5).
1.7.4 Fossils used in reco11struction of palaeogeography

Fossils may give valuable information about the distribution of past continents and oceans.
Some organisms are exclusively marine and presence of their fossils in older rocks
indicates existence of sea in that place. Migration of terrestrial plants and animals is not
possible from one continent to the other due to presence of oceanic barrier among them.
But if fossils of such plants and animals are found in rocks of certain age in widely
separately continents of today it definitely indicates that those continents were close
together or connected with one another by some land bridges at that particular geological
time.
1.7.5 Fossils used as evidence of prehistoric life and organic evolution

Study of fossils gives us valuable information about the origin and evolution of organisms
of today. Perhaps fossils are the chief basis of the theory of organic evolution of Drawin.
From the study of fossils it is possible to show how from primitive simple organisms
appear more and more complex forms of today. Study of fossils may directly demonstrate
evolution of amphibia from fish, evolution of reptiles from amphibia and evolution of
mammal and bird from reptile and so on. There are fossils called missing links like
/chthyostega (between amphibia and fish), Archaeopteryx (between bird and reptile). Fossil
can demonstrate the existence and/or dominance of some animal groups at particular time
of the earth history. It shows dominance of trilobites in Palaeozoic times, ammonoids
and dinosaurs in Mesozoic times.
1.7.6 Fossils used as economic tools

The practical use of fossil is its application in exploring some economic deposits. Many
resources are associated with some fossil organisms. Coal deposits are generally associated
with continental rocks containing abundant terrestrial plant fossils indicating existence
of a forest under warm and humid climate. Oil deposits are found mostly associated with
some microorganisms like foraminiferas/diatoms/coccoliths etc.
Chapter 2
SYSTEMATIC PALAEONTOLOGY
2.1 CLASSIFICATION AND NOMENCLATU~E ble types of animals and plants in

Biologists and palaeontologists have to study mnldumberad , It with two lines :
any scientific .mvest1gat1on
. · o f organism
· s . They shou e ed .
. . es of categories from sma11er to larger units
(i) Grouping of organisms mto seve~al. ty~ . . f ho logy anatomy, behaviour,
. · ·1 'f d d1ss1milanttes o morp '
observing the s1m1 an 1es an . . lled taxonomy or classification
physiology and many other characters. This aspect is ca
of organisms. .
. . ll h t 't ·11 be internationally
(ii) Naming of each taxonomic group sc1ent1fica y so t a I _w1 . loorecognized.
has two
This is called nomenclature. In other words, systematic b1ology/palaeonto oY ·
basic parts : taxonomy and nomenclature.
2.2 PHYLOGENETIC CLASSIFICATION . . .

In most of the natural classifications of plants and animals greater emphasis is given on_
evolution of different organic groups so that different subdivisions will indicate some sort of
evolutionary or phylogenetic relationship among themselves. A classification intending to reflect
such relationship is largely dependent upon some kinds of similarities and differences as one
may designed simply for indentification. This type of subdivision of organisms is called
phylogenetic classificatio11. In biological sc.iences it is possible to trace out such cha~acters
reflecting evolutionary relationship, between two diagnostic groups by direct observation of
living animals or plants. Obviously, smallest subdivision of one group have undergone only a
slight evolutionary transformation from the other similar subdivision of the same group such
as the difference between a man and an ape within the order primate. But greater the groups
have evolved far from their common source, greater would be their morphological differences
as primates differ considerably from ungulates although both are placed within class mammalia.
For a palaeontologist, erection of a phylogenetic classification is not so easy since the
information coded on fossilized shell does not give a complete record of the morphology and
life habit of that organism. There may be further complications for existence of such phenomena ,
like homeomorphism, dimorphism or polymorphism, taphonomic alterations. In spite of all these
difficulties palaeontologists always try to erect a phylogenetic classification for any organic group
as this is considered the most scientific and natural one.
Another technique called cladism has been adopted by some taxonomists for a phylogenetic
subdivis~on. ?f an organic gr~up (Henning, 1966). It is found that organisms usually exhibit
some pr1m1t1ve and some derived characters. Recency of common origin could best be shown
by possessio~1 of common derived characters. Generally, closely related groups must show
common de~1ved characters. For vertebrate, presence of a vertebral column is a primitive
character which does not reflect any close relationship among the different groups of vertebrates.
12
SYSTEMATIC PALAEONTOLOGY 13
But all mammals possess several derived characters (such as hairs on skin) common to aJI which
obviously indicate their origin from a common source. Cladism attempts to provide an
objective methodology for determining and showing graphically the recency of common origin
of related taxa, based upon primitive and derived characters. The assumption is that every parent
species always splits into two daughter forms after some definite time interval. The proposed
relationship, worked out from comparative morphology, is represented graphically on a
cladogram in which such dichotomous branchings are arranged in a series of nested hierarchies.
In graphical representation the taxa in question are arranged in lines. If species A and B are
related closely, the two are connected to a common point below the line. This junction point
of A and B represents their common ancestor. If species C is considered to be more distantly
related with A and B, it .is joined to a point still lower but connected with the common ancestor
of A and B. A cladogram is thus synonymous with 'classification. There may be two fundamental
problems with this method. There is no reason to believe that a large population of a species
will always split dichotomously instead of splitting into several species after some time.
Secondly, the elimination of subjectivity has not been possible in choosing primitive and derived.
characters especially when a large number of characters are invloved. However, most .
phylogenetic classifications assume some well defined models. The most common one is a tree
like structure, often called phylogenetic tree. The main character of this is that it is constantly
branching one and a branch never rejoins the ancestral branch or with some other branch. A
palaeonotologist can achieve a phylogenetic classification after a detailed study of fossils of an
organic group on its morphology (external and internal), habitats, temporal and spatial
distribution etc. But he cannot call the scheme proposed by him a final one. In fact, such a
classification is an expression of our knowledge of a given time and this may be modified in
future with further investigation and increase of our knowledge with discovery of new fossils
and new techniques for study of fossils.
2.3 PHENETIC (TYPOMORPHIC) CLASSIFICATION

It is far easy to classify fossil groups based on some observed morphological characters which
may or may not have any phylogenetic value. Such divisions may be based on one or more
diagnostic and readily identifiable morphological features. For example, based on dentitian, fossil
bivalvia is divided into a number of subdivisions. Such a method of subdivision is called
plzenetic or typomorphic classificatio11. However, there is a lot of subjectivity in selecting such
characters. Taxonomists have used different methods to minimize this subjectivity to get more
natural groups. Numerical taxonomists have tried to make the classification more objective by
opting for quantified phenetic resemblance as the reali~tic guide for natural grouping. According
to them, if many characters can be measured, quantified, specified, computed and the data are
represented by the use of the cluster statistics, similarities and dissimilarities between clusters
could be used to recognize similar or dissimilar groups. Numerical taxonomy may be useful
in some cases but subjectivity cannot be totally eliminated since the operator has to select and
give more importance to some of the characters to be measured for getting a meaningful result.
2.4 TAXONOMIC CATEGORIES

Linn_aeus in 17~9 used only 6 taxonomic categories viz. kingdom, class, order, genus, species
and variety from higher to lower rank. Later, two more categories are added : 'phylum' between
PALAEONTOLOGY
1-1
d enus At present, the most commonly used
,dngd m and class and 'family' between c;;derfian -7 ge;ius and species.
hierarchy are kingdom ph)'lum. class, o er, am, y, . . .
. . . rtificial and subJecttve but ideally they,
• h t· onomtc categories are a . . h'
Alth ugh most o f t ese ax . . s· . r spec·ies are grouped wit m a genus
... . · I t Ionsh1p. 1m1 1a ·· . . . '
as fur as possible.. reflect e,olut,onary re a d ·nto a class and similar classes into
. ·1 f ·1· ·nto an order, or ers 1
similar genera mto a famt Y, am, ,es 1 . l'ke subphylum subclass, suborder,
· such categories 1 '
a phylum. For some organic. groups I ss have been also proposed.
subfumil subgenus. superfamtly, superorder, superc a · . . .
.· . l ·od f l ·cal nomenc 1ature cover many P rocedures used . m dealing with
.
Intematl na _c es o ~oo og, es for formation of categories upto generic
higher taxononuc categones. There are several rul . d at least to the next higher category.
level. One rule is that a new category mu~t ~ ass1g~el . . this genus to an established
For · · ample, for creation of a new genus, 1t 1s essentm to assign
famil to which it belongs.
2.5 CONCEPT OF A SPECIES · . . . .

In any type of classification there should have a unit and species 1s take_n as the basic unit
of taxonomic subdivisions.
In biological science, a species may be defined as a group of individuals (natura.l po~ulati.o~)
which are actually or potentially able to interbreed among themselves produc_mg 1d~nt1cal
offsprings. Potential interbreeding suggests to consider only natural man1festat1on of
interbreeding.
This biological definition of a species is hardly applicable in case of a fossil population
where a knowledoe 0
of the phenomenon of reproduction is difficult, if not impossible to assume..
Thus morphological similarities among the individuals of a fossil population is the only critenon
for assigning them to a species. This would mean that species defined in palaeontology are
mostly artificial. In the phenomenon of homeomorphism unrelated genera/species may develop
identical external morphology due to adaptation to a similar life habit. Again individuals qf
one species may also exhibit contrasting morphology as in case of dimorphic/polymorphic
species. Therefore, in palaeontology most of the species are recognised as morphospecies.
Complication may arise from another direction. Most of the larger plants and anim.'l.ls at the
time of their fossilization become fragmented. Leaves, flowers and fruits of a plant were
detached from stem and fossilized separately. It is also difficult, sometimes impossible, to
reconstruct such fragmented fossils into a complete plant or animals. For this reason,
palaeontologists are allowed to recognise each of these fragmented parts as a separate genus
and species. These are called / orm genera and form species. For example Glossopteris is a
form genus and there are large number of form species within this genus. Again. since fossil
species also have limited distribution in the time dimension, from their inception to their eventual
extjnction, they are also called c/1ronospecies.
2.6 NAMING A SPECIES : BIONOMIAL SYSTEM OF NOMENCL.\TURE

Most matters concerned with the naming a new species are under the control of Jmemational
Commission of Zoological Nomen~lature which administers and updates the codes of zoological
nomenclature that ~re equally ap~hcable to. fossils and living forms and are also applicable for
naming all other higher taxonomic categories.
Regarding the establishmen! of a new species in the literature, the most important rules cover
the following topics : choice of a name, publication of the name with description and
illustrations, and designation of type specimens. Linnaeus ( 1707-1778) at first introduced the
binomical system of nomenclature in naming a species and this method is still accepted.
A species is designated by a name which has two parts. The first portion of the name
indicates the genus to which the species belongs and the second part is called the specific or
trivial name. This is called bionomial system of 11ome11clature. Both of these names are single
greek/latin words or latinized english words, so that it becomes understandable throughout the
world. The complete name of the species should be fol lowed by the name of the authors of
the species and the year of its first publication; e.g. the scientific name of cat of Bengal is
Fe/is benga/ensis Kerr, 1792. This means : the generic name of all cat is Fe/is; it has a number
of species of which one is 'bengalensis' (latinized name of Bengal); the author of this species
was Kerr who first published and described the species in 1792.
A few other rules as regards the naming of a species are as follows :
(a) The generic name must be initiated with a capital letter and trivial name with a small
letter. Both the names should be underlined at the time of writing and when published
they should be represented by italics.
(b) There is a considerable latitude of choice of words to be used as generic/specific names
such as Iatinized name of some eminent persons, name of some localities (usually the
place from which it is first reported) or a word indicating a diagnostic feature of the
species etc. For example :
Waagenophyllum indicum A Permian coral; generic name in honour of German
palaeontologist 'Waagen'; specific name from 'India'
(latinised) from where it ~as reported first.
Homo sapiens Homo (lat) : a man; sapiens (lat) : sensible or wise (name
of modern man)
Arachnophyllum murchisoni : A Silurian coral; arachnos (gk) : spiders web; phyllon
(gk): leaf-like; murchisoni : after Murchison, geologist
and founder of Silurian and Permian system.
(c) There must be one author who has erected the new species. However, more than one
person may participate in the official authorship.
(d) When an author of a species has assigned it to a genus different from that to which it is
correctly attributed the name of the author has to be enclosed within parenthesis after
subsequent modification of the name. For example, the modified name of the ammonoid
species Ammonities simplex Von Buch is Tornoceras simplex (Von Buch).
(e) Subgeneric name when introduced is written in the same manner as the generic name
and should be put in between generic and trivial name. For example, Lepidocyclina
(Eulepidina) dilatata. However, a subgeneric name may stand alone without generic name
like this : £ulepidi11a dilatata. Generic and subgeneric names can stand independently
but a trivial name cannot. The latter must be preceded by generic/subgeneric name.
Palae(Geo)WP-3
1
,, - _. a· _.
PALAEONTOLOGY
16
be written in the same manner
ecific name an d s h ou Id H
(t) Subspecific name may fo II ow a sp . . hanged to a trionom e.g. 01110
tandard
as specific name. In that case, th e s • ' bmonmn is c
sapiens sapiens. . . · d'vidual variants

. I for denoting s1mp1e m I
(g) The status 'variety' is often used m palaeonto o~y d ft the sepcific name by adding
. · may be designate a er .
or morphotypes. This vanety name . Ki / ritef goweranus f. ventrahs .
the word 'var' 'f' (form) or 'm' (morphotype); e.g. epp e . .
• ' . . cies the name of the author should
(h) At the time of publication of a newly discovered spe
be followed by 'sp. nov' or 'n. sp'. .
. ·mihr to some known species
(i) An author may find that his collected specimens are v~ry SJ • ' • t that species with
. I h h n assign his specimens o
but fail to confirm 1t. n t at case, e ca . . tic and specific
addition of such word 'cf' or 'aff' (meaning affinity) m between gene ·
name. For example, Halysites cf wallichi.
2 7 L AW OF PRIORITY-HOMONYMS AND SYNONYMS .

· cannot h ave.t wo d I ffierent names
• According to the codes of zoological nomenclature, a spe:1e~ .
or two different species cannot have the same name. This 1s also true for all ~thcr h1gh~r
taxonomic categories. There are a vast number of scientific names in the natural _history ~nd it
often happens that same species may get different names by different authors at different times.
Similarly, it also happens that different species may get identical names by different auth_or~.
Such incidents happen due to ignorance of scientists of each other works. In such cases, 1t JS
thus necessary to have some fixed standard by which the name which is to survive can be
- chosen. In the codes or nomenclature this standard is called law of priority. According to this
law, the name which has been proposed first has to be retained. For a particular species all
other subsequent (junior) names would be listed as sy11011yms. On the other hand lzomo11y111s
are identical names given for two .or more species. Here also senior homonym is retained and
the author/s of the junior homonym is allowed to change the name of his species in subsequent
time. Here is an example. Dunkar in 1869 proposed the name 'Fu.ms mayeri' for a modern
gastropod. Aldrich in 1886 described a Tertiary gastropod by the same name. Thus a case of
homonym arose and according to the rule of priority the name was retained for Dunker's
specimen. Aldrich subsequently changed the name bf his species as Fusus ottonis in 1897. Later
in 1904 Grabau concluded that Aldrich's species should be correctly assigned to the genus
Falcifusus, instead of Fusus. Then the .modified name of Aldrich species becomes Fa/sifusus
mayeri. Thus Fusus mayeri and Fusus ottonis would be its two synonyms.
2.8 TYPE SPECIMENS

A type in natural history represents one or more specimens which are used in describino
defining and illustrating a taxonomic unit. Type of a species is composed of individuals h· · 0 '
· bl · ·ri · . avmg
insepara e s1gni 1ca~ce m us1~g the name of the s_pecies, while the type of a genus should be
one of the man~ species belongmg to that genus. It 1s called ge11eritype. Normally once establish
the type materials cannot be changed. T.here are two kinds of type ·materials primary or
pro~erohl~pe anfdps~co11dary or hypolype. Pri1_nary type are generally selected by the original
autuor 1mse1 . nmary types of a new species may be of several kinds :
SYSTEMATIC PALAEONTOLOGY
17
(a) Holotype : A single specimen selected by the author ass uming it a~ an ideal form for hi s
new species is the holotype. The original description and illustration of the species gi en
by the author should be based on holotype.
(b) Syntype or Cotype : Instead of assigning one specimen as holotype, the author of a species
may select two or more specimens giving them equal status in describing hi s species.
These specimens are called sy11type or cotype.
(c) Paratype : Specimens other than holotype, which are formally designated by the original
author of a new species, are called paratypes.
(d) Lectotype : A specimen originally designated within syntype but subsequently chosen as
a holotype by the author is called lectotype.
(e) Neotype : If the original type material is lost or destroyed, the original author or any
subsequent author can select new types from the materials collected from same locality
and horizon. This is called neotype.
Secondary types, selected by any one other than original author, may be again of following
categories :
(a) Topotype : All subsequent specimens of the species collected from its type locality are
called topotypes.
(b) Plesiotype : Type materials of a species collected from any other locality by any
subsequent worker are called plesiotypes.
(c) Plastotype : Any cast of type specimen is called plastotype.
2.9 CLASSIFICATION OF ORGANIC KINGDOM

Within the organic kingdom there are tiny, often microscopic unicellular organisms to as
large as a whale. Many of these organisms or their related groups had left their fossils within
rocks. There are organic groups which are totally extinct at present and found only as fossils.
All these animals are subdivided into a number of phyla depending upon their morphological-
cum-structural complexity. Whittaker ( 1969) recognised five kingdoms of the organic world
which are : Monrea, Protista, Plantae, Fungi and Animalia. Mo11rea are simple unicelled
organisms with prokaryotic cells, Protista are unicellular organisms (both of animals and plants),
mostly microscopic with eukaryotic cells. Multicellular non-mobile algae and higher plants
(autotrophic) are included within the kingdom Pla11tae. The kingdom Fu11gi includes organisms
with cells without chlorophyll. The kingdom A11imalia comprises multicellular invertebrntes and
vertebrates which are also heterotrophic. Conventionally Animal kingdom is divided into 12
phyla. Table-2 shows this simplified scheme of classification of animals.
2.10 OUTLINE OF CLASSIFICATION OF PLANT KINGDOM

The two largest divisions of plant kingdom are based on the presence and absence of flower.
These two divisions are cryptogams (nonflowering) and pha11erogams (flowering). Cryptogams
include lower groups of plants without flower and seed. They reproduce mainly through spores.
r
PALAEONTOLOGY
TABLE-2
....~ SI.mp
. lified scheme of the classification of animals
Geological Modern Fossil
Phylum Diagnosis life-habit range representatives representatives
aqu ti Precambrian amoeba. radiolaria,
parasitic. to RecenL euglena. foraminifera .
foraminifera,
radio Iaria.
aquatic, Precambrian sponges calcareous or

manne to Recent. silicious
sessile spicules.
benthos.
aquatic Precambrian sea skeletons of

freshwater or to Recent. anemones, solitary or
manne; corals, colonial
lh; some anached or hydras, jelly corals.
pos.._~ hard cal- free living; fishes.
careous skeleton· solitary or
ilareral colonial.
symmetry.
ft animals; aquatic. Precambrian hook worm, unknown ,
trip blastic· crawlers to Recent. round worm, known only
bod., cavity with parasitic. flat worm. trace fossils.
ulh and anus;
i a eraI and
radial symmetry.
5.Annelida body soft; aquatic, Precambrian leeches, trace fossils as
coelomate terrestrial, to Recent. eanhworms. burrows,
segmented; burrowers, trails,
. organs crawlers. imprints;
differenciated·
scolecodonts.
radial symmetry.
6. Anhropoda se gmenced body aquatic, Cambrian to insects, trilobites,
ith jointed fresh water Recent. shrimps, ostracods
paired or marine, lobsters, estherids and
appendage ; calc/ terre trial, spiders, many others.
hitinou burrowers. centripeds.
kelewn· bilateral
Contd.
... . · - -----,----~-~-
.
- Phylum Diagnosis Life-habit Geological Modern Fossil

range representatives representatives
-7. Bryozoa unsegmented, marine Ordovician moss Fenestella
lophophore colonial. to Recent. animal s.
bearing; bilateral
symmetrical.
8. Brachiopoda unsegmented marine, Cambrian to lamp shells. Productus,
body covered sessile Recent. Spirifer.
with two unequal benthic,
valves on ventral often
and dorsal side; attached by
bilateral symme- pedicle.
try; lophophore
present.
9. Mollusca shelled animals; marine Cambrian to calms, snails, Nautilus, ·
unsegmented; sessile or Recent. nautilids, Ceratites,
shell bivalved or mobile, octopus. Belemnites.
univalved; burrowers,
bilateral fresh water
symmetry/ or terrestrial.
asymmetrical;
muscular. foot;
respiration by j
gill; often with -r
siphons.
10. Echinoder-
mata
bi lateral/radial
symmetry; spiny
marine,
sessile or
Cambrian to
Recent.
starfishes, sea echinoids,
urchins, sea crinoids,
I
skin; skeleton mobile cucumbers. blastoids,
below the skin benthos, cystoids. •
(endoskeleton) burrowers.
calcareous;
internal water
vascular system. '
11 . Protochor- bilateral; aquatic Ordovician acron worm, graptolites (?)
data primitive (marine) to Recent. amphioxus.
notochord; gill burrowing,
slit at some stage floating,
of life. auached.
12. Chordata permanent aquatic Ordovician(?) sea squirts, fossils of
notochord; gi II (fresh water to Recent. fishes,
slit at least at endoskeletons,
or marine), amphibias,
embryonic stage; teeth, skull,
terrestrial, reptiles, birds
dorsal nerve foot prints etc.
burrowing, and
chord, crawling, mamnnls.
endoskeleton swimmers,
ca-phosphatic. flying.
------------ --- ~--- --
PALAEONTOLOGY
20
· h fl d seeds Subdivisions of each of
Phanerogams include higher groups of plants wit ower an ·
these two groups are given below :
2.10.1 Cryptogamia
. d' · · b tween roof stem and leaf-
(i) Thallophyta : (Precam.-Rec.), thallus-hke; no 1stmct10n e • •
has two divisions :
(a) Algae : autophytic and chlorophyll-bearing plants; fossils available. e.g. Chara,
Diatom.
(b) Fungi: saprophytic or parasitic without chlorophyll; fossils available. e.g. Agaricus.
(ii) Bryophyta : (Precam.-Rec). leaf and stem differentiated but no root; fossils available;
e.g. mosses.
(iii) Pteridophyta : possesses distinct root, stem and Jeaf and internal conducting tissue but
no flower; a lot of fossil representatives; subdivisions are :
(a) Psilophytonae : (Sil.-Dev.). e.g. Psilophyton.
(b) Lycopodinae : (Sil.-Rec.). e.g. Lycopodium, Lepidodendron.
(c) Equisetinae : (Dev.-Rec.). e.g. Equisetum, Sphenophyllum, Schizoneura.
(d) Filicinae : (Dev.-Rec.). e.g. Fems.
2.10.2 Phanerogamia
(i) Gymnospermae : flower-bearing open-seeded plant; lot of fossil representatives within
Palaeozoic and Mesozoic rocks; subdivisions are as follows :
(a) Cycadofilicales : (Dev.-Jur.). e.g. G/ossopteris.
(b) Bennettitales : (Trias.-Cret.). e.g. Williamsonia.
(c) Cycadales : (Trias.-Rec.). e.g. Cycads.
(d) Coniferales : (Carb.-Rec.). e.g. Conifers.
(e) Cordaitales : (Carb.-Trias.). e.g. Cordaites.
(0 Ginkgoales : (Perm.-Rec.). e.g. Ginkgoites, Ginkgo.
(ii) Angiospermae : fruit bearing plants, seeds within the fruits; has two subdivisions :
(a) Monocotyledones : (Jur.-Rec.). one seed-leaf after germination; e.g. Paddy.
(b) Dicotyledones : (Cret.-Rec.). two seed-leaves after germination; e.g. Gram.
2.~l SYSTE~AT~C POSITION OF MAN IN ANIMAL KINGDOM

Kingdom : A01maha all animals.
Subkingdom : Metazoa all animals with multicelled structures.
Phylum : Chordata possessing notochord; dorsal nerve cord.
Subphylum : Vertebrata dorsal vertebral column.
Class : Mammalia bo_dy with _hair, nourishing of young by milk prod d . h.
m1lkgland m mother's body. uce wit m
- -- - - - ~
Order : Primate arboreal, brachiating digits, large brain .

Suborder : Anthropoidea monkeys, apes and man.
Family : Hominidae man and its immediate ancestors.
Genus: Homo man.
Species : Sapiens wise, sensible.
---,a, ~- --- --- ..··--- ....
Chapter 3
GRADE AND GROWTH OF ANIMALS
3.1 GRADE AND BODY PLAN : GROUPING OF ANIMALS INTO PHYLA

There are two aspects to differentiate the cooiplexity within animal's body. One is the level
of organization of body construction called grade and the other is the variation of the anatomical
body plan within a grade. Different grades are indicated by such criteria as whether the
organisms are unicellular or multicellular and for multicellular, whether they have two or three
well-differentiated tissue layers and whether they have a true body cavity (coelom) or not. At
any level of organization or grade, several distinctive body plans are common. Animals of same
grade sharing similar types of body plans are classed into a phylum. The different grades and
their body plans are as follows : (Fig. 3-1 ).
3.1.1 Unicellular grade .
It is made up of one-celled body layer or sometimes by only one cell. The body plans
are defined by the presence or absence of photosynthetic cells, cilia, and similar other
structures. They are taken as ancestors to all other organic groups and are included into
one phylum called protista. Some protistans may possess mineralised exoskeletons (e.g.
foraminifera, diatom, radiolaria).
3.1.2 Primitive multicellular grade
~~se ar~ early simpliest multi~ellular organisms possibly ancestors to other complex
hvmg animals. They are all extmct at present. However, among the Jiving multicellular
forms, sponges possess t~e most si~ple body organization, though they are not ancestors
to any other complex animals. Their body wall resembles a tissue but they have no well
defined organs. They are grouped as a separate phylum called parazoa (porifera).
3.1.3 Advanced multicellular grade
All high~r animal-groups possessing more than one cell-layer makin the bod II
grouped mto metazoa. g Y wa are
(i) Diploblastic grade : The body wall is differentiated int

t ·
ectoderm and inner endoderm. The body wall encloses a ; wo tissue layers, outer
· cavity) e.g. cnidarias. gu or enteron (gastrovascular
(ii) Triploblastic grade : The animals of this grade ·h ·

body wall : ectoderm, endoderm and a middle m:::te ara~terised by three-layered
types : rm layer. These are of three
(a) Acoelomatc : The phylum of this grade are worm-lik .

flat-worms. e without any coelom. e.g.
(b) Pseudocoelomate : They are also worm-l"k b

. b • e ut possess a fal b d .
formed m etween mesoderm and endoderm Th. . . f< • • se o y cavity
worm). The cavity is called blastocoel. · is is ound in nematodes (round
22
'. ~ . .~ - ··
GRADE AND GROWTH OF ANIMALS 23
(c) Coelomate : They possess true body cavity where coelom arises not from
blastocoel but develops as a space enclosed by measodermal layer mainly due
to splitting of mesoderm. This coelom is variously segmented, may be filled
up by organs like heart, gonad, kidney, stomach etc. Again the body plan of
,.
.
coelomate grade may be of four types :
(1) Amerous : Coelomic cavity is not segmented at all (e.g. sipunculida).
(2) Metamerous : Coelom is divided along its length into a number of
transverse ring-like segments each of which contains a pair of organs I
(phylum annelida and arthropoda). ~

(3) Pseudometamerous : This includes phylum mollusca, where coelomate
space is not divided and it does not envelop digestive tract. There may be
some duplicated organ systems within coelom.
(4) Oligomerous : This plan consists of a coelom that is divided into two/three
longitudinal segments each having separate organs and function. It has several
distinct groups of phyla : . (i) bracl,iopoda and bryowa with lophophore
(internal supporting organ) employed in feeding and respiration. (ii) ecliino-
derma with extensive series of canals, the water vascular system that function
locomotion, respiration etc. and (iii) cliordata with gill slits for respiration.
A scheme of subdivision of animals based on grade and body plan in given below
Grade and Body plan Phylum Examples
A. Unicellular grade Protista Foraminifera/Diatom
B. Primitive multicellular grade Porifera Sponges
C. Diploblastic grade Cnidaria Corals
D. Triploblastic grade i"
(a) Acoelomate grade Platyhelminthes Flatworms

(b) Pseudocoelomate grade Nemahelminthes Roundworms
(c) Coelomate grade
( 1) Amerous body plan Siphunculida Peanutworms -
(2) Metamerous body plan Annelida Earthworms
Arthropoda Crabs, Trilobites
(3) Pseudometamerous body plan Mollusca Mussels, Snails Squids
(4) Oligomerous body plan Bryozoa Sea mosses
Brachiopoda Lamp shells
Echinodermata Sea urchins
Chordata Vertebrates
Palae(Geo)WP-4
24
One layered
body wall
Unicellular grade
Primitive multicellular grade
L.s.
Triploblastic acoelomate grade
Triplobltilic coelomatc grade

FIG. 3-1 : DIVISIONS OF ANIMALS ON THEIR 'GRADE' AND "BODY PLA ..
(C.s. : Cross section; L.s. : Longitudina.l section)
''
_ ___. Growth line
Growth line
(a) PELECYPOD VALVE
(c) COILED SHELL

(b) CONICAL UNCOILED SHELL 1
· B. GROWTH BY ADDITION IN CRlNOID S1
A. ACCRETIONA~Y GROWTH OF MOLLUSC SHELL
FIG. 3 - 2 : TWO TYPES OF GROWTH IN ANIMALS
25
3.2 STUDY OF ONTOGENY

. . . d" .d . ulation Study of ontogeny reveals
Ontogeny is the hfe history of the tn 1v1 ua1 tn a pop · . h.
that individuals of different species exhibit different patterns of ontogenic growth: Even, w,t '"
. . . . . di
the md1v1duals of the same species different patterns an or ra e
. t of growth at different. stages
of ontogenac· development are seen. This · sometimes
· causes. d'111erence
" in external and/or internal
.
morphology among the individuals of the same population of a species. Thus sk~l.etons of two
individuals genetically related and living in identical environment when foss1hzed ~ay be
strikingly different simply because one was ontogenetically older than the ot~er. a~ the tim_e ~f
their death. Palaeontologists thus face a tough challenge in the identification of md1:1duals _wi_thm
a population belonging to a species for they may show a wide range of morphological vanatlons
due to the presence of individuals showing separate growth stages of the life, especially in such
case when the form is totally extinct at present. So the pattern of ontogenic development of
animals must be understood in order to recognise the range of morphologic variations displayed
by individuals of a species with the increase of their life-time.
3.3 TYPE OF GROWTH OF ORGANIC SKELETON

Skeleton-bearing organisms must show gradual growth of their skeletons to accommodate
the growing soft parts. As these skeletons are the chief materials of fossils we have to study
the effect of different types of ontogenic growths on the organic skeletons. For animals this
growth patterns of skeleton are of four types.
3.3.1 Accretion of materials

Most of the shelled molluscs increase their skeleton size simply by accretion of new materials
to the shell mainly to the outer or marginal parts of the skeleton [Fig. 3.2A(a-c)]. It has an
advantage of permitting continued use of skeletal materials deposited at earlier ontoganic stage.
It has the disadvantage that the form of the juvenile shell has to be retained or incorporated by
the individuals as part of the adult shell. Thus in such cases except size, the overall shape and
other broad morphologic features of the skeleton remain the same at any stage of life. This simple
type of growth is illustrated by coiled molluscs. For shelled molluscs (ga~tropods and cephalopods),
there is actually the growth of a hollow tube, uncoiled/coiled about an axis and widening towards
,(he aperture. Here new shell materials are constantly accreted along the apertural margin.
<?rowth by simple ma~ginal accretion of ne~ mate~ials is also found in other animal groups
particularly where shell ts external, representmg a smgle piece and serves in protection and
muscle attachment such as growth patterns of solitary coral, bivalved mollusca and brachiopoda.
One may consider the individual valve of a pelecypod or brachiopod as .,
. . . . ..
·1 d ho 11 ow cone
.. co1 e
openrng o~tward; their co1le~ m1t1al part, which f?rms a very s mall cone is called umbo. The
cross. sect1~n of each valve 1s marked by successive growth lines parallel to the shell'.
margm which are seen on the outer surface. s outer
However, the new materials are added not only along the t. •
as covering as a thin layer over the internal surface of shell o:~r edge of _the valve but_ ~lso
internally. This causes gradual thickening 0 f th , 1 ' so t ,lt growth lanes are not v1s1ble
size (Fig. 3-2b, c). e va ve towards unbo along with its increase of
I
PALAEONTOLOG Y
\
26
I
I
3.3.2 Addition of new skeletal part found in those organisms whose skelet?n~ cons_ist of
. A common method of skeletal growth, addit' of new skeletal parts. This as exhibited
several parts articulated softly or tightly, is the f '~7e echinoderms). The growth takes place
O
by the growt'h pattern of crinoid stem (a group shessba of the calyx. As new columnals are
I nals along t e ase
by successive addition of new co um d' d more down the stem. Further smaller
. us ones are asp 1ace
added at the calycal base, the prevao . t' plates This causes increase of length
I and adult pre-ex1s mg . . .
columnals appear between arger .. Fi _ B). Different crinoid species may mod1f
32
of stem as the animal grows by addition ( ~ th . different functional problems of the
this basic growth model in order to acco~m artat:he:: large columnals alternate with small
life. For example, parts of the _stem (up~ f) rt where all columnals are large and of
ones becomes much more flexible than_ its ower p~ is necessary in crinoids for adjusting
equal size This flexibility of stem at its upper pa . fi d
themselve~ with the water current and also to collect food as suspension ee ers.
3.3.3 Moulting · . · f th Th ·
Most of the arthropods including trilobites exhibit another basic mechanism o grow · 15
is periodic shedding off the entire skeleton and formation of .a new on~ t~ accommodate the
enlarging internal soft parts. This is called moulting or ecdysis. H~re w1thm_ the old skeleton,
the soft parts of the animal grow to a larger size and new cuticle which .underlie the old skeleton
hardens. Ultimately the animal in the larger size comes out by rupturmg the old skeleton and
gradual growth of a new skeleton. Growth here is thus rapid and episodic.
Growth by moulting has one clear advantage over the accretion and addition. Here the skeleton
of the juvenile animal need not forms the part of the adult body. Thus the adult has much freedom
to change not only its size but also its shape and some broad morphologic features after every
stage of growth. This, however creates a serious problem to palaenot61ogists to identify individuals
of the same species within a fossil population. Again, this type of growth has a distinct
disadvantage, that is its vulnerability to damage and predation during the period when the new
skeleton is yet to be sufficiently hardened immediately after each moulting. Moreover, the animal
has to extend considerable metabolic energy in replacing its entire skeleton at intervals. A
modification of this mechanism is found in the growth of some bones of higher vertebrates. The
form and structure of these bones change without rejection of old skeletal materials as they increase
in size. This form of growth has an advantage over moulting system but not has any disadvantage
that the organism is without a hard skeleton for some period of life.
3.3.4 Mixed growth

Many organisms however: display a combination of these two/three basic mechanisms of
growth. Crinoid stem, for example, shows a combination of two mechanisms : addition of new
parts and accretion to the old parts. Another example is the growth system of echinoderm
skeleton. Here addition of new plates within ambulacral and interambulacral areas is the principal
means of growth of skeleton size. At the same time all the existing plates grow by periph ral
or marginal accretion.
Coiled cephalopods also exhibit a combination of two modes of growth. In this case the
main shell grows in length and size by continuous accretion in much the ·same manner as found
27
GRADE AND GROWfH OF ANIMALS
. . ·te different biologically. The
in a gastropod shell. But these two groups of org.am~m are gm h t But the soft animal
soft animal of gastropod occupies the entire shell which is hollow throug ou .' h I t h mber
of cephalopod lives in what is known as the ~ody chamber which occupies t e as c a
. . h b · however separated from
covering l of 1 of the last whorl of the coiled shell. Body c am er 1s
2 4 . . · · · t moves outwards and then
the rest of the shell by a septum. As the animal mcreases m size, 1 d
. . wall or septum 1s
a new part1t1on . added penod1ca
. · 11 y to accommoda te the animal with the· . forwar
.
growth of the shell. This periodic addition of new septa has made the cep~alo~od she]~ internally
multichambered but all of them remain empty except the last one which 1s occupied by the
living animaJ.
3.4 GROWTH RATE OF ANIMAL

The rate of growth is different in different groups of organisms. Such differences are largely
hereditary; that is, they are under genetic control and the pattern of growth have been produced
by evolution. However, in all organisms there is a certain amount of variations in the rate of
growth which are not under genetic control but depends upon some specific conditions imposed
by the physical and biological environment. The terms "norm of reaction" is sometimes used
to express the amount of nongenetic variations possible in a morphologic feature or in the growth
rate of that feature.
In nearly all organisms the growth rate varies with time. That is, it changes during ontogenic
development. In fig. 3-3 the size of a hypothetical organism is plotted against time. It shows
that size of the animal in question increases at a slow rate at the beginning; it becomes much
rapid at the middle stage and in the final stage (adult stage) the growth rate is very slow, almost
nil. However, most of the plants and many animals do not follow this simplified and generalized
model of growth. In all the plants and in many invertebrates, growth continues throughout their
life-time but usually at a reduced rate making it difficult to recognise their true adult stage
(Fig. 3-3a). But in case of mammals and other higher vertebrates the adult stage of life is almost
without any significant growth and the upper part of the graph (Fig. 3-3b) virtually levels off.
When the growth rate of an organism is measured, the data obtained seldom yield a perfect
curve because of variations of some non-genetic factors.
3.5 ISOMETRIC AND ANISOMETRIC GROWTH

Most of the ontogenic study requires measurement of growth rate of one morphological
attribut~ in rel_ation to that of anot_her. ~or example, one can measure the change of length of
a shell m relat10n to the ~hange of its w_1dth and th~ growth relation between these two attributes
may be expressed graphically. From this, two basic types of growth can be defi d . · ·
d · · If h · 1ne . isometnc
an amsometnc. t e ratio of growth of two attributes of an animal does t h h
·
lh roug hout its · d . . no s ow any c ange
ontogenac evelopment 1t 1s called isometric growth (Fig 3-4 ) B ·f . .
of the growth of two attributes changes with ontogenic development it is : ·u ut, I • the rati_o
growth (Fig. 3-4b). In isometric growth, there is no overall cha f a ed ams~metnc
the animal grows uniformly in all directions. If growth of l h nge ~ shape of an animal as
isometric growth is plotted, it will form a str . ht . engt a~d width of an animal showing
(Ftg. 3-4a). But .m an amsometric
. a1g 1me passing throu h th . . .
growth the anim I h . g e ongm pomt
dimension with ontogenic development and h a . s o~s change of its growth rate in different
ence mvanably shows a change of its shape with
_ _ -=:.it.: _ _ __ _____,____,_....______,_.. _ _
-- "·,
~---
..
~/
..
,.
,..,
(
~
I I /
§
§
(I)
{I)
I - -
II
-
./ ~..,.
,,
'
___________,;;:._ TIME
TIME
{a) NON CONTINUOUS GROWTH (hypothetical animal)
{b) CONTINUOUS GROwnt (hypothetical plants)
FIG . 3 - 3 : CHANGE OF GROWTH RA TE

=
••
1
$ 1
g
i
i
"'::
:::,..
~
~
a
~
LENGTH . ::::,,., LENGIB----~- -.:
(a) ISOMETRIC (b) ANlSOMETRJC 2
FIG . 3 - 4: ISOMETRIC AND AN ISOMETRIC GROWTH OF ANIMAL t
-.c-
~.
7
~- -- . . _._. ~ · --~-:.L.A.· ---
• ~• _[ij/1 ' -
29
time (Fig. 3-4b). Anisometric growth is more common in animals and it can b~ expressed_ b:
the statement of Raup ·and Stanley (1985) that " s hape c h ange durina
o
ontogemc
·
growth
.
1s ,1
rule rather than an exception" and many authors incline to use the term al/ometry for this type
of anisometric growth.
3.6 CAUSES OF ANISOMETRIC GROWTH

The modern concept is that the change in form during ontogeny becom:s a p~rt of the
hereditary materials because of their functional value. Any change of shape of an animal_ or a
structure within the body of the animal along with ontogenic growth must have a funct1o_nal
and adaptive explanation. Sometimes these changes become rather abrupt as found dunng
metamorphosis of an insect which can be explained by sudden change of its mode of life. A
clear example is the development of adult mosquito from the pupa stage or tetrapodal frogs
from the aquatic tadpoles. Let us consider change of shape of some parts of an animal body
with response to the growth of animal itself. As a most simple case Jet us consider an imaginary
terrestrial vertebrate showing isometric growth in all its linear dimensions. If this animal after
a particular time would be able to double its linear dimensions it would cause an eight-fold
increase of the volume or weight of the animal. The femer bones of the legs of a tetrapod
have to bear the weight of the animal and if this bone maintains the same growth rate, the
cross-sectional area of the bone will increase as the square of its linear dimension. As the
strength of a bone increases with increase of its cross-sectional area, it is clear that animals
body even maintaining an isometric growth, shows increase of its weight more rapidly than
the increase of effective strength of its leg bones that have to support the entire body. Thomson
named this rules of scaling, inequality of growth of two parts of an organic body as "the
principle of similitude". In this case the animal can solve this problem only by increasing the
cross-sectional area of the leg bones more rapidly than the increase of its linear direction, and
this will result an anisometric growth of leg bone and consequent change of its shape. The
bones become thicker at its upper and lower end to increase areas of muscle-attachment. More
thickening compared to its length often causes torsion of the bones. This gives the bones more
strength so that they can support the growing weight of animal and function properly. This has
occurred during ontogenic growth of many terrestrial vertebrates including man. A few animals
to avoid the problem of similitude, grow by addition of identical structural units instead of
modifying the pre-existing one. Growth of a sponge is not caused by increase of the size of
the.flagellate _chambers that constitute its body but by addition of new such chambers of original
size. Vertebrate lungs also grow by this mechanism. Accretionary growth of a bivalved mollusc
is caused by addition of materials along leading edge of the valves but at the same time this
causes the thickening of older portion of valves. If new shell materials would not be added to
the interior of the shell along ~ith I_ateral and margin~! accretion, the shell would not only
becoi:ne larger but at the same time 1t would be _too thm and fragile to support the animal's
~ro;in~ soft parts. ;11~re ared~;ny changes f~nd m some structures of some animals especially
mh ob~s1 1 g_rou pfs w !c aref ht 1cu 1t to explain as we do not have sufficient knowledge about
t c 101og1ca1 unctions o t c organs/structures of the animal concern.
Chapter 4
SPATIAL DISTRIBUTION OF ORGANISMS
4.1 STUDY OF BIOGEOGRAPHY

Biogeography is the study of geographic distribution of organisms. Palaeobiogeograplzy
obviously deals with the same aspect except the plants and the animals concern are belonging
to the past geologic times represented by fossils.
A careful observation reveals that distribution of present plants and animals on the earth
surface has a close relation with the different aspects of geography such as climate of the earth,
movement of water and wind and other physiographic features like surface relief, latitudinal
distribution of land and sea etc. The ancient geographic conditions thus can be reconstructed
from the nature of the distribution of past organisms simply by applying the principle of
actual ism.
4.2 CAUSES OF CLIMATIC VARIATION .

A basic understanding of the present climate of the earth. is essential in the study of
biogeography. Ancient climate can be reconstructed on the basis of our knowledge about the
recent climates. The temporal changing pattern of the palaeobiogeography must also indicate
the pattern of the change of climate of the earth with time.
Temperature gradients and consequent climatic variations on the earth surface develop
because the sunlight shines the earth surface at a lower angle near the pole than near the equator.
Thus a beam of sunlight on a given cross-sectional area would spread over a larger area in the
pole providing less heat per unit area. This is just reverse towards the equatorial region. The
temperature of the earth is thus increasing from higher to lower latitude i.e. from pole to equator.
It is also known that pattern of rainfall is also related to the local distribution of temperature.
This can explain high temperature along the equatorial belt consequently causing rise of hot
air from this belt which on cooling at the higher level releases its moisture on either sides of
the equator. As higher air is also relatively cooler than the surface air, at any latitude, region
with higher altitude tend to be cooler. When such air moves up along the upside of the mountain,
it leaves most of its moisture on that side resulting a rain-shadow zone of dry climate on the
lee-side of the mountain. Further complexity is added by real nature of the earth's motion which
has a profound effect on atmospheric circulation. The primary effect is coriolis force. For this,
wind or water current coming from north pole towards equator curves in a western direction
instead of following a straight line. This will be just reverse in the case of wind movement
from south pole to the equator. For the same reason, 'trade wind' does not move in a straight
way from pole to equator. Wind also is of prime importance in determining the pattern of oceanic j
circulation. ~
1
4.3 GEOGRAPHIC DISTRIBUTION OF ORGANISMS-DISPERSAL

Observation of organisms in space at a particular time reveals that organisms are limited in
distribution within a definite geographic province. For example, the Pleistocene mammals of
30
• ••
- ~
-" \
SPATIAL DISTRIBUTION OF OR ANISMS 31
Asi3 3nd Australia differ radically. althou ,h. th •y w •r • adapt ·d 10 u similar environment. The
answer to this puzzle is 1ha1 lhe two ar ·as w •re s ·parnl d by s uways thar stood in between
these two continents as a geograpl,ic ba"ier. Oco 1ruphic distri bution of or ,unisms in horizontal
dimension of space is of great importune · to und rstund the geographic pattern and di. rributi n
of land and sea of the earth al a particular time.
Whenever a new pecies evolves in an area. it tends to spread our t all distant localities ro
avoid local competition with the increase of population. It then migrates from its point of rigin.
had the route of migration been open and available 10 curry it uninterrupted ly. This is ca lled
dispersal. But due to the presence of various types of nalurnl geographic barriers. the distribut ion
of a species is generally restricted. In general, easier intcrmigration of organi ms takes place
in between two areas of similar environment without any barrier and stronger the barrier greater
lhe contrast of faunas and floras exists between them.
4.4 SOME RELATED TERMS ON 'DISPERSAL'

Faunal/Biogeographic province : Biofacies variation can be grouped into larger geographic
areas covering substantial portion of the earth having a
broadly similar environmental condition. These are called
biogeographic provinces.
Co mopolitan A species which is universally distributed i.e. geographically
widespread. A general term that lacks genetic connotation.
Stenographic Species having a limited geographic range.
Eury geographic Species having a wide geographic range
Disjunct eurygeographic Species with wide geographic di stribution but occurs as
patches in favourable habitats.
Endemic Those forms which are restricted or exclusive to a particular
geographic locality such as in a single province or in a part
of a province.
Indigenous : · Forms originally descendant of a particular locality.
Immigrants Forms derived from a foreign locality but are contempora-
neous.
4.5 FACTORS CONTROLLING DISPERSAL

. The common _factors w_hich _help dis~rsal of organi sms are : (i) direct swimming (pos:ibl.
in case of aquat1c-nek~on1c ~namals). (11) attachmc~H to some moving objects (epiphmktons/
pseudoplanktons), floating animals (plank1ons). mobile lurvue (mcrnplunktons) and blowing ut
by wind (microscopic spores and pollen).
From the analy is of the above mentioned point it is clear thnt plnnktons and nektons of
the sea have greater chance
. . of wide dispersal · · B,enth os, are, ·Ill gcncrnl less mobal· • und usually
how a na~ow and li_m1ted ge~grnphic distribution. In spite of this fact, many inv~rt hrate
bent hos which are vag1le or ~ess1le show remarkable geographic distrihution for the r • ,son 1hn1
many of them produce mobile larvae (meroplank1ic).
Palae(Gco)WP-S
32 PALAEONTOLOGY
In general, marine organisms face less effoctive geographic barriers and so they can have
wide distrihution through sea water. On the other hand, terrestrial organisms have to overcome
variable types of geogrJphic barriers in the paths of their dispersal which they in many case
fail to do and consequently their distribution becomes much limited.
4.6 BARRIERS TO DISPERSAL

Barriers to dispersal result in isolation of organic populations and destroy chan~es. of
interbreeding between them. The barrier may be posed as much by submarine ridge or similar
landmass standing out on the sea-floor in case of marine organisms. It is posed by a se~ between
two landmasses in case of terrestrial organisms. Even within a huge landmass sharp differences
in climate (temperature), food supply and other factor may control dispersal of a pai1icular
organism. Sharp differences in temperature, salinity in two different zones o~ a s~a also cor~tr~I
dispersal and especialfy modify the course of meroplanktonic larvae. Considering these, it is
evident that agreement in past land floras and faunas between two continents (no~ separated
by a sea) at a particular time has been invoked to justify that either the two contm~nts wei:e
directly joined together or were connected by a land-bridge or like features at th_at time. ~his
is the basis of the idea of existence of a Go11dwa11a superco11ti11e11I which during Penman-
Mesozoic time yielded similar land plants and vertebrate fossils.
The discontinuous distribution of organisms may be brought about either by extinction of
the intermediate faunas or by bringing a barrier between two distinct areas. Often the two areas
separated by a barrier (say, two continents separated by an ocean) may be bridged allowing
free or partial intermigration of organisms of the rwo separatl!d lands. Such connecting land
bridges between the two continents may be classified into three types viz: corridors, filter
bridges and sweepstakes. A corridor is a broadly continuous connecting bridge existing over a
long time so that it permits an extensive interchange of floras and faunas of the two continents,
such as the connection now exists between Asia and Europe. A jilter bridge is more temporary
in duration and more restricted in extent and it allows some of the organisms holding some
particular characters and behaviours to pass through it. The Bering Strait acted as a tilter bridge
for some mammals during Plio-Pleistocene between North America and Europe. Only those
mammals who able to cross this bridge were capable of rapid movement and withstandino cold
weather. A sweepstakes bridge on the other hand, does not involve migration acro;s the
connection but rather it depends upon accidental transportation of some suitable smaller-sized
floras and faunas. Corridors and filter bridges operate equally well in either direction but
sw~epstake bridges operate only in one direction. Such migration takes place at present between
Afnca and Malagasy which allows accidental migrations of such small mammals like rodents,
shrews, small monkeys from Africa to Malagasy.
4.1 TYPES OF CLIMATES AND RELATED VEGETATIONS

Regional climates are intimately related to the large scale current system of · d . d .
. 'b · f · . wm
Th e d1stn ut,on o reg,ona1 c1,mates and related vegetation on the present e· rth · . ·h an ocean.
·
· · I h . . . . a 1s s own rn
Tab Ie-3. Not surpnsmg. . .
y, t e worldwide d1stnbut1on of certain types"· of terr ·t · I
. . . .
·
es na vegetation
tend to reflect t~e d1stnbutmn of thetr res~e~t,ve climatic condition. This relationship would
suggest that fos~,l plants could ~ u_seful md1cators of the ancient climatic conditions of the
earth. To a considerable degree this 1s also true for terrestrial vertebrates.
:n
TABLE-3
Climates and types of vegetation
Name of climate Type Vegetation
1. Wet equatorial belt equatorial air being heated near surface rises, equatorial and
cools and losses its moisture causing rainfall tropical rainforest.
uniformly throughout the year.
2. Trade wind littoral trade wind along the coastal hills rises up, rainforest.
cools, causing very heavy rainfall on the sea
side of the mountain.
3. Tropical desert cool air descends at tropics but becomes plants scarce;
(Lat. 20°-30°) warmed up again picking up more moisture desert to
from the surface preventing rainfall. semidesert.
4. West Coast desert westerly wind encounters interior mountain semideserts
ranges to the west and loses its moisture on steppes.
stoss side developing desertic climate on the
further west side of mountains.
5. Tropical wet-dry areas where moist trade wind rises up and tropical forest,
climate (East coast of crosses the hill, cools and sweeps in land woodlands and
continents between causing heavy rainfall. savanas.
25° to 30°N latitude)
6. Humid sub-tropical comparatively warm and humid. Summergreen
(comparable parts of deciduous forest.
S-latitude)
7. Marine west-coast moist wind prevents westerly wind from evergreen forest.
and mediterrenean coming from ocean, which thus rising up and
(west coast of dropping its moisture on the coast.
continents of middle
latitude 35°-65°N)
8. Humid continental steppe/grassland.
9. Middle latitude summergreen
desert and steppe deciduous forest.
Continental subarctic cold climate of continents and oceans near needle-leaf forest.
and marine subarctic arctic
Tundra continents at high latitude of north, ice arctic, tundra.

covered for greater part of the year.
Ice cap permanently ice covered areas of poles and without any
other high altitude areas. vegetation.
- - - -:-s
.=w.- -- 'l"" .,_ ' •
~
I
34 PALAEONTOLOGY
4.8 BIOTIC DISTRIBUTION

Biome can be defined as the total biotiG _association (both plants and animals) over a lar~e
area (say a continental landmass). Each 1Jiome is composed of a large number of organic
communities containing population of more than one species. It should be remembered that
though a particular climate may be found in two widely separated landmasse_s yet th~t does
not yield identical biomes in those areas. This is because of the presence of barriers to dispersal
of various kinds of organisms specially for plants and terrestrial vertebrates. Thus id~ntica~ fos~il
plants and to some extent terrestrial vertebrates belonging to a particular geological time m
two presently separated continents definitely indicate that those two continents were cl_ose
together and under the influence of same climatic condition during that particular geological
time. The significance of the distributions of some present day taxon association can also be
understood more clearly if we can trace geographic condition including climatic features of
the past and establish the change that had occurred. Marine realms present other sets ?f
problems. Pelagic forms tend to be more widely distributed than benthic forms. Some benth1c
species may have also wide· geographic dispersal through their nektic/planktic larvae.
4.9 LATITUDINAL CHANGES AND TAXONOMIC DIVERSITY

One conspicuous phenomenon of a present-day taxon as regard its dispersal, is its tendency
to display increase of species diversity towards the equator. The result is an increase in average
species diversity in communities in the same direction. There are however some exceptions.
Origin of this diversity gradient has been widely debated. Some hypotheses that have been
advanced are based on the assumption that all environments tend to be saturated with species
and this saturation level increases towards the equator and decreases towards the poles. One
suggestion is that higher number of taxa arises towards the tropics because of severity of living
conditions in the polar region. Another assumption suggests that rate of speciation is higher
towards the lower latitude. Another assumption is that environments of both high and low latitude
may potentially support the same number of taxa but the process at higher latitudes is frequently
interrupted by large scale environmental hazards so that the potential carrying capacity in these
regions is never approached. An idea expressed that taxonomic diversity increases towards the
equator because of increase of complexity of the physical environment towards this direction.
One important factor causing latitudinal variation of organic diversity in the sea is CCCD or
Calcium Carbonate Compensation Depth of sea water which determines the rate of dissolution
of CaC03 in sea water. Amount of dissolv~d Ca~03 in sea water primarily depends upon the
temperature of water. When other factors mcludmg the depth of the ocean remain the same
~emperature of w_ater is increasing from the poles to the equator. The solubility of caco i~
1
inversely proportional t~ the temperature of sea water. Thus warm equatorial sea water ·has
greater power or supply mg _CaC0 3 com~are. to its power of CaC03 dissolution. That is the
reason why calcareous organisms are dommatmg towards the lower latitudes. It seems . . t
. d' I d' . . d' . fl apparen
that lat1tu ma gra 1ents m organic 1vers1ty re ect a variety of interrelated factors that are not
yet fully understood.
4.10 CLIMATES OF THE PAST

It is not easy to evaluate the cli'!1atic changes that occurred in the geological past. There
are a variety of tools for reconstructing ancie_nt climatic conditions, but mostly they give only
a partial picture and every method used has Its own limitations.
35
4.10.1 Botanical evidences
Flowering plants are particularly useful in the reconstruction of climate since Cretaceou
period (i.e. since their appearance). Most of these fonns appeared during Cretaceo-Tertiary times
and many of them have their successful representatives in the present earth . So observati on of
these present-day flora and their variation in distribution in response to the present climatic
belts of the earth will help us a lot to interprete the climate of the Cretaceous and Cenozo ic
period by observing the distribution of similar .floras in their foss il-fonns.
Careful and large scale observation of the present day flora have show n th at so me
morphologic cum anatomic features of different plants may be useful cl imati c indicato L
regardless of their taxonomic position. Utility of some leaf characters has been sum me ri ed by
Dorf (1970) which is shown in Table-4.
TABLE-4
Leaf morphology and climate
Leaf characters Climate
I. Non-entire margin (toothed, indented, lobed) Cold
2. Entire margin Tropical
3. Large leaf, thick leaf, pinnate venation Tropica l, hum id
4. Small leaf, palmate venation Cold
5. Palmate compound leaf Tropical
6. Needle like leaf/pinnately compound leaf Cold
7. Apex with drip-point Tropi a l ith hea y rain fa ll.
When a climatic belt shifts, the corresponding flora also tend to shift with it. In some cases
the flora could be extinct especially when the shifting of a cl imatic be lt is too abrupt or severe
for successful floral migration or if the required climatic conditi o n for the ex isting fl o ra tota lly
disappears. Thus shifting of a climatic belt with time can be read from the nature of fo sil
floral migration. Dorf has shown that overall climate of western Europe, we tern U.S .A. had
changed from Palaeozoic onwards but one thing is clear that for these two region; cl imate has
undergone a net cooling since Early Cenozoic.
Even for greater part of Mesozoic (Triassic and Jurassic) whic h predate the ori gi n of
angiospenn flora, gymnosperm and pteriodophytes are employed in the reconstructi on of cli mate .
Certain ferns and gymnosperms belonging to cycads are largel y re str1cted to tropi ca l and
subtropical areas of today and probably lived in the similar condition in the past. Con ife rs , o n
the other hand, are largely restricted in cold-climatic temperate zones. Annua.I rings in th e tree
trunk are conspicuous features of temperate latitude but are found weakly de e loped wi thin
trees of humid region with heavy rainfall throughout the year. At least a season al vari ation of
rainfall in essential for different rate of growth of vascular bundles that produces annul ar rin g
in stem. There may be however some complications such as : some fl ra of equatori al be lt of
higher altitude sometimes resemble very much with the flora of arctic region of lo\! er e le ation.
36 Pt\],AEONTOLOGY
Not only leaves but also pollen and spores which may be often treated identical to the level
of form genera are of great value in palaeoclimatic interpretations especially in the interpretation
of climates of the Quaternary period. Study of distribution of modem spores and pollen has
shown that this often faithfully reflects the general geographic distribution of the modern plants
(Davies, 1967). Palynological data from North America clearly indicate that towards the end
of Pleistocene, flora successfully shifted northward and southward with the shifting of isotherms.
It also appears that large part of continental shelf of North America of today were emergent
during Pleistocene and supported a terrestrial flora.
4.10.2 Information from marine life

Some marine invertebrates are also of considerable value as indicators or temperature regime
in the ancient seas. The most useful living group in this regard are hermatypic or reef corals.
These colonial animals live in a symbiotic association with a group of algae. Zooxanthelle in
their skeleton. For this reason these reef corals are not only restricted upto the photic zone of
the sea but also grow near 30° latitude around equator (tropical seas) where the water
temperature seldom falls below 18°C. It appears apparent that similar temperature limit have
obtained since Triassic when appeared reef-builder scleractinian corals. Due to seasonal variation
of temperature. the rate of secretion of CaC0 3 by solitary corals may vary resulting in some
minute growth rings on the epitheca of coral exoskeleton. This is generally more pronounced
in non-tropical condition.
The record of some fossil planktons within the sediments of the sea may provide some
valuable information for the reconstruction of Cenozoic climate. In particular. this record sheds
light on the pattern of waxing and wanning of global temperature that caused by advanced and
retreat of Pleistocene ice-sheets. Coiling direction of some foraminiferas (planktic) that raindown
on the sea-floor after death is also found affected by the influence of temperature. Planktonic
fonninifera species Globorotalia menardii is found in the equatorial region. The abundance of
tests of this species relative to those of other forminiferas reflects climatic history and
temperature fluctuation in the sea in Pleistocene. (Ericson & Wollin. 1968). During glacial
episodes the relative abundance of this species is very low whereas during interglacial episodes
the species became relatively more abundant than others. Fortunately, in this case the accurate
chronology is provided by the record of reversal of magnetic polarity within sedimentary rock.
Some planktonic foraminiferas are found exhibiting change of the coiling pattern of their tests
with fluctuation of temperature of sea water in response to global climate as seen during
Pleistocene glacial and interglacial periods. The planktonic forminifera species Globigerina
pachydema are found producing dominantly dextrally coiled tests in warm water sea but mainly
sinistrally coiled tests in cold water.
Varying ratio of isotope 0 18 and 0 16 in foraminiferal test provides an independent method
for distinguishing Pleistocene glacial and interglacial periods. During evaporation of sea water
more isotopes of~ 16 are lost as they are relatively lighter than 0 18 . However, these 016 isotopes
come back to sea water through normal hydrologic cycle keeping 0 181016 ratio more or Jess fixed
in sea water. Normally, organic shells with oxygen show same isotopic composition as that of
the local sea water. But during a glacial period an enormous volume of the evaporated sea water
is transformed into permanent body of ice in which more, 0 16 isotopes remain trapped when the
18
cold sea water becomes enrio'~ed in 0 isotopes causing increase of ratio of 0181016 in water
SPATIAL DISTR/LlUTION OF ORGANISMS 37
and also within the organic skeletons produced at that time. Emiliani ( 1966) indicated from the
analysis of calcareous foraminiferal tests, the recurrence of glacial and interglacial pe riods
throughout the Pleistocene Ice Age. Tests of foraminifera usually show same isotopic composition
as that of local sea water. Sea water during cold climate is generally enriched in the heavier isotope
018 because during warm climate water evaporates more rapidly and more lighter isotope 0 16 is
Jost at that time and subsequently trapped within ice.
4.10.3 Evidences from continental sediments

Observation of present-day depositional basins at different climatic zones and the nature of
the deposited sediments in them leads to some generalization that can help us to reconstruct
past climate. A correlation between these two has been shown in Table-5.
TABLE-5
Sediments and climate
Sediment characters Climate
1. Tillites, varves, green shale, unaltered K-feldspar in sandstone, cold, glacial.
few fossils.
2. Laterites with brown nodules/altered K-feldspar in sandstone. moists tropical.
3. Nodular layered deposits of calcium carborates (caliche). warm, semi-arid.
4. Red sandstone, . dune cross bedding, barchans, interlayered arid, desert.
evaporities.
Chapter 5
STRATIGRAPHIC PALAEONTOLOGY
5.1 STRATIGRAPHI SUBDIVISIONS AND UNITS .

· h' 1· . lly mcuns science of layered rocks (strata : layers). But more precise ly
St \llSnll I ' I1CIU
~ • .
~tmt igrnphy or historical geology concerns with the study of vano~s types of roc~s oft.he earth
surfl e. their ·lussification into ordered units, their arrangement m a chron0Iog1cal order and
fin.ill from uvuilubl • data interpretations of historical events of the earth. These interpretations
incl ude n t only , ... cognition of past geological events but also deduction of the hi story of the
past lif" in •1 chr nological manner. Fossils within rock are the basic clues for such interpretati on
of past rgani history of the earth.
Stratigruphy has three branches viz : litliostratigrapliy, biostratigraphy and chrono-
straJigraphy . All these branches involves investigation of rock strata by classifying them into
meaningful objective units, larger and smaller which are called stratigraphic units (Fig. 5.1 ).
Lilhostratigraphy concerns with the subdivision of rocks on the basis of some observable
lithological characters expressed by succession of rock layers. The common lithostratigraphic
unit may be, arranged in a hierarchical system, such as group, formation and bed. A bed is
the smallest division of rock sequence showing some distinct and unique lithological features
as regards its colour. texture, structure and composition. A group of beds constitute a formation
showing a common genetic history and structural pattern. A group ranks above formation and
is composed of two or more formations formed within a single large territory or basin.
Fos ils are the basis of biostratigraphic subdivision of rock sequence, and the general term
for a.II biostratigraphic units is 'zo11e' which may be of different types but each zone must be
distinctive from others by its fossil content.
The purpose of study of chronostratigraphy is to organise the rock sequence with respect to
their geologicaJ age. Thus this classification needs a concept of geological time. Geological
time involves the time span between the formation of the earth upto the present day. For a
chronological arrangement of geological records and events. geological time need to be
subdivided and standardized as far as possible all over the world. The conventional hierarchial
system of geolagical time units are : eon, era, period, epocli and age from higher to lower
r~n~ .. These ar_e obvi~usly abstract units and hence cannot be called stratigraphic units (objective
d1:1s.mns). A _rime units can be .m ~c objective by referring the portion of rock that was deposited
within that time span. Thus d1v1 s1ons of rocks corresponding to each time unit constitute the
chron.ostratigrapl,ic units or time rock u11its. In other words each time unit mentioned above
has its corresponding chronostratigraphic unit.. The relation between this two is shown in the
Table-6. A generalized geological time scale is also shown in Table-7.
38
• Ji Q[ Q. ii SI . .d@. !L# I.JS.ii, 411 11 . . j p a t $ll! .: t
STRATIGRAPHIC PALAEONTOLOGY 39
TABLE-6
Geological time units and corresponding chronostratigraphic units
Examples from standard
Geological time units Chronostratigraphic units geological column
Eon Enothem Precambrian, Phanerozoic.
Era Erathem Archaean, Proterozoic,
Palaeozoic, Mes0zoic, Cenozoic.
Period System Cambrian, Ordovician, Silurian,
Devonian Carboniferous,
Permian (all within Palaeozoic).
Epoch Series Lower Cambrian, Middle
Cambrian, Upper Cambrian
(all within Cambrian).
Age Stage Olenella Zone, Paradoxide Zone
(both within Lower Cambrian).
'Age', the smallest unit of time is generally defined as a portion of time which was inhabited
by an index fossil. Thus the division of rock within which the fossil occurs in a sequence
constitutes 'stage', the smallest subdivision of chronostratigraphic unit.
5.2 LAW OF FAUNAL SUCCESSION: THE BASIC PRINCIPLE OF STRATIGRAPHIC

PALAEONTOLOGY
In biostratigraphy fossils are successfully used in fossiliferous Phanerozoic rocks in their
biostratigraphic subdivision, correlation and age determination. It is possible to use fossils for
these purpose because it has long been known that there is a definite relationship between the
fossils within a bed and its relative position in the time sequence.
In 1669 Steno proposed the law of superposition which assumes that in a sequence of beds,
younger strata are laid down on the top of older strata. Hence, it can easily be followed that
more and more older fossils should be found towards the bottom of the sequence with younger
fossils towards the top. However, the first conclusive statement in this regard came from William
Smith ( 1769-1839) who proposed another fundamental principle, the law of fa1111al succession.
According to this law, assemblage of fossils in a rock sequence exhibits some change in a
systematic manner from a simple types in older rocks to more and more complex type towards
the younger rocks. So each segment of time represented by a body of rock always shows a
distinctive assemblage of fossils which is different from that of the overlying and underlying
beds. This pattern of fossils in a rock sequence helps a stratigrapher to identify rock bodies
showing identical fossils even located at widely separated areas and to assign them to same
geological time. In an area where rocks are folded and deformed, application of law of
superposition may be difficult due to occasional inversion of beds. Faunal succession can be
safely used here provided the rocks contain fo ssil::;. Faunal succession is considered as
fundamental basis of all types of stratigraphic application of fossils. Smith however failed to
l 1.1l.1l' ({jl'IIJ\\'l'- ti .
40 PALAEONTOLOGY
give any possible explanation about the pattern of temporal change of fossil characters wit~in
rock-sequence. The proper explanation came from Charles Darwin, about I 00 years after Smith
when he proposed his theory of organic evolution.
5.3 PRINCIPLE OF UNIFORMITARIANISM AND FOSSIL

One of the basic principles in geology, proposed by Charles Lyell in 1830 is t~e_p~inci~I~
of uniformitarianism. But this uniformity doctrine actually was older than Lyell for it 1s 1m~ltc1t
in earlier writings of Hutton, Leonardo and Buffon. The basic idea is that the sa_me physi~al,
chemical and biological processes occurred in the past as those of present. From this, geo~ogis~s
arrive at the familiar dictum 'the present is the key to the past'. Although an assu~pt1on, 1~
certainly agr~es with observation and serves usefully in the interpretati?n of _the_p~st h1_story of
the earth. If.fossils are derived from past organisms, from biological umformitanamsm it seems
logical to assume that life processes were the same in the past as they are at present. Th~s
palaenotologists in many cases apply this principle in the interpretation of ecology of a foss11-
group, just comparing their morphology with their living counterparts of the present earth
observing their nature of habits. Similarly, this principle has been successfully applied in the
interpretation of palaeoenvironment and palaeogeography from fossil record.
But uniformity of nature must not be assumed in a rigid sense because only a static earth
could be completely unchanging. Earth is clearly a dynamic body and there is a considerable
fall of intensity of the dynamic processes in the present earth. From this, it can be visualized
that all the earth processes now operating were similarly operated in the past but definitely at
different rate and/or intensity. Rivers flows at different rates carrying different amount and kinds
of sediments; rocks are weathered in different ways as temperature, proportion of moisture in
atmosphere and available chemical elements in water and air are different today. Like physical
and chemical processes biological processes must vary in their rates and interactions. Moreover,
man's knowledge about the earth's processes is very much limited by his short period of
existence. So an event that occurs but once in billion years may remain unexplicable or a process
that acts very slowly over a long period may be unrecognised. Thus uniformitarianism or
actualism (as called by Kelvin) is a working principle essential to geologists but not an invariable
rule or scientific law.
5.4 TIME DISTRIBUTION OF ORGANISMS

When temporal distribution or time range of organisms is closely studied in a geological
succession of strata, it becomes clear that the organisms made progressive changes with the
advance of geological time after their appearance. The empirical observation to this effect was
formulated as a law (law of fauna/ succession) by William Smith and this was an important
recognition of the stratigraphic significance of fossils. As an explanation of the distribution of
fossil organisms as noted in geological succession, a hypothesis known as catastrophist concept
was suggested by the school of Cuvier, a pioneer French palaeontologist in the early nineteenth
century. This hypothesis presupposes that each major unit of geological time was terminated
by a worldwide catastrophe which extinguished all the life forms of the earth at a time.
Organisms in a modified forms were then restablished at the beginning of the next unit of time
by what was called an act of 'creation'.
The catastrophist concept was however gradually undermined and finally overthrown by
Charles Darwin's momentous theory of organic evolution in mid-nineteenth century. The
doctrine of evolution is based on three fundamental concepts known as variation, heredity and
natural selection. The most important implication in the doctrine of evo'lution is that a species
after it is born at a place evolves into more and more modified forms and may be ultimately
replaced by a new one. Anyway, it is destined to an end and the geologic range of species
would depend on the rapidity with which it evolves or is substituted by a new one. Thus a
species is always restricted within a definite interval of geological time, and once extinguished
it never reappears on the earth. It is relevant to point out here that the rate of evolution is deemed
to be very important in determining the time range of an organism and usually the rate of
eyolution is understood in terms of its degree of morphological specialization. However, it is
suggested that a stable environment produces a relatively slow evolutionary tempo, while rapidly
changing environment promotes rapid evolution.
Usually, when a new organism or fauna appears in a region it is taken to represent the
evolution of that form in that place. However, it may also indicate the removal of a barrier
somewhere else or shifting of an environment favourable to that organism existing elsewhere,
and not its evolution. So also the disappearance of a fauna from a region may record either its
extinction or merely its extermination. The term extinction is applicable to the case where
disappearance is due to evolutionary causes, such as the case of ammonities and dinosaurs which
had died out of this living world and are no longer present in any comer of the earth. But
when a fauna disappears from a locality owing to some external causes but survives elsewhere,
it is said to be exterminated from that locality but not extinct. For example, forms like giraffe,
gorilla, chimpanzee were present in Upper Siwalik times in the Himalayan region of India but
then they were exterminated from that region due to some unfavourable conditions (glaciation)
and at present they are surviving in Africa.
It has been also noticed that an isolated geographic area undergoing practically no
environmental change often helps an indigenous fauna to survive even after it has been
exterminated everywhere else on the earth. Such a place is called asylum for the persisting
fauna. The island of Timor, for example, has yielded from the Permian rocks a rich fauna of
crinoids and blastoids which are otherwise known from Early Carboniferous from the other
parts of the world. Some primitive mammals like platipus, kangaroos are found exterminated
from all continents in the Early Tertiary times, except Australia where these fauna are persisting
still now due to geographic isolation of Australia and the latter becomes asylum of this fauna.
If conditions change in such a way that a fauna preserved in some asylum is allowed to
disperse again, it might reappear in a region from where it has been exterminated earlier. Thus
that fauna in that locality would occur in the regional sequence of strata at two levels though
absent in between and such a fauna is called recurrent fauna. Again this fauna by' chance,
might ~nter a ne~ regio~ ~here its arrival and range would be quite different from those areas
where ~t h~d survived oragmally; such fauna is called Jieterochronous fauna, e.g. Schizob/astus
(blasto'.d) 1s t~e marker of Early Ca~boniferous in North America by its first appearance but in
Tim?r island It appears for the first time in Permian strata where it is taken as a heterochronous
fossil.
42
TYPE OF STRATIGRAPHIC EIJUIVAl r,,NJ 1 IME

UNITS NAME OF COMMON UNITS
OIVt ICA'
GROUP
LfllfOSTRATIGRAPHIC FORMATION
MEMBER
BED
RANGEZONE
CONCURRENT RANGE ZONE
BIOSTRATIGRAPHJC ACMEZONE
ASSEMBLAGE ZONE
'
lNTERVAL ZONE
EONOTifEM EON
ERATIIEM ERA
SYSlEM PERlOD
CHRONOSTRATIGRAPmc SERIES EPOCH
STAGE .AG£ I
CHRONOZONE a· .
t
FIG. 5 - I : DIFFERENT TYPES OF STRATIGRAPHIC UNITS
' ' Lycopsid
' '
Bryopbyta ' '
Psilop~id
'i,'
Ch~
~)ta
I
1' ' '
FIG. 5 • 2 : A BROAD PHYLOGENETIC T Aufotropbic bac:taia
GROUPS OF PLANT REE SHOWING EVOLUTJON OF DlFFEREXT
5.5 MAJOR LINES OF EVOLUTION OF PLANT AND ANIMAL GROUP

Plants : Whatever may be the origin of virus and bacteria, the different groups of algae
must had arisen from some autotrophic bacteria. But whether all the different groups of algae
arose independently from a common stock or from different groups of bacteria is difficult to
presume for lack of any direct evidence. Cyanophytes or the blue green algae are considered
the earliest plants in the sea as evident from fossils (algal stromatolites). Structurally, they are
also very simple with cell without a distinct nucleus (prokaryotes) and plastids. They could be
ancestors of all other advanced groups of algae. However, it is more or less accepted that earliest
land plants arose from some groups of green algae or chlorophytes. These two groups not only
show similar mechanism of photosynthesis but also chlorophytes is the only algae which made
for the first time a successful transition from marine to freshwater and even to moist surface,
a logical step towards the life on the land. French botanist Liginer proposed a speculative theory
assuming that ancestral green algae, some branching filmentous forms, were mainly tidal-flat
dwellers. With elevation of coastal land much of these flora became extinct but a few of th e m
were able to survive penetrating soil and developing primitive root system, thus capabl e of
absorbing water and nutrients from the soil. These plants gradually developed an inte rnal
conducting tissue system to carry water from root to leaf and ultimately became the first group
of vascular land plants, the psilopsids. Psilopsids are considered as the ste.m-form of all other
higher groups of vascular plants like lycopsids, sphenopsids and pteropsids. Pterospsids we re
first represented by ferns or filicales and later by gymnosperms and angiosperms (Fig. 5-2).
Animals : Algal group euglenophyta (at present represented by the typical genus Euglena),
a descendant of blue green algae shows some advance characters over its ancestor. They show
not only an organized nucleus separated from the cytoplasm by a membrane, but also chlorophyll
pigments within some ovoid bodies, the chloroplasts. But unlike the green algae it is provided
with cellulose cell wall. Besides this, cells are also provided with one or two flagella and instead
of being passive floaters like most of the other phytoplanktons they are active swimmers. Th ey
also exhibit a gullet at its anterior end, though they art! autotrophic. Near this gullet there is
also a red-pigment often called eye-spot which seems to be sensitive to light. There are also
reports of sexual reproduction in a few species, besides the common mitotic division of cell.
In other words this group, as a whole shows a curious mixture of plant and animal
characteristics. For this reason, zoologists ;,itend to classify some groups of engleno-phytes
within the protozoan flagellates and the later are considered as the earliest group of protozoans
evolved from such green algae. As such a transitional phase is found to present among earlier
plants and animals, many biologists intend to group all these simple animals and plants (most
of which are unicellular) into a single phylum, the protista.
Once the protozoans were established they soon became greatly diversified. Most of them
developed dimorphism and sexual reproduction. Though, there is no direct proof, but origin of
metazoans from protozoan ancestry is more or less accepted by all. There may be two broad
possibilities as regards the origin of first metazoan . Repeated nuclear division within a cell in
a organized manner might have led to a multicellular body. The second method is the
~ifferentiation of individual cell, within a colony of protozoa leading to interdependence and
individuality as found within the porifera, the earliest metazoan group, so much so that it is
often debated that whether porifera is a true metazoan or a colony of unicellular organisms. It
L- ~ - - - - - - -- - -----
~
.fl;.
Brachiopod
/rtho~
Bryozoa
Mollusca
Hcmichordate
Annelid
Bi.laterally symmetrical
- - - - - - - -- - - - - - - - - - - - - - - - -·- - - - - - - - - - - - - -- - - -- - - - - - - - - - -
Trochphorc-like ancestor
. Radi.aly symmetrical
------- Ec h modenn
Hclminthid
Dipleu-likc ancestor C oclenteras
Planula-like ancestor
Porifera--------
Flagellate-protozoan
?
iChlorophyta (green algae)
~
~
~
t""'
0
C)
FIG. 5-3 : BROAD PHYLOGENETIC RELATIONSHIP AMONG THE MAJOR ANIMAL GROUPS "<::
srRATJGRAPHIC PALAEONTOLOGY
45
is possibl~ thaht. ahnchestor of t~e true metazoan could be a blastula like colonial flagellate
prot ozoan m.w 1c t ere was d1fferentiat·ion of somatic
· and reproductive
· cells. It was somewhat
.k
l1 e p/a11ula larvae produced by som h · ·
e worms or elmmthes. Coelenteras are considered
. of such. an. ancestor· Most of th ese ear1·1er metazoans are radially
derl·vatives · · ·
or b1rad1ally
symmetrical. Helmithids were evolved from some multinucleate ciliatas (protozoan) which
become a~oelus w~rms _by t~e formation of cell-membrane. The rest of the metazoans may be
group.ed m two divergmg Imes. One line culminates in the phylum annelida, arthropoda,
brach10poda and mollus<.:a. and the other line ending with the phylum echinodermata and
chordata. Because of the widespread occurrence of trochophore larvae among the helminthids
and annelids, such an ancestor might have given rise to all the former groups of invertebrates.
Echinoderms-chordates probably arose from some planula-type of primitive worms (Fig. 5.3).
5.6 HISTORY OF LIFE THROUGH AGES

Stratigraphic history of the varied types of organisms appeared in different geological times
may be discussed on following headlines.
5.6.1 Chemical traces of life

Biochemical and astronomical studies provide a foundation for hypothesis concerning the
origin of life, but we must look to the preserved fossils for direct evidences of past life.
Radioactive dating of meteorites indicates that the earth was consolidated from cosmic dust at
least 4.6 b.y. ago. The oldest crustal rocks so far discovered have a radioactive age of about
3.6 b.y., i.e. about I b.y. younger than the earth itself. However, traces of life have been recorded
from rocks of 3.4 b.y. age. Earliest organisms are mostly prokaryotic algae and bacteria. They
left some sorts of chemicals traces within the rocks. The most common of these are the
stromatolites exhibiting branched or layered structures within rock, commonly made up of
calcium carbonate or silica. They are widely distributed in older and younger Precambrian rocks
of the world. Most of the palaeontologists believe that these biogenic structures are produced
by some photosynthetic blue-green cyanophytic algae. Living counterparts of such algae in the
recent tropical seas are producing similar algal structures. Structurally primitive (prokaryotic)
cyanophytes are considered as the most oldest group of plants preserved within ancient
sedimentary rocks. Some well-preserved stromatolites may exhibit fragments of original algae
that can prove their origin from such plants. Besides, there are no known means of producing
such structures non-biologically.
A second kind of chemical trace is provided by some complex carbo11-compoun.ds extracted
from some Late Precambrian sediments but there is confusion about their nature of origin. The
first possibility is that these carbon compounds did not originate at the same time as the rocks
but instead, seeped into the sediments long after their deposition or were formed in place later
by some bacterial activities. The second lies in proving whether these compounds are the
products of organism or not, as such compounds might have been produced non-biologically
at early times in the earth history.
Biochemists are now convinced that these extracted carbon compounds, though much simpler
in structure than amino acids, fats and carbohydrates, are nevertheless too complex to have been
produced non-biologically. Further evidence of their biological origin is provided by a distinctive
46 l'AIAl•XJNTU/,0(; y
property nf th curhnn compound prmJuc cJ hy or ,1111is111s. lo ma11y carh ,,. ..compoun<Ji;, rm,I ~<.; ul · i
ar• com11..)s •J of lung spirnl dlllins of curbon li11k ·d to oth ·r clcrncnt . Such chains <1 ·<.: ur i11
two forms that. show 1hr sani, ·omposi1in11 a11d prop ·rli ·s. 111 <inc f<>r,n lhc chHi11 spin,1."l tu rh,~
lrft and uth ·r it spirals to the right. 111 nun· hiolugic11I carhon· l.:Ofllf)OUncJ thcs · twc, kit1cJ of
chains nr • found t occur in equal propnrtiu11, half left-spiral ·d and half ri ,ht-spiraled. If ,wcv ·r.
organisms for r ·asm1s yet to he undorstood constru ·1 dwins thut spirn ls only in I ·ft dir ·cticm.
Such carhon comp unds rolnte th• plnnc of poluriz ·d light wh ·n I okcd ul t.hrou -,h a polarh, irw
micr scope and are ·alled optically active. On the other hand, 11011-hiolo ,;cal carhon c.;omp<,und~
do not rotate the plane of polarized light and arc called <>plically i11active. A high pcrccnta •c,;
of optically active carbon compounds in ancient sediments is, therefore, a stro11g evidence <11
lhcir orj.!anic origin.
5.6.2 Earliest recor,ls offossil:;·

In addition to these chemical traces, there huve been frequent reports of actual fossilizi;cJ
purts. impressions, trails, tracks of organisms from some Precambrian rocks. I l ,wevcr, uclut1 I
affinities of many of these organisms are doubtful. A brief accounts of some ancient fo.._ . _ ;1,1,
an! given below.
(i) Fig Tree Chert Formation

Tht! oldest rock in which fossils have been so far recorded is the Fig Tree Chert
Formation of South Africa dated as 3.4. b.y. Microscopic observation has shown the
presence of tiny microspheroids similar to some modern blue green algal materials. Their
size-range is within the limit of modern prokaryotes. This oldest layer of chert formati on
with organic traces indicates that early chemical evolution of the earth had passed into
organic evolution and also suggest that life and formation of layered-rocks came almo-"1
simultaneously in an oxygen-free atmosphere. '
(ii) Gunflint Chert flora

~xpose? along the shore of Lake Superior in northern Minnesota and adjacent Ontari o
,s a ~cncs of ~recambrian sedimentary sequence overlying older volcanic and granitic
terrain · Gunflmt Chert Formation lies at the middle of this sequence, composed of ab ut
ten feet la~c.r of alternate while, red and black cherts. Gunflint Cheri, dated about
2 b.y., _exh,b11s dome-shaped stromatolite masses . Significance of Gunflint flora was
rec g~ ise.d by Tayler and Barghoorn ( 1954). They dissolved some cherty r cks in
~ydr~f ~onc aci~ and found a residue of tiny filaments and sphcrtcal bodies which were
,d~nr,~i.ed ~~ fra~nacnrs of primiti_ve ulgac. These were invcstiguted under clt:ctron
m,croscopc and tour hroad categoncs huvc been recognised .
(i) Thin threads with w·all-l1'k
• c· p·Lt rt 110n
·,.· rescm
, bl 'mg some modern blue green al ,ae.
(ii) Spherical bodies, liomc resembling present day bacteria and some unicellular blue
green algae.
(iii) Siar-shaped hodics of unknown uflinity.
(iv) Umbrella or parnchu1c-strnp •d bodies of unknown affinity.
47
(iii) Ediacara fauna-the oldest d .
recor of ammal t ,·1 (F.
The first record of undoubted f .. , . ~ss1 s 1g. 5-4)
1
Precambrian rocks of Edica . h~~s, ~ ot soft bodie? animals have been obtained from
1 01
marine sandstone Tile Ed. ra South Australia. The fossils come from a shallow
· rncare fauna ·Is . d' ·
the beginning of Camb · b ra ,ometncally dated as 640 m.y., antedating
?.
reported from Scandina~;n y about 10 m.y. Similar type of faunas have been later
a, southwest Afnca, Russia and India.
The Ediacara fauna, containin . b .
. 11· · t ti . g .t out 1400 specimens belonging to about 30 genera
fa mg m o Ive mam categor"1es o f two maJor . phyla cnidaria and annelida, are as follows:
(a) Rounded
. impressions
. w·tt
' l rad'iatmg
· grooves resembling modern jelly fishes. Such
fossils also .occur
. m you nger Ph anerozo,c
. rocks. Important genera are Cyolomed11sa
and Medusuutes.
(b) Im~res~ions of stalk-like fronds with grooved branches also resembling th<! primitive
Cmdana pennatulaceans. Some genera are Charniodiscus, Ptridiniwn, Rangea etc.
(c) (1) Elongated worm-like impression with a horse-shaped head and a segmented
body resembling some living annelids e.g. Spriggina.
(2) Rounded flattened worm-like impression with a central groove and strong
segmentation also resembling some annelids e.g. Dicki11so11ia.
(d) Some oval-shaped impressions with T-shaped grooves and some circular impressions
with three bent-arms radiating from centre, both of doubtful affinity e.g. Precam-
bridium, Tribrach.idium.
5.6.3 Precambrian-Cambrian boundary

There was a drastic change of organic types with the beginning of Cambrian evident from
the fossils found in Cambrian and Precambrian rocks. In contrast to the record of traces of
simple plants and soft bodied animals within Precambrian rocks, cambrian fossils are represented
by almost all the advanced invertebrate groups, such as trilobites (arthropods), cnidarias,
primitive molluscs, brachiopods, and earliest echinoderms. Moreover, most of them appeared
very abruptly ·which at first sight may be difficult to explain.
There are two explanations : One, organic evolution of this time was very rapid due to intense
selection pressure operating at that time. The second explanation is that the acquisition of hard
shells as opposed to an unmineralized integument, must have brought the bearers thereof to a
'fossilized threshold'. This acquisition of mineralized shell took place approximately the same
time and spread from one taxon to the other. About the origin of this hard skeleton it has been
said that calcareous and phosphatic materials which usually constitute the hard parts, originally
developed as excretory products accumulating over the animal's skin, gradually making it hard.
Once it was formed, quite accidentally, the shell was used as a base for safe muscle attachment
an~ also to protect the animal from predators. This would give these shelled animals a
tremendous selection advantage. Once such a feature was achieved by one group, the selective
pressure to other phyla to develop such hard coverings would be significantly increased.
Consequently, most of the invertebrate groups soon developed similar hard skeleton.
Palac(Gco)WP- 7
PALAEONTOLOGY
48
Medusinites
Cyclomedu.sa Sriggina
Rangea
Tribrachidium
Dikinsonia
FIG. 5 - 4 : SOME ELEMENTS OF EDIACARA FAUNA
I I'\
-,I\:
""'
E
F ASSEMBLAGE ZONE OF D, E & F
o() ACME ZONE OF D
~f'
ASSEMBLAGE ZONE OF D & C

C
~'
CONCURRENT RANGE ZONE OF 8 & C
B
J\
RANGE ZONE OF A
A
FIG. 5 - 5 : DfFFERENT KINDS OF BIOSTRA TIGRAPHIC ZONES
.-
~, .
I•
..;
srRATIGRAPHIC PALAEONTOLOGY 49
some auth~rs a~vocate som~ drastic change of environment as an important factor resulting
. rapid diversification of organisms at the beginning of Cambrian. But it does seem that definite
,n · l . l 1 .
level of phys10 o~ica comp exity_ was attained by many animal groups that enabled them to cope,
first with excretion and, later with secretion of various inorganic minerals used to make their
skeletons. Unfortunately, there is no fossil record from which one can infer about the origin of
these complex groups of invertebrates within the earliest Palaeozoic rocks. All the animal groups
in Cambrian were already clearly separated and distinct. Apparently they might have some
common ancestors from which they arose, that might lacked hard parts or they were so rare that
their fossils are yet to be traced. Comparative study of the animal groups indicates that bryozoa
and brachiopoda are more closely related. Similarly mollusca, annelida and arthorpoda appear to
be related as do the echinoderms and chordates. Protozoa, porifera and cnidaria are more primitive
groups of Precambrian and may be the ancestors of other advanced phyla.
5.6.4 History of Pla11ts

The land plants of the group trachaeophyta probably arose from some green algae by
developing root system; the stem also developed vascular system for gathering water and
nutrients. These fossils are known from Ordo-Silurian beds. The first great expansion of land
plant led to evolution of ferns and related forms which were dominant throughout the Palaeozoic.
At the beginning of Carboniferous these early seedless plants were mostly replaced by early
gymnosperms (pteridosperms). They had dried seeds and pollen for reproduction and throughout
the Mesozoic they were the dominant group of land plants (age of gymnosperms). Gradually,
these open-seeded group of land plants were overshadowed by a third and final group of plants,
the angiosperms. They developed suitable adaptation for more successful reproduction and
dispersion in land by producing seeds which are enclosed or covered by hard integuments.
Fossils of this flowering plants first appeared in Late Jurassic time, but attained climax during
Cenozoic with the decline of gymnosperms (age of angiosperms).
S.6.S History of Invertebrates

First record of invertebrate fossils was found in Ediacara fauna which is mainly represented
by three phyla : protozoa, cnidaria and annelida. But in the earliest Cambrian rocks appeared
arthropods and brachiopods. The earliest most arthropods, the trilobites, which made their first
appearance in Lower Cambrian, became dominant in Middle Palaeozoic and extinct at the end
of Permian. Inarticulate brachiopods appeared in Lower Cambrian along with trilobites followed
by articulate group from Ordovician onwards. Brachiopods attained their climax at Permo-
Carboniferous and declined after Permian. Owing to the dominance of these two groups,
~ m a y be referred to as · ~ a11d b~~Ordovician marks the
first appearance of many other groups of invertebrates such as anllfozoans (tabulates and
tetracorals), echinoderms (sessile pelmatozoans) and molluscas. The Middle Ca mbrian Burgess
Shah! Formation of British Columbia, Canada has yielded an exceptional fauna enriched in
~hropods (trilobites and others) crinoids, numerous annelids etc. Even the soft bodied early
invertebrates are also preserved here.
Mesozoic witnessed some drastic change of faunal characters. Most of the representatives
of Palaeozoic phyla became either extinct or declined in number. The extinct forms are trilobites,
most of the inarticulate brachiopods, all tabulates and tetracorals and all sessile echinoderms
PALAEONTOLOGY
50
. · Of brachiopods with gradual increase of number
(except crinoids). There was .a drastic decline h· d.' A iong the cephalopods, all the
d . I cypods and cep a 1opo s. n
of molluscs, such as gastropo s, pe e ·' .. b d· the other hand, ammonoids,
. . ·1 ,~ ·1 d . ·oss the Permian oun ,try. 0 n
nautilo1ds except Naur, us a1 e to c, · · b b d · nt and 'tttained their climax
. p I oic gradually ecame a un a '
which remained su bd ue d in a aeoz ' . t th end of Cretaceous and hence
. . b I . , tely they became extinct a e
111 Jurassic-Cretaceous ut u uma
ri · . f · ·d , · tie
:1
.
1 af!e of ammomtq;1 1n
M sozoic cnidarians are represented
e. ,
~esozo1c may be re e, re to ,ts ~~ • ~ , d . th end of Cretaceous gradu a l
b ~ r o u ps scleractinians and alcyonanans. owar s . d~ d
dominance of foraminifera, a group of protozoans has been 111 icate ·
Cenozoic m·1rks the appearance and dommance o mo.
· f st of the modern representati ves of
.
0
different inverte~r:lte phyla. These are protozoan foraminife~a, numerous typ~s . f co:o~i ~: cor~~
of scleractinians and alcyonarians, numerous arthropods, mnunerables vanet,es O gas ropo s
and pelecypods and varieties of benthic echinoderms.
5.6.6 History of Vertebrate .

Vertebrate certainly arose from some sort of invertebrate anc~stors b~t as ts so often the
case. the exact ancestral group remains yet unknown to us. Earliest fossils of fis~-bones are
found within Middle Ordovician rocks which were fully differentiated from their .invertebrate
ancestors. Graptolite~ the well-known fossil group from Ordovician-Silurian, accordin.g to so'?1e,
may represent some earliest group of protochordates. It appears that some soft-bodies n:1a~ine
animals possessing primitive backbone (notochord) were related to ancestral vertebrates. Similar
Ji ving marine organisms belonging to protochordate group are Ianceolets. These are somewhat
specialised descendants of the earliest Palaeozoic ancestor of vertebrates.
Palaeontologists have the opinion that echinoderms might have a close relation to earliest
vertebrates, because at larval stage they are similar to some protochordates and are different
from other invertebrates. It may be possible that chordate and echinoderms arose from the same
unknown ancestor in the earliest Palaeozoic times.
Although, first trace of fish fossils are found from Middle Ordovician, but they flou rished
and dominated during Silurian (age of.fishes). Devonian was the time when vertebrates first
made their transition to land, for the oldest fossil amphibians are found in Upper Devonian
rocks of Greenland and eastern Canada. In adapting a life on land, the early vertebrates faced
the problems of reproduction, water retention and respiration in air that were earlier sol ved by
land plants and land invertebrates. In addition, they found an unique problem of locomotion
on land. The lobed fin fishes, crossopterygians, with their complex fin muscles were ideally
suited to develop into elongated flexible limbs to support the movement of anima ls on land
and they are considered by many as ancestors of amphibia. One of the ancie nt-most amphibian
fossils is lcthyostega of Devonian which possessed well-developed limbs together with a tail-
fin like a fish. Trueamphibias appeared in Upper Devonian. They attained the ir climax in Permo-
Carboniferous (age of amphibias). It is suggested that mode rn amphibi as are all envol ved in
Triassic-Jurassic as descendants from the earliest Palaeo zoic forms known as Iabyrinthodonts .
The transition between amphibias and the ir desce ndant re ptiles is less clear from fossil record
because the principal differences between them are not skeletal but in the mode of the ir
reproduction. From fossi I record it appears that by Late Carboniferous pe riod the first g roup
of reptiles, evolved from some labyrinthodont ancestry, were the cotylosaurs that subsequently
gave rise to all other reptiles, hence sometimes called stem-reptiles. Reptiles, althoug h first
I --~ .
SI
appeared in Devonian-Carboniferous became dominant in Mesozoic and the latter sometimes
is called the age of reptiles. There was appearance and diversification of variable forms of
reptiles in Mesozoic which include, thecodonts, dinosaurs, pterosaurs, dicynodonts, cynodonts,
and many others. Many of them attained huge dimensions (giant reptiles) and were adapted to
various modes of life (land dweller, aquatic, flying in air, burrowers etc.). However, dominance
of reptiles came to an end with the end of Mesozoic when all large reptiles suddenly became
extinct and there was an abrupt decline of the number of reptiles in the Cenozoic period.
Mammals possibly arose in Late Triassic from cynodont group of reptiles (mammal-like
reptiles). The skull and teeth pattern have indicated that those reptiles were both herbivorous
and predatory carnivorous like mammals. During Triassic, they progressively grew more
mammalian features and one of them gave rise to the earliest mammal. Again it is seen that
the significant difference between reptiles and mammals is mainly reproductive and physiological
rather than skeletal and that causes difficulty in pointing our their transitional stages from the
fossil-record.
The shelled reptilian egg was an important advance over the amphibias and fishes. But
mammals totally abandon the production of external eggs. Instead, the egg is retained within
the body of the female where embryo develops and is protected before being born active. Along
with this internal embryological development, mammal has developed another specialization
that permits the young animals get nourishment from milk produced by the mother in her
milkgland.
It is possible that earliest mammal evolved from one of the various mammal-like reptiles
prevailed during Upper Triassic, or different groups of mammals may independently evolved
from different stocks of mammal-like reptiles, a phenomenon called polyphyletic origin.
Although, like mammals birds are also warm-blooded yet from their egg-laying reproduction
and general anatomy, birds can be accurately said as feathered reptiles. From fossil evidence,
it appears that bird first appeared in Jurassic from some thecodont reptiles. The Jurassic
Archaeopteryx, a fossil bird exhibited feathery wings (modified anterior limbs), beaks (with
teeth like a reptile) and a tail.
Though birds and mammals both appeared at different times in Mesozoic but they only
became dominant in Cenozoic eriod after the fall of reptiles.tenozoic is thus sometimes called
the age of birds and mammals Diversified groups of birds and mammals appeared at diffrre~lt
times of Cenozoic adapted to variable modes of life. Among the wide varieties of Cenozoic
mammals, there are marsupials (kangaroos), edentates, insectivors, primates, ungulates, rcxh:nts,
bats, whales etc. The major animal and plant groups appeared in different geologici.11 perit-xJs
are shown in Table- 7.
5.7 USE OF FOSSILS IN STRATIGRAPHY

Stratigraphic use of fossils may be discussed on following heads :
5.7.1 Biostratigrapliic zo11atio11 (Fig. 5-5) .

From the nature of distribution of a taxon it is clear that each spedes, evolved from an
ancestor, has a beginning and if is extinct today it has also an ending. From such an extinct
PALAEONTOLOGY
52
TABLE-7
Geological time and major groups of organisms
Age
Vertebrates Invertebrates Plants
Geological time (approx.)
before
Recent
C Recent
E Pleistocene dominant birds, dominant molluscs, dominant
N Pliocene bony fishes and echinoids, corals flowering plants.
0 mammals, few and foraminifera.
z Miocene reptiles and
0 Oligocene
I Eocene amphibias.
C Palaeocene
65M.Y.
M dominant reptiles, dominant dominant
E Cretaceous fishes, few cephalopods gymnosperms
s Jurassic amphibias, first (ammonoids), and ferns; first
0
mammals and birds. pelecypods, angiosperms.
z Triassic
0 gastropods; few
I corals, echinoderms
C and brachiopods.
200M.Y.
p Permian first fish, amphibia dominant dominant ferms
A
L Carboniferous and reptilia; fish brachiopods, arthro- and early
A Dev_onian dominant. pods, corals sessile gymnosperms.
E Silurian echinoderms and.
0
z Ordovician nautiloids; few
0 Cambrian cepalopods, gastro-
I pods and pelecypods.
C
545M.Y.
PRECAMBRIAN Ediacara fauna annelids,
coelenteras
(640 M.Y.)
Fig Tree chert flora (earliest plants)
(3.4 B.Y).
ORIGIN OF EARTH 4.6B.Y.
species occurring in a geological column two boundary lines may be obtained, the lower,
marking its beginning while the upper marking its end. These two boundaries also can demarcate
a portion of geologic time; a time before the evolution of the species, the time during which it
existed and the time since it became extinct; Thus any rock containing that species must have
been deposited within the time when the species existed. This division of rock showing the
total vertical range of a taxon is called taxon range zone. Range zone of a species at a locality .
may represent its total time of duration in the earth or may _represent its local range .which
could be only a part of its actual range because it may be possible that a _taxon may not app~ar
and/or expire everywhere on the earth exactly at the same time. The total time range _of a s~cies
is possible to infer from the extensive data about stratigraphic occurrence of the s~c1es obta.med
from various parts of the world. A range zone is generally named from th.at particular f~ssil. A
range zone, based on such a species which shows only a local occurrence 1s known as te,I zone·
Normally several species occur together in a rock sequence which may help to erect several
range zones. Portion of rock may be found marking overlapping occurrence of two/r~10~e range
zones and such a portion rock constitutes a concurrent range zone. Overall association of a
group of fossils in a portion of rock sequence may constitute an assemblage zone. Acme zone
or peak zone is a body of strata in which maximum abundance of a particular species is found
but it may not be its total range. Interval zone is the intervening portion between two
biostratigraphic zones where fossils are either indistinctive or altogether absent.
5.7 .2 Correlation and age determination of rocks by means of fossils

Correlation is to establish time equivalence of two or more spatially separated stratigraphic
units. Time correlation, though only approximate, can be undertaken, by various methods related
to physical properties of rocks. Older Precambrian rocks directly may be dated by adopting
various radioactive decay methods. But for younger fossiliferous Phanerozoic rocks fossils are
the main tools of correlation and age determination. Most of the radioactive methods except
radio-carbon are difficult to apply in younger rocks because radioactive minerals are not only
rare here but also for the fact that common radioactive isotopes mostly have long half-life. On
the other hand, as radio-carbon has a very small half-life, its successful application is only
possible for fossils of Quaternary rocks.
The most common method of correlation by a fossil uses lower and upper boundary of a
tax.on range zone marking the first and last appearance respectively of a taxon. First appearance
and extinction of a taxon is generally taken as a worldwide synchronous event and they may
indicate approximate same time-level at different rock-sections where the taxon occurs and all
such zones may be considered time-equivalents. Caution should be taken because a tax"· .nay
also appear at different time-levels owing to its migration at different localities at di~.~·rcnt times.
Similarly, a taxon may disappear at different localities at different times. It may ut' exterminated
from on~ locality ~ue to some unfavourabl~ conditi_ons but may continue i11 some other place
under suitable environment. So, before regional or intercontinental correlation by this method
one has to determ_ine the a~t~al stratigraphic range of a particular species by analyzing its
~curren~es at various localities of the world. Taxa found to be very useful in regional and
mte~c?ntmental correlation are the i11dex fossils or guide fossils. An index fossil is one which
exh1b1ts very short vertical range, wide geographic distribution, greater ecologic tolerance,
~b~nda~ce, rap.id rate of evolution and some diagnostic morphologic features. floating or
~wimmmg. marine organisms, also called pelagics, have always a greater chance to be ideal
~ndex fossils as they have capability of wide dispersal within a short time-interval. ivlany
invertebrate
an . benth·1c forms prod uce mob"I Ie I arvae t h rough which
· they can attain wide
· d1spersa
· I
. db_mdex _fossils among them are not uncommon. A similar term, zone fossils. sometimes used
in iostrallgra h . ·I d I . h. h . . . .J
P Y me u e t 10se w 1c exh1h1t short time range but may or may not show w1c e
........_,
.......~_,,{, ... ' -:· . ~ . . :_. ·' ,., , , . ,.._. ·.• . : ,. ·~,·~1
54 PALAEONTOLOGY
geographic dispersal. ln this sense, all index fossils are zone fossils b~t _the reverse. is not alw~ys
. t rat·1graph"1c zone b as ed on a locally occurrinlTe zone fossil 1s ca11ed
true. A b10s .
teil zone
. .
which..
. f l
may b e use d f or corre Iat10n o oca 1 secti·ons
.· Names of some common mdex fossils of Ind1.1
are given below.
Age
~ Name of fossil Animals group
Arthropod (trilobites) Lower Cambrian
Redlichia noetlingi.
Cnidaria (coral) Lower Ordovician
Halysites wallichi
Pentamerus oblongus Brachiopoda Upper Silurian
Calceola sandalina Cnidaria ( coral) Middle Devonian
Waagenophyllum indicum Cnidaria (coral) Permian

Otoceras woodwardi Mollusca (cephalopod) Lower Triassic
Macrocephalites macrocephalus Mollusca (cephalopod) Middle Jurassic
Stygmatopygus elatus Echinoderma (echinoid) Upper Cretaceous
Miscellanea miscella Protozoa (foraminifera) Upper Palaeocene
Assilina exponens Protozoa (foraminifera) Middle Eocene
Pellatispi ra madaraszi Protozoa (foraminifera) U ppcr Eocene
I
/ Lepidocyclina dilatata Protozoa (foraminifera) Middle Oligocene
Miogypsi11a globulina Protozoa (foraminifern) Lower Miocene
5.7 .3 Interpretation of past environment from fossils

Environment usually has some impact, feeble or strong, upon the organisms, which sometimes
may be reflected by some peculiarity of their morphologic characters. Effect of environment
becomes pronounced on organi!)ans living in some localized region such as marine benthic
animals, terrestrial plants and animals whereas control of environment on planktic or nektic
animals are less pronounced. However, any interpretation of past environment must be done
from in situ fossil assemblges. Environmental interpretation is based on principles of
uniformitarianism. That is we have to observe the present environments of the earth and their
various effects on animals and plants now living under such conditions. Then we try to match
them with their fossil counterparts. For example, reef corals are found at present in the tropical
sea at a shallow depth (about 35m) and in warm ~water (around 23°C). These may help to
understand environment of formation of Cenozoic and Mesozoic reef builders. Colonial corals
of Palaeozoic times such as tabulatas and tatracorals have no living counterpart. Yet it can be
maintained from the present observation that formation of biohermal masses has necessiated
sea water rich in dissolved CaC0 3 (above carbonate compensation depth or CCD) and therefore
relatively hot water, and such condition prevails only in shallow zones of tropical seas. This
promotes growth of reef corals in tropical sea of the present day.
Observation of some marine bcnthic animals now living in the sea shows that many organisms
exhibit restricted depth of occurrence as some are sensitive to light and some have restricted
salinity tolerance. Echinoids of present sea prefer mud-free clear and agitated water for proper
functioning of their vital water vascular system.
Terrestrial animals especially plants are _highly affected by the local en ir nment und exhibit
many features in response to temperature, humidity, seasonal variation of rainfall "t . For
example, annual rings found in the wooden stem of many plants indicate an alternate dr an 1
wet season within a year. These are produced by alternate dark and light coloured bands foun J
in cross section of the wooden stem. Dark bands are the result of excessi e gr \ th f \lllSl'lllar
bundles during rainy season when excessive water is available from soil. Poor growth of xyl "m
tissue resnlts in a light coloured ring during the next dry season with . carcity of water. Thus a
dark and a light ring together mark one year and it is sometime possible to al ulate th age
of the plant by counting the number of rings present in the stem (for u es of plants in cl imal i ·
interpretation see chapter 4 ).
S.7.4 Palaeogeographic reconstruction by fossils

Fossils sometime help to reconstruct broadly the past geographic elements of the earth. In
this respect, terrestrial plants and animals become especially important because of their restrict ·d
dispersal capacity. A sea in between two continents act · a, a barrier which prevents migration
of terrestrial animals and plants of one continent to the other. However, observation has shown
that during some geological times the terrestrial fossils of everal continenL which arc now
widely separated from one another by seaways exhibit a striking similarity. This can be explained
only by assuming that those continents must be closely a'. ociared together or connecred with
one another by land bridges during that particular time. Thus rhe two major floral and fauna!
group found during Permian-Cretaceous times of the world may lead to the as. umption of
existence of two supercontinents at that time. The northern one comprising Europe, North
America and Asia (excluding India) is named us Angarala11d and a southern one comprising
India, South America, Africa, Antarctica, Au tralia and New Zealand is named as
Gondwanaland. Fossils of particular age may indicate occurrence and position of land, sea,
rivers in a particular continent at that time. That the pre ..ent Himalayan m untain was initially
a sea is known from the profuse fossils of unbounded marine animal s obtained fr rn the
Himalayan Palaeo-Mesozoic rocks.
S.1.S Fossils as evidences of organic evolution

Fossils give some direct evidences in support of the theory of organic evolution. This theory
states that most of the advanced organisms of today are the products of gradual transformati on
of some simple, primitive ancestral forms of the geological past brought about by such factors
as variation, heredity, struggle for existence and natural selection. Careful study of fo ·siL from
Cambrian rock upto the recent past can trace the history of evolution of one group of animal ·
from the other. Fossils have proved the evolution of amphibias from a fish ancestry and evolution
of mammals and birds from reptilian ancestry. Sometimes this transformational history is clearly
documented by a fossil bearing somewhat intermediate characters between the ancestor and
descendant. /cthyostega was an early amphibia having not only limbs like other terre. trial
vertebrates but also possessed a fish-like tail-fin. The earliest bird Archaeopteryx al o exhibit ·
some characters co111 mon to both reptile and bird, such as a beak like a bird with teeth like a
reptile. Besides, the history of evolution of different groups of animals from their earlie t forms
to the most recent forms can also be traced by fossils left by those organisms. One of the best
known records of evolutionary history has been exhibited by the horse by a series of fossil ·
l'.1l.1~1(il'IIJ\\'f• X
56 PALAEONTOLOGY
preserved within Cenozoic rocks of North America. This record begins with a small, primitive
four-toed horse Hyracotherium of early Eocene times and cultuninates in the modem horse
Equus appeared in Upper Pliocene which are large-sized animals with one toed legs provided
with hoofs.
5.7.6 Fossils as indicators of economic deposit

Fossils not only are of value in conducting purely scientific studies but have also use in
applied geology. Since many of our important resources are associated with sedimentary rocks,
fossils within them may be sometimes helpful to locate such rocks on the earth surface. For
example, mining geologists use the fossils to date the strata below and above the rock containing
the resource. Coal deposits, commonly associated with terrestrial plant fossils, may be located
in the same way. Economic use of fossils becomes increasingly widespread with the exploration
of petroleum. A knowledge of fossils is the basis in the search for the oil-bearing rocks. In
this respect, microfossils are commonly used by petroleum geologists because these small sized
fossils are not likely to be broken by the drill-bit and their study is possible from drill core
samples. Foraminiferas, a group of protozoan microfossils are particularly used in India and
many other countries where oil is mainly located-within marine Tertiary rocks where these fossils
are ~undant. Microfossils like diatom algae, fish teeth and scales and spore-pollen are also
used m other cases.
6.1 STUDY OF PALAEOECOLOGY AND ITS LIMITATIONS
Ecology is defined as the study of mutual relations between organisms and their environments.
Literally it means study of organisms where they live. The place where an organism lives is called
its habitat.
Palaeoecology is the study of the past organisms and their environments. The study of
palaeoecology is much more complex than that of ecology because in case of ecologic study
of modem organisms we can directly observe the animals and their habitats and one can easily
understand their mutual relationship. But in case of dead and past organisms which had left
the evidences of their existence in the form of fossils in rocks, this relationship becomes highly
distorted for several reasons such as :
(i) Many organisms living in an environment may not be preserved for various unfavourable
conditions.
(ii) Organisms might have been removed after their death from their original environment
to some other places where they were fossilized.
(iii) Different organisms living in different contrasting environments might have been
fossilized in one place by chance.
(iv) Most of the soft parts of an organism which in many cases reflect their habitats and
mode of life have been lost during preservation processes. So it is extremely difficult
for a palaeontologist to interprete about the life habit of a fossil organism only from its
hard-part morphology.
6.2 PROCEDURE FOR PALAEOECOLOGIC INTERPRETATION

In spite of these several limitations, palaeontologists have tried their best to interprete the
ecology of the fossil organisms. For this, they have to adopt various indirect methods. These
interpretations are based on a fundamental principle in geology known as 'principle of
actualism'. Thus in the first step towards the palaeoecologic interpretation, palaeontologists try
to correlate the fossil-organisms with their living counterparts. Fortunately, many fossil organisms
have their living representatives in the recent earth. In case the organism in question is totany
extinct, there may be some living forms showing a very close resemblance with the extinct
fossil-form .
Once this relationship is established, the second step is to study the ecology and habitats of
these living organisms. It is essential to note their hard-part and soft-part morphology and their
respective functions in relation to their life habit in a particular environment.
In the third step, palaeoecologists will look for the fossils of that organisms preserved in
their original environment. This type of assemblage is called biocoe11 osis . Within such an
57
;.,'
VI
00
29.2 70.8
i'
~
,..;;.
:i NON-MARINE
ENVIRONMENT MARINE ENVIRONMENT
~
;>:
4000
Ul '
..J
Ul
~· .
2000
o-.----- l\h3h. C . . . .~ ....,. ,.. . . , . ,..,,,, OCEANOPELAGIC -- I l
2000 ~
i'
..._..
t 4000
BATIIYPELAGIC
I
~
Ul
0
6000 ..._. :5 I
~ ABYSSOPELAGJC
8000
10000
0 10 20 30 40 so 60 70 80 90 100
(cumulative pcrccntcgc of earth surface)
FIG. 6 - I : MAJOR TYPES OF MARINE ENVIRONMENT AND RELATED MARINE HABITATS

REPRESENTED BY A HYPSOGRAPHIC CURVE
.. ~ ;
PALAEOECOLOGY 59
assemblage, palaeontologists wiJJ note the various morphologic features of that fossil-organism
and the other associated fossil forms and try to understand the type of environment in which
they live. Then ecology of the fossil-organism in question can be interpreted after a careful
comparison with the data obtained from the ecologic observation on the related modem forms.
6.3 ENVIRONMENTS ON THE PRESENT EARTH SURFACE

The deposition and formation of rock; on the earth surface (sedimentary rocks) is mainly
controlled by various geographic features. The sum total of these \'arious factors is the
geographic environme11t. Each environment on the earth surface is characterized by various
physical, chemical and biological features by which they can be defined and identified. The
nature of rock-types and the organisms are directly related to the type of environment. So rocks
and fossils in them are good indicators of past environments. The task of reconstruction of past
environments is best approached by a careful study of rock deposits and the characteristic
organisms found in the present environment of the earth by application of principle of actualism
(uniformitarianism).
Various types of environment found in the recent earth are as follows :
6.3.1 Marine enviromnent (Fig : 6-1)

Neritic Environment on the continental shelf portion of the sea from 3m. to
200m. depth zone of water.
Epineritic ·ShaJJower part of neritic zone upto 30m depth.
Infraneritic Deeper part of neritic zone beyond 30m ::4epth.
Photic zone Zone of sea water lighted by sun (generally upto 50m).
Bathyal Environment of sea on the continental slope zone (from 200m to
2000m depth).
Abyssal Environment of deep sea zone beyond 2000m depth.
6.3.2 Non-marine enviro11me11t

Fluvial Environment of the running water or river.
Lacustrine Environment of lakes (usually freshwater).
Swamp Environment of an enclosed small lake filled up by organic muck.
Desertic Environment of closed and open deserts.
Glacial Environment of permanently ice-covered region.
6.3.3 Mixed e11,viro11me11t : A zone encroached by both land and sea

Shore environment Environment of zone of sea between high and low tide.
(Coastal or Littoral)
Foreshore Extends from lowest low water to a highest water level where occurs
a well marked break in slope caJled berm.
Backshore Extending from berm hack to the furthest point reached by waves on
the beach, where there is usually a wave cut sand-cliff called 11ip.
~- ...... & • •.• • • . -~
PALAEONTOLOGY
60
Environment of a wide shore affected by tide and fed by large
Tidal flat
quantities of sand and silt brought by rivers into sea.
Environment of an enclosed saline lake near the shore, cut off from
Shore lagoon
the main sea by a sand bar; water of the lagoon is fed by the ocean at
the time of high tide.
Environment of a wide funnel-shaped river-mouth in which tide causes
Estuary
daily reversal of the river current.
Environment of mouth of a river at the junction with sea, where large
Delta
amount of sediments are deposited by river.
Brackish water Environment of a water body formed by a mixture of saline and fresh
water.
6.3.4 Special types of environments

Paludal A swampy environment where bacteria are active; ideal for
transformation of vegetal matters to coal.
Spelean Environment of open underground space where deposition takes place
from underground water.
6.4 TERMINOLOGIES RELATED TO ECOLOGY

Some common terms related to ecology are as follows :
Biosphere The space occupied by the living organisms.
Habitat The place/environment where an organism lives.
Fauna A natural assemblage of animals living together.
Flora A natural assemblage of plants living together.
Biota A natural assemblage of flora· and fauna living together.
Biocoenosis An assemblage of biota in their original place.
Thanatocoenosis An assemblage of biota brought together after their death merely by
chance.
Biotope The area inhabited by a biocoenose.
Biome Different
. biocoenoses o f a 1arge territory
. .
(biosphere . thus
1s · the largest
baome).
Ecosystem Referred to the whole biome and its environment.
Ecologic niche A place
. . or environment where a particular
. .
orgamsm .
finds the optimum
d
con ihons to be flourished.
Population
Individuals of a species in a locality.
Community
Populations of two/more species.
l Diversity
Refers to richness of community.
Biomass
The amount of 1·avmg
· matter .mcludmg
. stored food .m an ecosystem.
/ Dominant species The most conspic uous, species
,· . m . a community. (usually > 50%). ·
l
.... _; .' . ~ ~ -: .. '
PALAEOECOLOGY 61
companion species Other species forming 10% to 20% of the community.
fortuitous species Species showing < 10% of the community.
Biofacies Biologic aspects of a rock, indicating variability of limiting factors
controlling distribution of organisms in space.
faunal province Larger geographic areas characterised by one distinctive types of
biofacies.
Stenotopic Species adapted to a narrow range of environment.
Eurytopic Species adapted to a wide range of environment.
Bionomic subdivision of organisms :

(i) Life Habits (Marine)
Benthonic Organisms living on the sea floor.
Vagile (Vragnant) Benthos capable of movement on sea floor.
Sessile Benthos which are fixed on the bottom of sea.
Epifaunal (Epibiont) Benthos living directly on sea bed on hard or soft substrate; may be
sessile or vagile.
lnfaunal Benthos living within the substrate (burrowers/borers).
(Endobiont)
Pelagic Organisms living in sea-water.
Nektons Pelagics capable to move by swimming (swimmers).
Planktons Pelagics not capable to direct their own movements (floaters).
Phytoplanktons Floating microplants.
Zooplanktons Floating microanimals.
Epiplanktons Move by attaching themselves with some other floating or moving
(Pseudoplanktons) objects.
Nektobenthos Can swim as well as crawl on the sea-bottom.
Meroplanktons Mobile larvae of some marine invertebrates.
Oceanopelagic Pelagics living very close to sea-water surface. (planktons dominating)
Neritopelagic Pelagics living in water of neritic zone.
Bathypelagic Pelagics living in water of bathyal zone.
Abyssopelagic Pelagics living in water of deep sea.
Epibiont Animals attached themselves to other organisms.
Epizoans Animals attached themselves to some plants.
Epiphytes Plants attached themselves to some other organisms.
(ii) Food Habits (Marine)

Carnivorous Mainly feeding on flesh of other living or dead organisms.
Predators Directly living on flesh of other animals catching them in living
conditions.
PALAEONTOLOGY
62
Living on fle sh o f on
ly dend and decomposed animals. . .
Scavangers
. from
Fed by sel ctmg . t he su rrounding water-suspended microorganisms
Suspension fe ders
or detritus. · d h
d
Fed on the detritus and ea d microorganisms depos1te on t e sea
Depo ·it feeders
bottom.
(iii) L,fe Habits (Te rrestrial)
Fresh water Ll' ving in river and fresh water lake. .

d f h
Estuarine/ . . . ·howing
L1vmg m a zone, s a mixture of saline an res water.
Brackish ,, ater
Desert/ Living on or within sand of a desert or sea beach.
Sand D,, ell er
Forest dwe llers Animals living in dense/moderately dense forest.
Arboreals Forest dwellers but mostly living on tree branches.
Arctic!fundra Animals living in arctic or polar regions.
Alpine Organisms on the top of high mountains in both of temperate and
tropical region.
Grassland/Steppes/ Animals living in plain lands covered with heavy growth of grasses.
Savanas
(iv) Food Habit.s (Terrestrial)

Cami vorous-Predators Hunting animals; living on flesh of their preys.
Carn ivorous-Scavangers Animals living on the flesh of dead/decomposed animals.
Herbivorous Animals living on the plant-materials.
Browsers
Animals living on leaves/or fruits of large plants/or shrubs.
Grazers Eaters of grass, small shrubs, and herbs.
Omnivorous
Living on both animals' flesh as well as plant materials.
Saprophytes
Plants living on dead/decomposed orga.nic mass.
(v) Biologic Associations
Mutualism
Or~anisms ~f two different groups remain intimately associated and
derive benefit from each other.
Commensalism
Organisms of two different groups are intimately associated in such a
way that one derives benefit from the other while the other is neither
helped or not harmed. ·
Parasitism
Organisms 0 f t d' &.c
. wo 111erent groups ure associated in such a way that
one derives full benefit at the expense of other.
·· ,· . ~ ~-
' ' " .
PALAEOECOLOGY
63
6.5 FACTORS CONTROLLING THE OCCURRENCE AND ABUNDANCE OF AN
ORGANISM IN ITS. ENVIRONMENT
Organisms ' both animals and plants , 1·1vmg
· ·m an ecosystem are not ·mdependent of each
other. In fact, th~y are surr.ounded by a host of abiotic and biotic factors which are measured
in te~ms_ of ~hystcal, chemical and biological properties of the environment that actually limit
the d1stnbut1on a~d ~~undance of a particular species in a community. These factors are hence
togeth~r. called li~itmg factors. Physico-chemical factors include temperature, pressure,
availab1hty of sunhght, oxygen, water depth (in case of aquatic organisms), salinity (in case of
marine organisms) and nature of substrate. Biological (biotic) factors mainly concern with the
food-relation among the organisms in an ecosystem.
6.5.1 Abiotic factors

(i) Temperature : It is one of the most significant limiting factors in nature. Life can exist
within a specified range of temperature. However, temperature tolerance of different
species is different. It also varies with the different stages of development of the same
species. Temperature tolerance can be distinguished in three levels.
Maximum Uppermost limit of temperature that a species can withstand.
Minimum : Minimum level of temperature tolerable to a species.
Optimum : Temperature at which a species shows its maximum activity.
(ii) Pressure : Air and/or water pressure on the earth surface is variable. Air-pressure on
mountain tops is minimum and it is maximum near the sea-level. At the deepest part
of the ocean the water pressure may be about one thousand atmosphere.
(iii) Light : Light is essential for most animal habitats specially in case of plants where
their vital food-preparing process photosynthesis is directly depending upon the
availability of sun-light.
(iv) Water depth : The effect of water depth upon the distribution of aquatic organisms
especially marine animals/plants is considerable and vital. In fact, it is the main factor
controlling the distribution of marine organisms because this factor directly influences
all other factors like hydrostatic pressure, temperature of sea-water, light intensity, salinity
of water, and oxygen availability. All the factors except sunrays vary with the increase
of water depth. Because no sunlight is available beyond the photic zone of sea, all
photosynthetic plants are restricted only upto this zoney.
Non-availability of sunlight and increase of water pressure with water-depth affect the
distribution of many types organisms. A striking example is the distribution of reef
building corals in the present tropical seas.
Corals are a group of marine and sessile i~vertebrates showing two ecologic types; one
group living individually are found at variable depth of s~a. They are cal~ed 11011-reef
or ahermatypic corals . The other group forms dense colo~1es and are restricted only to
the photic zone of sea. They are called reef or l,ermatyp,c corals. They form most of
the tropical reefs of today. Why are reef corals restricted only to the shallower part of
the sea? Close observation has revealed that the skeletons of these corals are encrusted
Palae(Geo )WP-9
64 PALAEONTOLOGY
by the growth of a green algae known as zooxanthelle, living with the corals. by
mutualism They take shelter on the coral's skeleton and supply CaC03 to corals required
for makin~ coral's skeleton. As these algae cannot grow without sunlight, so they, at
the same time restrict the growth of reef corals wit_hin the photic zone of se~. This
information may be successfully utilized for the fossil reef-corals of the geologic past
which probably exhibited a similar life-habit.
(v) Oxygen : In general, oxygen is constantly renewed. to the deep sea by worldwide
circulation pattern. Cold surface water rich in oxygen smks downwards at the poles ~nd
then passes equatorward along the sea-floor. Thus deeper part of sea-water contains
higher oxygen. In contrast, unless very shallow, the ocean water mass, stirred by ~aves
up to a considerable depth, becomes stratified and thus the deep layers develop deficiency
of oxygen. This condition excludes many kinds of benthos and seems to account for
the decrease of diversity of many organic species away from the shore.
(vi) Salinity : The effect of salinity on distribution of sea-organisms is also considerable.
The different forms are :
(i) Stenohaline : animals with a narrow tolerance range.
(ii) Euryhaline : animals with a board tolerance range.
Most of the marine forms are stenohaline often living near offshore. Many freshwater
forms can Jive in brackish or hypersaline water. However, extrapolating backward in
time from a single Jiving species to a related fossil species with respect to its salinity
tolerance may not give always correct information. The genus Lingula (brachiopod)
appears to have been euryhaline for million years past but today it lives in near shore
habitat, often in brackish wafer.
(vii) Substratum : One of the most important relationship between substratum and
distribution of benthics concerns their feeding mechanisms. Deposit feeders are more
abundant than suspension feeders in a muddy bottom whereas the reverse is true for
sandy or rocky bottom. Substratum is al&o important concerning the methods of
attachment and movement of benthic animals. Rocky bottom is preferred by encrusting
forms, or forms that are fixed by some of their organs for better and safe attachment.
On the other hand muddy and sandy bottoms are preferred by burrowers. ~ _ ..-
6.5.2 Biotic factors-Food relationship among organisms

The activity of all living organisms is centred round their food habits. Within an ecosystem
many constituents, both living and non-living interact. In most communities, organic and
jnorganic compounds are synthesized from environment by the green plants through the process
photosynthesis. These are called producers. These synthesized organic matters (food) are
consumed by animals called consumers. Plants are hence called autotropl,s (preparing their
own food) and aJJ animals are heterotrophs (depending on plant for their food). Among the
animals there are herbivores (primary consumers), carnivors (secondary consumers) and top
or large carnivors/omnivors (tertiary consumers). Parasites, saprophytes, scavangers may enter
into the system by variety of ways.
PALAEOECOLOGY
65
complex orga~ic m~terials deri~ed from producers and consumers are again broken down
and transfor~ed mto simple organic compounds after their death by some microorganisms,
,nostly bactena, called redu.cers, decomf!osers or transformers. The simple organic compounds
produced by them are agam rel~ased m. the e~vironment from which producers once again
incorporate them as raw matenals. This entire cycle is called food chain or food web
(Fig. 6-2a).
Some common features of all food chain are (i) plants lie at the centre (ii) food chain cannot
be a close circuit but -~-ears vario~s ramifications because of existence of various types food
habits of consumers. (111) hypothetically this food relation in an ecosystem can be shown by a
pyramid (food pyramid) where producers form the base of the pyramid, followed successively
by primary, secondary and tertiary consumers. So the number primary. secondary and tertiary
consumers is gradually decreasing in a balanced ecosystem (Fig. 6-2b).
6.6 METHODS OF ECOLOGIC STUDY AND INTERPRETATION

Methods of study of palaeoecology may be divided into two groups : autecological and
synecological. The former method devotes to the study of individual forms and the latter
considers study of communities with all the interaction among the two or more individuals.
A details of different methods of autecological study is discussed here.
6.6.1 Autecological study

This involves detection of life habits morphofunctional analysis, different life activities of
animals and plants and analysis of the environment in which they lived. Some common methods
are as follows :
(i) Direct evidence through preservation
The most direct way to learn the life-habit of extinct fossil animals is to observe them
preserved in the midst of some life activities. For example, some animal fossils are found
attached to some types of substrates or to some other organic shells. Fossils of two
mutualJy associated animals are not uncommon. The chief problem in using epibionts
(an organism attached to some other organism) as indicators for the life habit of their
host species is the one of establishing whether that attachment took place before or after
the death of the host. In the last case, however, no interpretation of life habit of the
host is possible. In the other case the relationship may be established and in such cases
location and orientation of epibionts on the host may be useful points. Many epibionts
live above sediment-water interface to get their food and other requirements. In that
case, if the host species remains exposed partly. t~ water ~nd buried partly to mud
(partially infauna)). the limit of the growth of ep1b10.nt species .on the host-s~ell ~ay
indicate the original sediment-water boundary and this may enltght on the onentat1on
of the host species at its life time.
In situ preservation shows that fossils of some platyceratid gastropods are attached. to
the calyces of some crinoids; apertures of the ga~tropods are fitted on the an.al openmg
of the crinoids. This may indicate that the snails fed on the faecal materials of the
crinoids.
66 PALAEONTOLOGY
I SUN
I
~
I PRODUCER J
TERTIAR y CONSUMERS
SECONDARY CONSUMERS
5850000
PRODUCERS
(b) PY11AMm SIIOWING APPROX/1.fATE NUMBER OF ORGANISMS TIIAT COULD FORM A

/ BALANcEo ECOSYSTEM IN AN ACRE OF LANO (not to scaJc)
FIG. 6 - 2 : FOOD PYRAMID & FOOD CHAIN
... ,.·. ... .. ·.~.....

•• •• •• • • • •
• ••
• • • • ••••
•• • • • •• • • •
• ....
·:-. ... ,. . . .
• ••••••••••••• • • • • • •
• .. -· ...
• • • • • •• •• • • •••• • • • •
• • • •• • • • • • • • •••••••••••
• • •
• • •
• • • • • • ••••
• • • • • • • ••••
• ••
• ..·:.:• ...
• •• •
• • •
• • •
• • •
• • •
.
- • •• • • •• •• ••• • •
(a)RANooM
.··*,....... .........
•
•':.•.\
• • • • • •••••
....
• • • •••••
#• :
.......
••••
•••
• • •
• • •
• • •
• • •
• •
• • ••
(b) AGGREGA TEO
• • • • • •
(c) UNIFORM
FIG. 6 - 3 : NATURE OF SPATIAL DISTRJBUTION OF A SPECIES - POPULA TfON
PALAEOECOLOGY
67
(ii) Homology and analogy-Homeomorph·
d. ism
A l~ss •;ect a~proach to understand the life habit is to infer them by homology and
an~ ogY· n sue( c ases, we have to consider at first, the case in which fossils and recent
amma 1groups re 1ated or unrelat d) th be · ·
. . . e at ar s1m1lar structures which are closely related
but not necessarily
h . mdent1cal and are J·udged t o h ave ha d a common ongm. · · Th ese
structures . avmg co~mon mode of origin but different functions are called homologous .
Such as wmgs of a bird, forelimbs of a man, pectoral fins of a fish etc. In other instances,
som~ structures ~ay develop in two unrelated groups of animals which also serve similar
or different funct.10n but ~ave distinctly different mode of origin. These structures are
calle~ l,omoplastic. Once 1t is established that homoplastic structures also perform same
funct1?n they are called analogous. For examples, wings of birds and wings of insects.
Sometimes same external morphology is attained by related or unrelated group of animals
showing same life habit. The phenomenon is called homeomorphism, or morphological
convergence. For example, richthofenid brachiopods and rudistid pelecypods (both
extinct) are morphologically similar to a solitary conical corals. This indicates that the
former two extinct forms were also sessile like coral and are attached by their larger
valves. Homeomorphism may be found within animals of same geological interval
(isochronous) or or different geologic intervals (heterochronous ).
(iii) Functional morphology
Inference on functions of certain morphologic features of some fossil groups is possible
especially when it is coupled with the inference of their habits derived from sedimento-
logical evidences. However, there is no single method for such inference. For inter-
pretation of function of a fossil- structure a common procedure is as follows : first,
postulation of some hypothetical functions for the similar structures in recent groups of
animals. Next, testing each hypothetical function in terms of whether the function in
question would be useful to the living organisms in the light of their known life habits
and habitats and whether the functions are mechanically feasible in terms of their
morphology and life habits. The most reasonable function is thus chosen on this basis.
The conclusion is then be applied to the fossil structure in question by homology or
homoplasy. In any approach to functional morphology it is important to remember that
a particular structure may perform more that one function. For example, mollusc-shells
serve a protective function in addition as a support to muscular system. Extremely thick
shells make molluscs invulnerable to many types of predations but at the same time in
causes Jess mobility of the animals and many such pelecypod shells are found in sessile
condition. On the other hand, high mobility, most common among molluscan species
having thin shells permits the organisms to escape from predators.
Sharp, narrow and highly elongated pelecypod shells are characteristics of burrowers.
Long siphon in a pelecypod reflected by its internal deep posterior pallial sinus is also
indicative of deep burrowing habitat. Siphons in these case~ a~e project~d ab~ve the
sediment-water level to collect water and food. Elongated sltt-ltke pores m ponferous
zones of some clypeasteroid and spatangoid e~hinoid~ indicate presence of respiratory
tube-feet which in turns indicate their burrowing habit.
. ! _'
'I I • • • • Ir' , . • . . ... ,;- . ./
-·
PAUEONTOLOGY
68
(Iv) Blologlc ocUvity-Pnllchnologlcul data . organi ms have left fo ·sil-evidence
d by skeletal parts. many · -
Even when not r_.prct. enl • ·. . . ft ace fiossils like track , trails, burrow ·,
. d of It fe m forms o r . .. .
of their •,ostencc an wuy _ . t'on of some life acnv1ue of orgam m
·1 oduccd by preserva a .
bores etc. A trace f oss1 s, pr . . beha· viour such as its boring, burrowing
. b t the organism s - .
therefore can reveal muc h a o~ . d' , . behaviour and habits of animals a the
or truiling nmure. These are directly in icatmg
ure mostly preserved in situ.
(v) Envlronmcntul fuctors . can be tackled by the study of the

. f b' t'c
1 factors of the cnv1ronmen 1
Reconstruction o a 10 · ·. . f h hologic features and arrangement.
host sedimentary r~k but exar~m~taon _o t, ~ ~no:ative. The main aim is to get ~ me
of the fossilized animals _i~re_ als~ so~et~;e:n::~; (for marine forms), nature of water
ev .,dences on depth of d1stnbut1on o t . t
·
current, •
temperature, salinity and oxygen ava1 a • aty o f the environ men .
-· 1 bT
.
. . .. . come finer t wards open sea and certain structures like
Broadly elastic sediments be · i . B t these feature s
ripplemarks, and cross beddings are considered shallow _water_ eatures. u ·
are not conclusive evidence for depth . However rain prints, mudcracks are more
conclusive features for littoral zone.
Depth : Observation of bathymetric distribution of pre.ent day animals may. help us in
some way but it is occasionally found that organisms had often changed their depth ~f
occurrence with progress of geologic time. Recent work has . ug~ested that benth1c
foraminifera are much more sensitive to depth and as many as 1x depth zones are
established based on their depth-distribution. However, the best biologial criterion may
be the biomass where the quantity of total population is dimini shing with depth and
with increase of distance from the coast. Here, caution should be taken against the post-
mortem transportation.
Water current : Present day observation has shown that many forms orient themselves
in response to strong water current either passively or actively to get some benefits.
Some crinoids are found holding their arms perpendicular to strong current at shallow
depth but spread them out in low or moderate current at greater depth.
Interpretation regarding water current is mainly based on sedimentological data obtained
from some current oriented features like cross beddings, current ripples etc. However,
references are also made to palaeontological data as and when available. Some organisms
as they have preferred types of elongated shells (such· as Orthoceras, Belemnites,
Turritel/a etc.) align themselves with their long axes parallel to the direction of dominant
current prior to their burial.
Temperature : It is found that distribution of organisms is highly influenced by thermal
conditions of. the environment. Also, various morphologic features may develop in plants'
and animals' body in response to extreme temperature condition. Observation of present
day organisms of arctic, tropical and desert regions may supply us a lot of information.
Rhythmic variation of temperature is sometimes reflected by some cyclical structures
on some organic bodies. The best example is the annual rings found in woods of many
/',,tv.£OECOLOGY
69
higher plants. Xerophytic plants be .
(e.g. cactus). Many molluscan sh c,~me spmy ~it~ very small leaves or without leaf
0
thickness, coiling pattern and col e .s ~ foramrnifera tests show variation of size,
. oration m respo t ..
ammals, commonly taken as ind · nse o temperature cond1t1on . Among
tcator of warm w t h .
most important. Their assoc· t' . a er, ermatyp1c corals are by far the
ia ton with a var' t f
develop upto the limit of ph t' ie Y o green algae has forced them to
o 1c zone.
Salinity : Distribution of species . . .
monospecific assembl b may vary with vanatmn of salinity of sea water. A
. d' t· age, a sence of stenohaline forms, dwarfed fauna are some features
m 1ca mg some extreme salinity cond.,t.mn.-
~~yg~'7t"°n : Organisms in a badly aerated environment are often small in size with
.ms e .5 or tests; often the corresponding sediments in such case becomes black shales
with pynte.
(vi) Study of population of a species

Another aspect of aut~cology is to study of popuh1tion of a single species. Such a study
may reveal a generalized ecological behaviour of a species removing the overprints
caused by some local factors, or taphonomic processes. Density is defined as the average
number of individuals in an area. But density as measured in a fossil species may not
correspond to its original density due to taphonomic overprints. More important is to
study the spatial distribution pattern of the population. It may be random meaning that
if an area is divided into a number of contiguous cells, the probability on number of
individuals being in any one of the cell is the same (Fig. 6-3a).
The more common spatial pattern is termed patchy or aggregated (Fig. 6-3b). In a
random distribution the arithmetic mean of the number of individuals (X) in a cell equals
2
the variance (a measure of spread about the mean S ). In an aggregated distribution
S 2 > X (i.e. variance greater than mean) consequently the ratio of S
2
/x will be one in
random distribution but greater than one in aggregated distribution. Spatial distribution
pattern is difficult to determine by visual observation; it must be determined statistically.
More observations are required for finding the density of an aggregated distribution than
in a random distribution.
The reason of the aggregated distribution is difficult to interpret. Different species are
aggregated for different reasons. Oysters are aggregated on a suitable hard substratum.
Some require selective food zones. Echinoids aggregated at a clear water zone for the
better functioning of their water vascular system. Young individuals are often aggregated
around their parents.
Rarely, over a small area some organisms may show uniform distribution (Fig. 6-3c)
where variance 52 is less than mean. Artificial plantation is an unusual example. Some
calms who live burrowing within the sediment, feed through their long siphons or tubes
by ingesting the sediment in a circular ways ar~und themse!ves. This m°?e of _feeding
encourages the adjacent individual to stay ou_ts1~e t~e feeding ra.nge of its neighbour
and this might have resulted in a near-even d1stnbut1on of the animal.
. " . • ~.':_ . .... -JY . ... - - - - ,~ · - • ' I.. ·~,
-&. - "
- - '
70 PALAEONTOLOG y
6.6.2 Synecological studies or multispecies ecology

Ecologic studies concentrated on more than one species is called synecology where we have
to study the community composed of more than one population of species. Here, instead of
recording density of a single species, we have to record the densities of different species in
each sample. The result is a table of numbers called matrix. Various statistical methods may
be employed to analyze the density of different species within a matrix or between two martices.
Sometimes ecologists prefer to express the relative abundance of different species in a
community from percentage rather than densities. The relative abundance of a species is obtained
as a percentage by dividing the number observed for that species by the number of the total
individual and multiplying it by 100. This is called species frequencies. Community is named
from one/two of its most dominant species(> 50%); others are called companion (10%-20%
and fortuitous species (< I 0% ).
·' l . ·. - -
PART - II
INVERTEBRATE FOSSILS
Chapter 8
PORIFERA (SPONGES)
8.1 SYSTEMATIC POSITION
Sponges are multicellular animals with remarkable simple organization (cells not organized
into tissue). The grade (primitive multicellular) of organization is considered in between protozoan
and metazoan and hence they are sometimes called parazoans. Possibly, they arose from protozoan
ancestry but remained an evolutionary blind ally, not ancestral to any other metazoans.
8.2 MORPHOLOGY (Fig. 8-1)

In general, they are marine sessile benthic animals and filter-feeders. Most have a unicelled-
body-wall made up of colloidal gelatinous matters called epithelial cells (pinacocytes). This is
normally lined by horny materials (spongill) or hard calcareous or sillicious spicules. These
spicules are normally preserved as fossils.
A typical sponge has an upright body. It is bag-shaped or sac-like with a central cavity,
called paragaster, opening at top by osculum. Outer layer is perforated by numerous tiny pores
called ostia all of which lead internally to some incurrent canals and thence to some chambers
(flagellate chambers) which are lined by collar cells or clwanocytes and the latter enclose
radial canals. Exhalant passage leads from these chambers to the central cavity called
spongocoel, and then goes outside by osculum. Each collar cell is globular with a cylindrical
neck (collar) bearing a flagellum which moves constantly inside the cavity causing entry of
water inside through the external os(ia and ultimately to the chambers to central cavity by small
internal ostia and exit of the same from central cavity through osculum. Incurrent canals, lined
by choanocytes, lie in between two adjacent flagellated canals (flagellate chambers) and are
connected with the latter by fine pores called prosopyles. Cells which occur near this pore region
are called porocytes. In between collar cells and pinacocytes occurs mesenchyme layer with
free cells inside the cavity often called amoeboid cells of ameobocytes.
~ of organization of sponges are of three types. In the simplest grade, called asco11,
the sac-like body is merely single-chambered with outer epithelial pinacocyte cells and inner
side lined with choanocytes. A higher grade-form called syco11, has a number of grouped ascon- .
like chambers with a central opening. Majority of sponges are of leucon (rl,ago11) grade in
which a number of sycon-like elements open into the large central cavity (paragaster).
There are about 1500 living genera of modern sponges of which 80% are marine. Mostly,
they are living in shallqw water intertidal zone. A few fresh water forms are found with stalk-
like structures. Fossils of complete or partial sponge bodies, mats of spicules or isolated spicules
are found in rocks extending back to Cambrian or even Precambrian.
97
...
PALAEONTOLOGY
98
Osculum
Flagellate chamber
a.Ascon 0st1um
Epithelial cell layer
(pinacocyte cell)
b.Sycon
Flagella
Collar cell
(choanocyte) ~ ... , ,
~ '\:
Jncunent~ //
.... _ /
d. Flagellate chamber
Mesenchyme Choanocytes
Porocyte
Apopyle or
internal ostia
Endo-pinacocytc
c. A diagra111a1ic sectional view of the body walll
FIG. 8 _ J : MORPHOLOGICAL FEATURES OF PORIFERA (sponge)
""I:)
~-.
i
~
~
~
C;: ·::" "·:· ·::z -
VJ
C::: ::, C!~' ~a ;!:a ~
~
Monaxon ~
Hexactine
Tiiaxon Pcntactine
Tetraxon Octactine Polyactinc
FIG. 8 - 2 : DIFFERENT TYPES OF SPICULES '°

,C)
"
PALAEONTOLOGY
100
U CL~IFICATlO
T,, subphyla of sponge are distinguished :
i Gibtinosa : Pinacocyde.s overlie a middle gelatinous layer called mesenchy"!e in which
spicule""5 are secreted by sclerocytes cells wherein arnoebocytes wander. Spicules often
become tuning fork shaped.
\ii) ·uda : It has neither pinacocytes nor me.s enchyme. Choanoaytes are born in netw~rk
of filaments called trabeculae arranged as chambers. Spicules are five rayed or s1x-
ray-ed.
8.4 SKELETO S (Fig. 8-2

Skeletons of sponges are secreted by some specialized cells called spongocytes or
scluocytes. The skeleton is composed of crystalline or microcrystaJline calcite/aragonite or silica
in opaline form and are called spicuhs. Skeletons may occur also in the forms of spongins
whi ~h are fibre-like commercial sponge) strengthened by foreign inorganic particles.
omenclarure of spicules depends on (a) number of axes (such as monaxon, triaxon,
1arru.o-,, etc. (b number of rays or needle-like terminations corresponding to the direction of
growth such as m.o nactine. diactine. triactine) or (c) size (megaschre and microsclere). A
simple megasclere spicule is monaxon which may be monactine or diactine (needle-like· straight
or curved spicule). Mona.xon can be divided on the shape of spicules (such as oxea, tylote,
styk. acanthostyle etc.). ·
8.5 GEOLOGICAL HISTORY

Porifera_ is a long ran~ing gr~up and the existing forms show a little change of their
morphol~g1cal features since thetr appearance in Cambrian or Precambrian. Most of th
PaJaeo_zo1c sponges had silicious skeletons. Report of calcareous sponges are found fro;
Devom.an onward. ·
Some important sponge fossils are as follows :
Silidous sponges
Palaeozoic Effe.lia (Cam.), Protospongia (Cam) Vauxia (Cam) B h. .
( 0. rd) ff' · · ' . • rac wspong,a
. •. md,a (Ord.-Carb.), Astraeospongia (sil.) Hindoceras (Carb )
Tuusv,lla (Carb.). ' ·'
Mesozoic
Coeloptychium (Cret.), Ventriculites (Cret.), Coscinopora (Cret.).
Calcareous sponges
Palaeozoic G· 1·
myco ,a (Carb.), Meandrostia (Carb.). Cotyliscus (Carb.).
Mesozoic Ste11,spongia
· (Jur.), Corynella (Jur.).
Spo~ge-Jike fossils : Receptaculites (Cam.), lshadites (Cam.~.
Ten1ary sponges are represented by detached . .
sponges have wide distribution in marine d a~d aso_lated spicules and similar isolated
an sometimes m freshwater deposits.
atn is ot\Stl
. I _ _ ,....
v J' . .
O ,.• "\\{'
h .the n otthea'.) l ,, - ·
Chapter 9
CNIDARIA
9,1 GENERAL CHA~CTERS (Fig. 9 . 1)

Cnidarians ar~ the ~implest of all true metazoans. They belong to diploblastic grade where
tissues are orgamzed m two .layers forming the body-wall, the outer 'tctodeaa and the inner
e n ~ n between. them IS present a jelly like layer, called mesogloea through which runs
a simple nerve net (Fig. 9-lc). The body-wall encloses a single hollow gastrovascular cavity
for which the phylum was previously known as Coelentera(a (Gk.-coel : olfo~ ~ nd enteron :
gastrovascular cavity) that included both ctenop ores and cnidarians. At present, they are
considered as separate phyla. The new name CnipJJJ:ia refers to cnidoblasts which are a series
of stingi!!_g cells found within ectodermal layerl£_nidoblast cells are very sensitive to water
vibration and help to catch small organislT)s (passing close to the animal) making the hair like
n~m~tocyst (within cnidob_last)_ ac_tive, killing ~he prey~ injecting a poisonous liquid present"
w1thm the cells through this stmgmg thread (Ftg. 9-ld-e). The single body cavity has only one
opening acting both as mouth and anus. This is surroun ed by a ring of tentacles. The cavity
is lined by endoderm which in some groups (anthozoans) may be infolded to form some radial
partitions called mesenteries making increase of area of food digestion ·(Fig. 9-1 a-b ).
The lower group of cnidarians exhibit phenomenon of altem ation..Jl/ ener, tjpJJ. They
develop two forms : the fixed polyp and the free medusa alternating each other in the life cycle.
Polyps asexually give rise to medusae which reproduce sexually and the zygotes develop into
polyps. However, this phenomenon may be reduced or even entirely eliminated in some advanced
forms where the animals are mainly represented by the-= polypoid or med_r:i~9l_d stage.
The phylum includes animals living both in sea and freshwater condition. Marine forms are
either solitary or colonial. Animals are sessiieat polypoid stage but planktic or nektic in
medusoid form. Their total geological range is Precambrian to Recent.
9.2 BROAD SUBDIVISIONS

Cnidarians are grouped into three classes : hydrozoa, scyphozoa and anthozoa, based on their
phylogenetic relations, nature of life cycle and nature of soft parts and _h ard parts (exoskeletons).
A. Hydrozoa (Fig. 9-2a)

Hydrozoans are bell-shaped/vase-like animals with tentacles surrounding the mouth. They
usually exhibit both polypoid and medusoid stage in life cycle. They sometimes possess chitinous
or ~alcareous exoskeletons. Many posses s foot-likes structure for attachment._ Some forms are
sohtar~, other forming reef-structures (e.g. Mellepora). They are mo~tly marme _but a few are
found m fresh water (e.g. Hydra). The geological range of hydrozoa 1s Precambrian to Recent.
Archeocryptolaria is a common fossil of Cambrian and Ordovician.
8 • Scypbozoa (Fig. 9-2b)
S~yphozoans are mostly free-swimming umbrella-shaped medusae, exclusively marine, with
a radial symmetry. The sessile polyp stage is highly reduced . The common examples are jelly
101
PALAEONTOLOGY
102
_.,._ _ _ _ g
.........::-----9
1.....::.....:::--,~~--wi~~~...
2--~
t? C1lila----6
"~f-----4
~
3------+.~~,..;q.
4-------,;~....,..:,.. :89
(b) A pORTION OF TRANSVERSE SECTION
THROUGH pOL YP SHOWING
DEVELOPMENT OF SEPTA WITIUN
MESENTERIES
(a) A CUT AWAY VIEW OF A CORAL POLYP SHOWING

RADIAL ARRANGEMENT OF MESENTERIES AND
SKELETAL ELEMENTS
l ORAL DISC 2 TENTACLE 3 GULLET 4 MESENTERY 5 MESENTERIAL FILAMENT 6 SEPTUM 7 BASAL
DISC 8 ECTODERM 9 ENDODERM
(c) BODY WALL OF A CNIDARIA

SHOWING DIPLOBASTIC
STRUCTURE
l ENDODERM 2 MESOGLOEA
~ NfRVF NFT 4 FC'TODERM
:'i C'NIDOBl. ·\ST CELL
Operculum 1~~21
Shaft-~~...7 -
Nematocyst-i~l¥-C~~
Coiled threadl--fl...,..lf--~-.1
Lasso.-.....,..._"'
l..lmo
.---Nucleus
(d) CNIOOBLAST AT REST (e) CNIDOBLAST WITH NEMATOCYST

DISCHARGED
FIG. 9-1: WALL STRUCTURES AND INNER FEATURES OF ANTHOZOAN POLYP (coral)
\
~ -
. ... j ., '~-- . , -.
-- --- - i:.-.~.
cNJDARIA 103
s a few of which are found in the fos sil record. Brooksella is a Middle Cambrian fo
ti,s he , . P b. rm.
Their geological. range 1s recam nan to Recent. Precambrian Ediacara faunas poss ibly contain
impressions of Jelly fishes.
c. Anthozoa (Fig. 9-1 a, b; 9-2c)
The Anthozoans are animals with polypoid stage only. They are exclusively marine and they
include such animals like corals, sea anemones, gorgonians and sea pens. They may be solitary
or colonial, living individually or in groups respectively. A solitary form may be radially or
bilaterally symmetrical. Skeleton-bearing colonial groups often form large reef-structures in the
sea. The enteron of anthozoans is divided into compartments by longitudinai partitions called
mesenteries growing in pairs by infolding of endodermal layer that extend inward from the
wa o t e enteron. Towards the base, the mesenteries are free and above, near mouth they
unite with the gullet (a passage from month to enteron). Some anthozoans, like corals secrete
calcareous exoskeleton but others lack it (such as sea anemones). Vertical wall-like skeletal
elements grow from inwardly folded segments of ectoderm, located in between mesenteries
called septa which extend inward from the periphery and reach near the centre. Septa are
intercepted by some transverse partitions called tabulae. They have geological range from
Ordovician to Recent.
From the viewpoint of palaeontologists corals are the most important among all the groups
of anthozoans. They possess hard exoskeleton and left numerous fossils in the geological record
from Ordovician onward. Many corals have their living counterparts in the present seas which
are forming the present day reef-structures. So, interpretation of past environment and
palaeoecology has been successfully made from the study and comparison of recent and foss il
corals.
9.3 MORPHOLOGICAL FEATURES OF CORAL EXOSKELETON (Fig. 9-3 - 9-7)

Simple/Solitary coral A coral living independently.
Compound/Colonial coral A coral where many individuals live together as a colony.
Corallite Cup-like or disk-like skeletal structure secreted by each polyp
around and under itself, also called tlzeca.
Corallum A complete skeleton of a coral, whether a single theca of a
solitary coral or united thecae of a colonial coral. For a solitary
coral corallite and corallum are the same but in a colonial coral
corallum refers to the entire skeleton of the colony while corallite
represents the skeleton of the individual of the colony.
Apex The tapering end of a solitary corallum by which it is attached
to substratum.
Base The expanding or growing side of a coralluni.
Basal plate (disc) Larva of coral after settling to the sea bottom and fixing it to
the substratum, secretes a thin calcareous plate, which gradually
becomes thickened by addition of skeletal materials (Fig. 9-l a).
Prototheca As polyp, attached to the basal plate, grows upward, margin of
basal plate also grows upward and becomes the periphery of a
tiny calcareous cup called prototheca.
Palae(Geo)WP-14
l
PALA£0NTOWGY
\04
(c ANffl()ZOA (eonl)
b) SCYPHOZOA (Medusa)
G. 9-2 : LONGITUDINAL s1::cno

OF THREE MAJOR GROUPS OF
CNIDARIAN POLYP
7
1---~ 3
4---
5---.
~-w--3
"
1
10
11
..--s,
~~~r---6
(b)CROSSS~
(c) SEGMENT OF CORALLUM IN

• THREE DIMENSK>NAL VIEW
(a) A 8fMPlE CORAU..UM
fIO. 9 - 3 : BASIC MORPHOLOGIC FEATIJRES OF AN lDEAL SIMPLE CORAL SKELETON
L BASE (C.tx) 2. APEX 3. MAJOR SBPTIJM 4. MINOR SEPTIJM 5. DlSSEPIMENT 6. COLUMELLA

7. FOSSULA a.SEPTAL TRACE 9.0ROwnt LINE 10.AXIAL STRUCTURE 11, TABULAE 12.lllECA
13.mrnECA
CNIDARIA / 105
/,
Septum As a polyp grows, the lower part · of its inner wall becomes
wrinkles forming mesenteries; some vertical plates from theca
called septa arise radially from basal plate in intermesenterial
position (Fig. 9-1 a, b; 9-3) from infolding of ectodermal layer.
Calyx The basal forms a cup-like depression on which the polyp rests.
It may be deep or shallow and often closed by a lid-like
operculum; the floor of calyx is obviously made up of upper
edges of vertical septa.
Epitheca The upward and outward inflection of basal plate constitutes the
epitheca, which forms the outer surface of corallum and is well
seen in solitary corals. In colonial coral epitheca sometimes
covers the entire corallum. Epitheca may be marked by
successive growth stages of the corallum (tranverse growth lines)
and also sometimes by some long.itudinal ridges divided by
furrows. These ridges mark the junction of septa with epitheca.
Theca The distal parts of septa become united forming the inner wall
of the skeleton called theca. ,
Thecarium Interior space enclosed by the theca.
Shape of corallites In solitary corals, starting from an ideal conical form, corallum
(Fig. 9-4) may be horn-shaped (ceratoid), cylindrical, disco1dal, turbinate,
hat-shaped, pyramidal, slipper-shaped, scolecoid and others.
Corallites of colonial. corals are less diverse and may be tubular
(circular or elliptical in online), prismatic (polygonal outline),
conical or cylindrical.
Type of colony (Fig. 9-5) Colonial corals show a wide range of patterns produced by
various types of arrangement among the individual corallites. The
major types are as follows :
(a) Massive colony : Corallites mostly polygonal, closely packed together leaving no space
in between them. There are several kinds of massive colony.
(i) Cerioid : Corallites joined with one another but each retains its own outer wall and
identity; sometimes the wall of two adjacent corallites are connected by fine
perforations called mural pores. This is mainly found in tabulate corals e.g.
Favosities.
(ii) Plocoid/Astraeoid : The wall between two adjacent corallites becomes fused forming
. a common wall; but septa of each corallite remain unreduced, retaining their identity
e.g. Astrocoenia.
(iii) Thamnasterioid : Plocoid type, but septa of adjacent corallites become confluent,
often twisted making the boundary wall of each corallite indistinct e.g. Orionastraea,
Thamnasteria.
106 PALAEONTOLOGY
u~IB\l~~:s
CYLINDRIC
TUBULAR
PRISMATIC PYRAMIDAL
c
DISCOID
CERATOlD
TROCHOID
SCOLECOID
CONICAL
FIG. 9 - 4 : SHAPE OF CORALLITES
Cerioid
Astracoid Thamnastcroid
m.p :Mural pore p.s: Pscudoscpta
Aphroid Mcandroid
Corallitc·
Ftieicullllcd
Phaccloid
FIG . 9 - 5 : DIFFERENT TYPES OF COLONIAL CORAL..
s ---..
i;trcam ""--:· ...i.:"h ,n1· m,. .... -
CNIDARIA
107
(iv) Ap~roid.: Septa of each cora.llite become short and adjacent corallites are joined by
a d1ssep1metal zone. This is mainly found within some hexacorals e.g. Pachyphyllum.
(v) Meandroid : Corallites are joined in linear series each runs irregularly over the
surface like convolution of a human brain or river meanders e.g. Meandrina.
(b) Chain colony (cateniform) : Tubular corallites are joined only along an axis or line
forming a chain-like pattern in plan view; chains are bifurcating through budding causing
lateral growth of the colony e.g. Halysites.
(c) Fasciculated colony : Corallites are separated from one another but maintain a rough
parallelism. There are two types.
(i) Phaceloid : Corallites are joined by some lateral connecting processes or tubes. e.g.
Syririgopora.
(ii) Dendroid : Subparallel corallites are produced by tree-like lateral branches e.g.
Waagenophyllum.
Type of septum : ·
(a) Based on size and distinctiveness of septa (Fig. 9-3).
(i) Major septum : Larger-sized, extending from periphery to the centre of calyx.
(ii) Minor septum : Smaller-sized, developing in between two adjacent ·major septa, not
reaching upto centre.
(b) Based on ·genesis or time of development (Fig. 9-6b).
(i) Protosepta : The first group of major septa which develop at very initial stage within
the cavity from basal plate. These are six in number and named as cardinal (one),
·counter (one) lying just opposite to cardinal, a pair or alar septa on either side of
cardinal, and a pair of counter-lateral septa on either side of counter septum.
(ii) Metasepta : All other subsequently developed major and minor septa.
Tetracoralla A group of corals in which the six primary septa are aligned in such
a manner that the calyx exhibits four areas or quadrants, two cardinal
quadrants marking areas between cardinal and alars and two counter
quadrants between counter and alars. Metasepta are added in two pairs
at a time within cardinal quadrants and two within counter quardrants
giving a broad bilateral symmetry of calyx.
Hexacoralla A group of corals in which the six protosepta are aligned in six equal
quadrants; two between cardinal and alars, two between counter and
counter-laterals and two between counter laterals and alars. Six
metasepta are added at a time, one in each quadrant giving a six-fold
symmetry of calyx.
Tabulata Corals are devoid of any true septum. Tabulate corals are all colonial.
PALAEONTOLOGY
108
lI
(iii) LAMELLAR (iv) TRABECULAR SEPTA

(ii) FIBRO - NORMAL
COVERED BY L.AMELLAR
(1) TRABECULAR TISSUE
(a) MICROSTRUCTIJRAL ELEMENTS OF SEPTUM
CT
CT er er CTL er crt
(iv) CR
CR
er er
(ix)
(viii)
(vii) CR
(i) .. (v) : RUGOSE (vi) - (ix) HEXACORAL

CR:CARDINAL SEPTIJM CT: COUNTER SEPTUM A: ALAR SEPTUM CTL: COUNTER LATERAL SEP'IUM
CRF: CARDINAL FASSULA MT METASEPTA
(b) PRIMARY SEPTA AND SEPTAL INSERTION IN CORAL
ZIGZAG RHOPALOID
LAMINAR CARJNATE SYNAPTICULATED FENESTRATE
(PFRFORATF.O)
P.E : PERIPHERAL EOGE; AX.E , AXIAL EDGE: C : CARINA; S : SYNAl'TICLi LE: I': PERFORATION
{c) SF.PTAL SURFACE ORNAMENTATION
r Exoscpt11m
~fcscntcrial Pair
Entosc1,1un,-..-':li-.q
(e) A PORTION OF TRANSVERSE SECTION
JNSERT SEPTA (f) CONFLUENT SEPTA
(d)
FIG. 9 - 6 : MORPHOLOGY OF SEPTUM
CNIDARIA 109
Fo~ula (Fig. 9-3) Protosepta of many letracorals or rugose corals al matured stage
became shortened and degenerated produciff\!: depressions or gaps in
their positions on calyx. These are called fossulae . lilphrentis bears a
cardinal fossula in the position of its cardinal septum.
Entosepta and Exosepta (Fig. 9-6e) : In some hexacorals, two types of septa may be seen;
one group lying inside the mesenterial pair called entosepta and the
other lying outside such pair called exosepta.
Insert and Exsert septa (Fig. 9-6d, f) : Normally, the growth of septa is limited within the
margin of calyx which are called insert septa. But in some hexacorals,
septa may extend beyond the margin of calyx, called exsert septa . In
some colonial (massive) rugose and scleractinian corals, septa of one
corallite become continuous with those of adjacent corallites and the
septa are called confluent.
Microstructure of septa (Fig. 9-6a) : Three microscopic components or tissues usually constitute
three types of septa.
(i) Trabecular : Consists of parallel or fan-like needles called
trabecules, each of which is composed of serially arranged
radiating whorls of fine fibres of calcium carbonate (often called
fibre fascicles).
(ii) Fibro-normal : Tissues exhibit a median dark line bisecting each
septum and projecting from it are narrow parallel fibres of calcite.
(iii) Lamellar : Usually occur as cover over a septum, made up of
trabecules or fibro-normal tissues.
A septum may be composed entirely of fibro-normal or trabecular
tissue or a trabecular median part and fibro-normal peripheral part
(Fig. 9-6a iv).
Septa) surface ornamentation (Fig. 9-6c) : Septa) surface may be smooth, paper-like (laminar
septa) or may be variously ornamented as follows :
(i) Carinate: Septa bearing alternate ridges (carina) and furrows
which when become angular called zigzag septa.
(ii) Synapticulated : Septa) surface may be provided with spines or
rod-like structures called synapticules often interconnecting the
two adjacent septa. ·
(iii) Fenestrate/perforated : Septa may be provided with perforations
especially when trabecules are loosely connected.
(iv) Rhopaloid: Abnormal thickening of septa towards the centre.
. ,.. "'1 ., ) ~
I\ f!~ t .._ • ' ,4 J L •
· · - .a,
I'/\ I.I\ h'ON'l'W .nc; \'
\10
A-.;htl Struliturc
~ - - - - - (Spider's wrh)
(i) Top-view (II) Three d rnonstonll view
(a) AXIAL STRUCTUR E
(ii) Mid-calyx (Iii) Central (Iv) Throuat,out calyx

(i) Peripheral
(b) DISSEPIMENTS IN CALYX VIEW
Olu cplmcnt#iun
'fllbularium
(c) TABULAE (complete and Incomplete)
FIG. 9 - 7 : AXIAL STRUCTURE,DISSEPIMENTS AND TABULAE
.• ,,,
(I)
b) PERIPHERAL JNCREASI : <Iii>
a) AXIAl. INCREASE
(d) REJllVENUS I NCb IN A

S0t1TAR Y :ORAL.
Fl . 9 - 8 : TYPES OF AS EXUAL lU~PROll CTION IN 'ORA L
!hh
ftl11\\, a\on g ..,.,.,_..
cNIDARIA 111
Di~piments (Fig. 9-7b) : These are curved (convex inward), somewhat obliquely arranged
plates occupying the space between two adjacent septa. In
tetracorals, they may be present marginally, centrally or throughout
the calyx. In the last case, septa may be almost obliterated e.g.
Cystiphyllum. The peripheral zone where dissepiments are usually
best developed is called dissepime11tarium (Fig. 9- 7b) . In
Hexacoralla dissepimental plates may be present to connect,
adjacent corallites of a colonial coral which are called vesicular
dissepiments.
Pali/Palus (Fig. 9-7a) : Some peculiar growth system of exosepta in some hexacorals may
produce some vertical rod-like pillars (representing separated inner
edges of exosepta) near the axis of corallite. Often they appear in
top view as a star-shaped structure surrounding the central axis
(columella) .
Columella : A vertical rod-like structure extends along the axis running from
base to apex of a corallite; usually formed by dilation of inner septal
edges. Columella may be represented by a solid or porous rod.
TABLE-9
Distinction between Tetracorals and Hexacorals
Tetracorals Hexacorals
l. Six protosepta arranged in four quadrants Six protosepta arranged in six equal quadrants
giving a bilateral symmetry of calyx. giving a six fold symmetry of calyx.
2. Septal surface is usually smooth (laminar) Septal surface usually becomes synapti-
or carinate type, often provided with culated or fenestrate or both.
flanges or denticulations.
3. Solitary or colonial; colonial coral may Solitary or colonial; besides massive and
be massive and fasciculated. fascicuiated type, colonial form may be
meandroid or hydnophoroid (centre of
corallites are arranged around little. hillocks
or monticules)
4. Septa not divisible into entocoelic (ento- Septa divisible into entocoelic and exocoelic
septa) and exocoelic (exosepta) type; type with structures like pillars (palis)
5. Columella, if present, originated by Columella if present, is always of septal
various means. origin. .
6. Corallites in fasciculated colony are Corallites in fasciculated colony are separated
completely separated. by complex perforated tissues coe11ostewn
that consists entirely of exothecal di ssepi-
ments.
7. Septa are usually confined within the Septa are extending beyond the margin of
margin of calyx (insert). calyx (exsert).
Palae(Geo)WP-15
PALAEONTOLOGY
112
: Surrounding columella may lie a cylindrical/co~ical zone
Tabellae (Fig. 9-7 a)
traversed by some fine curved (concave inward) obhque plates .
Axial structure (Fig. 9-7a) . Columella tabellae and pali occupy the axial part of calyx
. forming a~ axial structure where a central columella becomes
surrounded by rod like palis and between them occur tabellae.
· a spi"der 's web at the
The whole structure is looked hke · centre ·
Tabulae (Fig. 9-7c) . These are transverse plate-like skeletal elements in the form of
. horizontal/subhorizontal/convex or concave upward p_lates,
extending partly or all the way across the axial area without
intersecting any axial structure. They are called c~mplete tabulae.
They may be incomplete or intersected by a~ial structur~. A
strongly tabulate axial zone is called tabulari~m. Tabula ts a
characteristic feature of this class, and is present tn all the groups
of corals including Tabulata where septum is absent.
9.4 CLASSIFICATION
Corals are classified mainly on the basis of following characters :
1. Number and arrangement of tentacles and mesenteries.
2. Number and arrangement of primary septa.
3. Presence and absence of septum.
4. Size, shape and relationship among the corallites.
Three subclasses are recognized within corals :
(i) Ceriantipatharia (Rec): Virtually unknown as fossil; solitary or colonial.
(ii) Octocorallia (Alcyonaria) (Ordo-Rec.): Poorly known as fossils, colony forming a flat
fan of anatomizing branches. In polyp, mouth is surrounded by eight stout tentacles
(hence called octocorallia).
(iii) Zoantharia: Solitary or colonial; skeletal structure secreted by basal tissues; septa present
(except tabulata). Four orders have been recognized in them.
Order 1. Tabulata (Ord.-Perm.) e.g. Halysites, Favosites
Order 2. Rugosa (Ord.-Prem.) e.g. 'Zaphrentis, Wentzellela
Order 3. Scleractinia (Trias.-Rec.) e.g. Cyclolites, /sastrea
Order 4. Heterocorallia (Dev.-Carb.)
9.S DIAGNOSTIC CHARACTERS OF THE FOUR ORDERS

Tabulata : Calcitic skeleton, exclusively colonial with slender corallites, often connected
by pores or tubular bodies; septa absent but some short spine-like structures (pseudosepta) may
be present; tabulae present, occurring commonly across the corallites horizontally or may _be
concave or convex upward. The order is named form this tabulae which become its most
diagnostic structures.
Rugosa : Also called tetracorals; the name 'rugose' refers to its wrinkled exterior; solitary
or colonial; skeleton calcitic; six major septa inserted serially in four quadrants; septa mostly
simple; sometimes with axial structure; tabulae present.
cNJDARIA ID
Scleractinia : Also called hexacorals; the name refering to its hard rocky skeleton (Gk-
skleros : hard and aktinos : ray); skeleton aragonitic; solitary or colonial with six protosepta
producing six equal quandrants; subsequent metaseptal insertion in multiple of six; septal wall
with various micro-skeletal structures; septa exsert, axial structures present; dominant reef-
builders of recent oceans.
Heterocorallia : A small branch of Palaeozoic corals differing fundamentally in structure

from the rugose as to warrant separation as an independent order; at the earliest stage four
protosepta meet to form a right-angled cross; so far a few fossils are reported; all are solitary
forms.
9.6 REPRODUCTION AND GROWTH (Fig. 9-8)

Reproduction in corals takes place both asexually and sexually. A new individual or a new
colony grows by sexual reproduction. This produces free swimming larvae (meroplanktic
planulae) which when become matured settle and attach themselves to some object on the bottom
and develop into new individuals or a new colony. However, a colony grows or multiplies by
asexual means where a corallite gives rise asexually to offsets. The term 'budding' is normally
used for this phenomenon. There are three main methods of budding.
(a) Axial increase : In this mode, a corallite splits by fission into two or more daughter
polyps within the calyx. Such budding is inevitably parricidal (i.e. destroys the parent).
(b) Paripheral increase : Here small offsets arise round the periphery of parent corallite. This
may or may not be parricidal.
(c) Lateral increase : This is the most common method of growth found in rugose corals.
This lateral bud may arise fully or partly external to the parent calices.
In many solitary tetracorals, corallites are often found to be constricted at irregular intervals
bearing a broad or narrow shelf where the septate older calyx is exposed. Above this, it may
expand again, before once more been constricted. These constricted bands probably indicate
scarcity of food-supply for some time when the polyp used its own tissues in order to stay
alive, and it became narrower than the existing form. The next period of increase .permitted a
normal growth when the calyx again became wider. This phenomenon of growth is called
rejuvenescence.
9.7 EVOLUTION
Phylum Cnidaria could have been derived from some colonial protistans through the
development of specialized cells and their organization into tissue. Leaving porifera aside,
cnidarias are considered the oldest among the metazoans from their diploblastic body-wall and
some other primitive features and hence they may be even ancestors to all other metazoans.
The probable relationship among the different orders of corals may be summerised as follows.
Two hypotheses have been proposed to explain the origin of corals or zoantharia. The first
postulates that living hexacorals, extinct Palaeozoic tabulates and tetracorals had a common
ancestor in some as yet unknown early Palaeozoic anthozoan stem. Absence of fossil hexacorals
in Palaeozoic may be due to their lack of skeletal parts. By Middle Triassic, most of them
.. . , . a: . .
--- ~ _ y
PALAEONTOLOGY
114
'----~-SEA LEVEL
r.11,.;:;,.:.._.,.:~-REEF
=_::::.:~OLCANIC ISLAND SINKING
_ __::~-- VOLCANIC ISLAND

(c)
(submerged completely)
FIG. 9 - 9 : THREE SUCCESSIVE STAGES OF GROWTH OF CORAL REEF

Ca) fringing reef (b) Barrier reef (c) Atoll reef
(.J
·s Q
N
0
C
u T
u
K
(.J
·s
N J
0
"'
()
~ T
p
I
C
(.J
·s D
N
0
.;
u s
c..
0
:
C
<G
·c
.0
I
.
...
I
e
<G
(,)
... .- .. ... ,,,.-
ct ... .-···
FIG . 9- 10 : GEOLOGICAL RANGE OF MAJOR GROUPS CNIDARIA AND RELATED FORMS
~ ,. \
CNIDARIA
115
developed skeleton-forming habit and they soon occupied the niches left by the extinct tet . I
and tabulates. · racora s
T~~ se~ond hypothesis holds that hexacorals descended directly from tetracorals. The
transition is supposed to have taken place in Late Permian-Early Triassic times.
A third opinion. is that, from an ancestral zoantharian stem appeared first tabulates which
lacked septum .. ~ts i:nay gave rise to tetracorals. There is distinct morphological similarity
betwee~ Ordov1c1a~ t~bulates and rugose corals. Rugose may be the ancestor of scleractinia or
both might have ongmated from a common group of sea anemones. The fact, that some Late
Palaeozoic rugose showed the insertion of metasepta between counter and counter lateral
protosepta like hexacorals may support the hypothsis that a tetracoral could be the ancestor of
hexacorals.
9.8 CORAL REEF (Fig. 9-9)

At present coral reefs are confined to tropical seas and are well developed in the Inda-Pacific
and North Atlantic Oceans. Scleractinians have build up massive wave resistant bodies often
of great thickness. Bulk of these reef materials of the present oceans are supplied by dead
skeletons of colonial corals, although innumerable other animal inhabitants of the reef zone
also contribute variously in forming the reef structures. Structurally, coral reefs belong to three
major types viz., fringing reefs, barrier reefs and atolls. Fringing reefs develop along the coast
line especially surrounding some islands, (e.g. coral reefs surrounding the Andaman Islands).
Barrier reefs develop away from the shore line often endosing a zone of water between reef
and the island. Atoll develops farthest away from shore forming a circular or horse-shore shaped
ridge encirculing a central lagoon. The Great Barrier Reef of Eastern Australia is a linear feature
parallel to the fault-bounded margin of the continent and the reefs grow on the edge of an
upraised fault. It is assumed that these three structural types may indicate successive stages of
growth of reefs around a sinking or rising island.
9.9 ECOLOGY
Almost all corals are sessile benthic in habit. Observation on fossil corals of Palaeozoic
(tetracorals and tabulates) has led to their separation into two distinct facies : a shallow marine
platform facies and a deep marine basinal facies. The former type of corals are complex, mostly
colonial, associated with carbonate facies. The latter, associated with shale facies includes mostly
solitary corals. This two-fold divisions of Palaeozoic corals is broadly analogous to the divisions
of scleractinians living in recent seas. One group includes structurally complex, mostly colonial
forms caJled hermatypic corals, which are the important reef builders in the recent tropical seas.
Others are ahermatypic corals, structurally simple and solitary forms living at variable depth from
shallow to deep marine zone (most common upto 500m where the temperature range of water is I
50° to 10°C). From ecological point of view, hermatypic corals are the most significant as they "
are restricted in the shallower part of the tropical oceans with depth ranging between l Om to
50m with temperature of water between l 8°C to 29°C. In hermatypic corals, the endodermal walls
are repleted with symbiotic algae, (dinoflagellate Zooxanthalle). These algae are so essential to
the metabolism of corals, supplying it with nutrients, oxygen and calcium carbonate, that
hermatypic corals can only be flourished in the photic zone of the sea (normally 50m depth) upto
which sunlight can penetrates as without sunlight algae cannot perform photosynthesis. Thus fossil
reef-corals definitely indicate a particular type of shallow marine condition .
...~
,,, • .,
,"Y..,. . "I
• • • •,
\. . ~ . ~• ... I ,---
. ' ~- ~ f
.
.'
.
·•···. ·. .., ·.f _:·.-~-
l • ..
· --··· < ~ -... _:- . ~~4;_

,. ,
. . •
. } - .
,
~ .. -. r,..'.
.
•.
; .
'
.
. ... .
_.,
PALAEONTOLOGY
116
Coral reefs have built up massive wave resistant structures and the symbiotic algae have
forced the corals to construct such reefs in the warm tropical sea within the limit of photic
zone. These reefs often become of great thickness and extent where bulk of the materials are
contributed by the corals. Such a huge rigid structure built up by corals of several generations
becomes the habitats of a host of diverse organisms, and they together form one of the most
complex marine ecosystem. This shaJlow zone of the sea, lighted by the sun, with constant
wave actions also becomes the site of considerable accumulation of planktonic protistas which
are the source of food of many other invertebrates and fishes. The coral reef's productivity,
calcium metabolism and carbonate fixation are very high. The range of habitats is such that
there is a high degree of specialization in the associated faunas which include fishes, different
groups of benthic molluscs, brachiopods, arthropods, echinoids, crinoids, bryozoans, etc. All
) these animal groups are found in zones of localized niches at various parts of the reefs.
'\
9.10 GEOLOGICAL HISTORY (Fig. 9-10)
First record of tabulate corals is known from Early Ordovician rocks. They attained their
) climax in Silurian-Devonian. They were declining in number since Late Devonian and became
e~tinct toward the end of Palaeozoic. Many of its species however, attained a widespread
dispersal and are used as index fossils such as Favosites forbest (Ordovician).
Tetracorals a~so appeared in Ordovician and attained their climax by Silurian-Permian times
but t~ey also fatled to cross the Permian boundary. Solitary forms like Calceola, Zaphrentis,
Cystiph!llum, Omphy'!"'1 a~pear to. be inhabitants of deeper part of the sea while the colonial
forms like Acervularta, Ltthostrotwn, Lonsdaleia lived in shallower part of th d
often found associated with stromatoporoid reef-s~ructures. e sea an are
Fossils of hexacorals are first recorded from Middle Trissic rocks Th d 11 ·
l
in numbe~ since their appearance and became dominant after Tertiar e~;ra ua y mcreased I
abundant m present tropical/subtropical seas where they are b 'Id' y. h ey are very much
structures. ui mg t e present day reef- /
·t
9.11 STRATIGRAPHIC USE OF CORALS
A. Corals as geologic age indicators
/ Although, scleractinian corals are mostly Ion ra .
tubulatas left many species which are used . dg- &ng1~g forms, Palaeozoic tetracorals and
h' ·t . as m ex 1oss1ls The wid .
t is sess1 e benth1c forms was possible through th . . e geographic dispersal of
known index fossils are as follows : ear meroplanktic planulae (larvae). Some well-
Name of species Age

Tabulata Favosites forbesi
Lower Ordovician
Halysites wallichi
Lower Silurian
Rugosa I
Calceola sandelina
Middle Devonian
Cystiphyllum cristatam Devonian r
Waagenophyllum indicus
Streptelasma corniculum
Permian
Middle Ordovician l
rt .... -
. , ... 11\t. along w1u ..... - --
Cf/JDARIA 117
On the basis of such index fossils Palaeozoic sequence of Burma has been biostratigraphically
zonated.
8. Corals as geochronometers
Accurate observation on many solitary rugose and scleractinian corals shows their epithecal
surface with fine growth lines (about 200/cm). Each of these lines indicates former position of
the rim of the calyx. Again the fine ridges may be grouped into some prominent bands or
annulations between which epitheca is somewhat constricted. It is assumed that these fine growth
lines represent daily increment and the coarser band monthly and still wider and broader
annulations represent yearly growth . Wells (1963) and Scrutton (I 965) working on Devonian
rugose corals counted the number of growth ridges per annual annulation and concluded that
a Devonian year had an average of 400 days. This figure also roughly corresponds to the
astronomical data. Thus the coral as a 'palaeontological clock' may provide a consistent check
on estimate of rate of change of the earth's axial rotation since Palaeozoic.
9.12 SOME CORAL-LIKE FOSSILS OF UNCERTAIN AFFINITY

A. Archaeocyathid (Fig. 9-11 a)
Archaeocyathids are a very early group of calcareous fossils found in Lower to Middle
Cambrian times. Although, they are grouped as a separate phylum, they are thought to be as
being of a grade or organization comparable to that of a porifera to which they might have a
biological affinity. From their associated fossils, -it is inferred that they were shallow marine
sessile organisms associated with stromatolitic bioherms or as isolated individuals.
Archaeocyathids are normally found as solitary organisms bearing superficial resemblance
to corals and/or sponges. The conical calcareous skeleton is called cup which is no more than
a pair of inverted cones, one inside the other, joined with each other by some vertical but radially
arranged partitions each called pariety or septum. The latter divides the outer annular cavity
inlervallum into a number of segments called loculae. The large central cavity enclosed by
the inner cone seems analogou~ to the paragaster of a sponge. The lower part of the cup is
usually expanded into a basal flange with finger like lioldfast. The outer and inner wall are
perforated by numerous holes (analogous to ostia of a sponge) arranged in some longitudinal
rows. Parieties are also sparsely perforated. The wall is usually composed or polymicro-
crystalline calcite. In addition to this there may be some accessory structures within intervallum
like some radial rods (synapticula;), perforated transverse plates (tabulae) or imperforated
arched plate called dissepiments analogous to those of corals.
,. Archaeocyathids are commonly classified into two groups. Such as 'regularies' and
tcgularies' depending upon the presence and absence of symmetry of the skeleton. Mostly
t ey are symmetrical.
adhNearly all archaeocyathids came from carbonate shelf sediments deposited in warm sea. Other
as, er~d to substrate by their holdfast or similar device. They are most commonly found ·
bu~tted with algal stromatolites and also with triolobites and brachiop~ds of Cambrian age,
archac:;aret~ found in association with sponges (ecological competitors?). Occurrence of
rocks ha:athid fossils are commonly patchy in nature. Their abundance in Lower Cambrian
· enabled paleontologists to use them as a stratigraphic marker of this age .
...
PALAEONTOLOGY
-----------r Central cavity

_ _ _ _ __,,uutcr wall
ll8
..__ _ _ _.,. Locu\a
~----"'arict)'
Inner wa\\
(a) STRUCTURE OF AN ARCHAEOCYATHEA
-
(b) CONULARIA -. Tabu\ac
- \, . Strong
Mamelon . A
- Pillar
-
... . - ,
I W'
- .,.
-- - ~ ;
i::::: ~
~ ~
Vertical Section Of Lakchia
Fractured Part Of Colony
Actinostona (Vertical Section)

Showing Equally Developed
Pillars & Lamcl\ac
(c) STROMATOPOROIDS
FIG. 9. 11 : SOME CORAL-LIKE FOSSILS OF UNCERTAIN AFFINITY .
CNIDARIA 119
B. Conularids (Fig. 9-11 b)

Conularids are a small group of extinct marine animals whose biologic affinity is still
uncertain. From their remarkable simple shell morph.ology it becomes difficult to find out their
actual systematic position. They resemble coelenterate polyps/medusae, more closely allied to
scyphozoans. Or, they may represent some simple uncoiled shell of gastropods. Recently, some
tentacle-like structures resembling those of corals are found preserved in a few fossil-samples
suggesting their probable affinity to corals. Association and mode of preservation suggest that
they were sessile at least at their earliest ontogenic stage.
A typical conularid has a thin, somewhat fragile chitinous shell, radially symmetrical, conial
or pyramidal in shape. The shell opens at the apertural end and closes towards the narrow
apical end by which it was possibly attached. There are combination or transverse and
longitudinal ridges and furrows on the outer surface. Prominent furrows are often found to be
present at the junction of two faces in case of a pyramidal shell.
Conularids are more or less restricted in Palaeozoic although a few are known to be present
in Triassic. Many species have short vertical range and wide geographic dispersal which might
indicate their pelagic nature like the adult scyphozoas .
. C. Stromatoporoids (Fig. 9-11 c)

Stromatoporoids are calcareous masses or layered structural materials found in marine
carbonate sequence of Cambrian-Cretaceous age but are most dominant in Silurian-Devonian.
They were important reef builders in Lower Palaeozoic. Stromatoporoids have been considered
by different authorities to be hydrozoans, sponges, encrusting foraminiferas, broyozoas, algae,
or as a separate phylum which has no living counterpart. The suggestion that they are hydrozoan
is based on the presence of such structures as astrorhizae which are somewhat analogous to
hydrorhizae of hydrozoans. The modern reef-building hydorzoan Millepora resembles in many
respects stromatoporoids. Skeletons of some recent encrusting sponges of the Pacific Sea named
as Sclerospongia resemble those of stromatoporoids. Many Palaeozoic stromatoporoids were
reef-builders and since they had geological requirements similar to those of reef-building tubulate
corals, these two reefs are often found together and some authors like to put stromatoporoid as
a type of tabulate corals.
The structure of the colony may be studied in horizonal and vertical section. The skeleton
of the colony is called coenosteum. Colony shows irregular layered structures forming thin and
flat sheets or rounded masses (mamelons).
In vertical section it shows some stout upright pillars joined and traversed by horizontal
laminae often called tabulae. Pillars define the individual chambers. In Actinostoma pillars and
laminae are equally developed whereas in labechia pillars are more stout and strong than
laminae.
Palac(Gco)WP-16
Chapter 10
BRACHIOPODA
10.1 INTRODUCTION .
Brachiopods are small sessile· benthtc · ·mverte brates h·avm· g an exoskeleton
· . . (shell) covering
· respect, they have some superfitcta
the soft parts. In this ·, I rese
, mblance with bivalved molluscs. . .
· · f
In both the cases, the shell 1s composed of a pair o mge vh" d alves · Both are fed
. by draw1110
. . 0
· · the shell and filtermg

water mto · off food part1c· 1es. B ut zoo Io g1·cally they are .qmte different as
regard their body plan, symmetry, ahgnment o f va Ives an d se veral other internal fe atures.·
· ·
Brachiopods are animals of triploblastic coelomate grade with oligomerous body plan .
A common character of all brachiopods (Fig. 10-1) is its possession of a shell made up of
two unequal but equilateral valves hinged along their posterior margin . Of the two valves, the
one, normally lying on the ventral side of the animal, becomes larger. This larger valve al so
bears internally a fleshy stalk-like body called pedicle or peduncle at its posterior end, used for
attachment of the animal. This valve is commonly designated as ventral or pedicle valve. The
smaller valve usually lies on the dorsal side of animal and internally bears the food-collecting
organ called brachia for which it is called brachia/ valve or dorsal valve. Early investigators
mistakenly thought that brachia was homologous with the foot of mo11usca (used for locomotion)
and hence gave the name of the phylum 'Brachiopoda' (Gk.-brachion : arm and podos : foot ).
Though, the valves are unequal they are equilateral and that gives the shell a bilateral symmetry
where the symmetry plane passes across the two valves cutting both equally along the anterior-
posterior line. Internally, the animal bears a complex food-gathering cum respiratory organ called
lophophore or brachia, supported by a loop-shaped calcareous ribbon called bracliidium which
remains attached at two points inside the brachial valve. Depending upon the nature of articulation
of two valves, brachiopods have two broad divisions. 'hiarticulata', where the two val ~es are
attached by muscle and 'Articulata' where the valves are hinged posteriorly by teeth and sockets.
Inarticulate brachiopods usually show epifaunal or burrowing life habit (Fig. l 0-2a, b ).
10.2 STRUCTURE AND COMPOSITION OF SHELL (Fig. I0-3a-f)

'f!1e shell is secreted by cellular epithelium. In all articulate brachiopods, the shell wall is
multlla~ered. These are : an outer non-calcareous ~erio_stracum, a middle calcareous primary layer ·
and a~ mner seco~dary la!er of calca_reous and inorganic materials. In inarticulates, secondary
la~~r 1s absent while t_he pnmary layer 1s composed of alternating bands of calcium phosphate and
ch1tm. The mantle which secretes the shell is in folded under the shell edge thus.. · t·v..
· 101mmo 0 a 0oenera I "
zone from which all the shell-secreting materials are produced Microscopi·c ob
·
three types of shell structures within articulates brachiopods (Fig. 10-3c-f).
- h
servat1on as s o,,
h ,11
(j) Impunctate : Here no perforation or cavity is found within the shell wall.
(ii) Endopunctate : Shell structure is penetrated from the inside b I I)'
. . y some Iarge regu ar
arranged e Iongate d cav1t1es (pu11ctae) normal to shell surface Th · b liar
111
outgrowths of the mantle called caecae . A coecum nearly rea h · th ey contha tu t~ ·e
c es e outer s e 11 sur ac
I ?O
, ......,.... ~
........ . ·.. J .. .....
. . . ··~
'
_... j .. . ::__ .,.
• ..,., .... _, . .· .:, , f.• •. i"
BRACHIOPODA 121
(a) NORMALLY ATIACHED SHEEL
M
Pv
Cp
-A
Pd
(b) LONGITUDINAL SECTION THROUGH THE SHELL
FIG. I 0-1 : ATTACHMENT AND INTERNAL ORGANIZATION OF A TYPICAL ARTICULATE

BRACHIOPOD
A : Anterior Abw : Anterior Body Wall Adm : Adductor Muscle Am : Adjustor Muscle
B : Brachidium Bv : Brachia) Valve Cp : Cardinal Process Dm : Deductor Muscle Fg : Foodgut
L : Lophophore M : Mouth Mc : Mantle Canal Mcv : Mantle Cavity Me : Mantle Epithelium
P : Posterior Pd : Pedicle Pv : Pedicle Valve Sw : Shell Wall Be : Body Cavity
Brachia Brachia) Vl!lve
Pedicle Valve
(b) EPIFAUNAL-PEDICLE OPENING

ON PEDICLE VALVE
Pedicle ---+---t::i
(altached)
(a) BUPROWER-PEDICLE OPENING SHARED BY BOTH THE VALVES
FIG. 10-2 : INARTICULATE BRACHIOPODS
PALAEONTOLOGY
122
Anterior
M:ugin
Ii
, J
Periostracum
Phosphatic Malcrial .
Organic Material
a. ARTICULATE BRACHIOPOD Anterior

SHELL IN CROSS SECTION Margin
\ .
b. INARTICULATE BRACHIOPOD
SHELL IN CROSS SECTION
(ii)
c. IMPUCTATE SHELL
(ii)
d. PSEUDOPUCTATESHELL
Caecum
(ii)
(c) ENOOPUNCTATE SHELL
f. A CAECUM CENLARGED)
FIG. 10- 3: SHELL MICROSTRUCTURE OF BRAC

. HIOPOD
(i) Cross Section, (ii) Top View Of Shell Aft E . .

. . er ross1on Of Periostracum
Epl : Epithelium, Pl : Primary Layer.. Ps . p .
.' >' . er1ostracum, SI : Secondary Layer.
,:_ .... .- --, . "i:\ . .. ..... "Wlllt, -
BRACHIOPODA 123
( Anterior
Anterio-Posterior
Line Of Bilatera1----••a.i.
Symmetry
Anterior
w Side View
T
Dorsal
View
Plication (a) ORIENTATION & DIMENSIONS

Hinge Line Umbo ...___~-....~.,,,,... _...Oll!"---..1
Beak 0
Pedicle..,._ __ Brachia!
Valve
Valve
Median Sulcu,~--.....1 Anterior Linc Of
----•Commissure
Pedicle (Ventral) View
Brachia! (Dorsal) View
(b) BASIC MORPHOLOOIC FEATURES
~oc
·, ~.2 QVO 1rcu ar
(Subcircular)
(c) SHELL-FORMS IN VALVE-OUTLINI;
(Transverse) Elliptical
Diconvcx Rcsupinate
Concavo-Convex Plano-Convex Convexi-J>lane Convexi-Concave
(S-Shaped)
(d) SHELL-FORMS FROM SIDE VIEW (Along Plane Of Symmetry)
Umbo Of P-Valve
Umbo?fD-Valve~(i)
Hmge Area
(ii) 0(iii) ())(iv) @(v) 0,(vi)
~~--uf 8-Valve
(Dorsal
Straight
b s
Erect . IIlly
SI 1g ..
:Strong Iy I ncurvc J View)
u erect lncurved
(e) NATURE OF VENTRAL UMBO IN SIDE VIEW
<::> ~//
Strophic Non-Alate Strophic-Alnte
u O·
Non-Strophic-Straight Non-Strophic Curved
(f) TYPES OF HINGE LINE
FIG: 10 - 4: EXTERNAL MORPHOLOGY OF BRACHIOPOD SHELL
-· . ,.
PALA EONTOLOc; y
124
\
I
I
I
;
, I
,,, /
~
"'
I
,/ • Straight
Acute Right Angle
B-lnterarea
Posterior Margin Foramen

Of P-Valve
Posterior-Margin elthyrium ,\t, 1 ,\,.; IJ..:llh) rJUIII .\l •, 1\ ,.; LJ..:l1h)llllllt

Of B-Valve Central-Delthyrial Basal-Delthyrial (On Beak) (Over Beak)
Apical-Delthyrial (Hypothyrid) (Amphithyrid) 1 f'l·rmc,nrhnid I 1F. rirh ~ricf l
(Mesothyrid)
Bisulcate
~ffRENGFFO~ &
Unisulcatc
Rectimarginate Uniplicate Biplicate
Muliplicate
(d) DIFFERENT TYPES OF ANTERIOR MARGIN (As Viewed Keeping Brachia( Valve Above The Pedicle Valve)
··- Growth Line~

Capillatc Costate Reticulate &
Tuberculate
Spinosc
Rugose
(c) DIFFERENT TYPES OF SHELL-SCULPTURES

• Ribbed
FIG. IO - 5 : EXTERNAL MORPHO!-,OGY OF BRACHIOPOD SHELL
..
~ ~ .. :', , -- ·
BRACHIOPODA 125
and is connected with it by a brush of tiny tubes filled with muco of saccharid~. Below
this brush are ~c!.J:_ls filled with glycogen and protein which hangs within the empty
cavity below. Caecae primarily function as storage chambers; they also function in
respiration and inhibit possible boring by predatory animals. It is also assumed that
the brushes can secrete an organic glue which may be used for repairing the periostracum
layer as and when necessary.
(iii) Pseudopunctate : Some stro homenid shells exhibit some thin and often irregularly
spaced taleolae which give a false impression of punctae within shell-wall, especially
in weathered samples where the outer periostFacum is lost. These are called
pseudopunctate shells.
10.3 EXTERNAL MORPHOLOGIC FEATURES (Fig. I 0-4, I0-5)

A. Shape of shell (Fig. 10-4c, d)
Valve outline
Ranging from circular, subcircular, triangular, rectangular, rhombic, elliptical etc.
Side view
Concavo-convex, planoconvex, biconvex, convexi-plane, convexi-concave, resupinate
(s-shaped) (conventionally, in such a description, character of brachia) valve is mentioned first).
B. Features associated with posterior side of valves (Fig. I 0-4b-f and I 0-5a-c)
Umbo : Posterior protruburence of each valve; ventral valve umbo is larger
and more compicuous than that of dorsal; umbo may be straight, erect
or suberect, slightly or strongly incurved (Fig. J0-4e).
Beak : Pointed extremity of each umbo which marks the beginning of shell
growth.
Beak ridge : Posterior lateral margin from beak to cardinal extremity; may be
entire, dentate or spinose.
Hinge line : Posterior line of junction of two valves; this may be straight or
curved; long or short.
Hinge axis : Line passing through the two points of articulation (teeth and socket
along hinged area); it may or may not coincides with hinge line.
Strophic shell : ·A shell showing straight and long hinge line, sometimes covering the
maximum width of shell; hinge line and hinge axis are nearly parallel
in this case (e.g. Spirifer).
Non-strophic shell : Hinge line curved, shorter than shell-width and not fully coinciding
with the hinge axis (e.g. Terebratula).
lnterarea : A plane or logitudinally curved surface lying between beak and
posterior margin of each valve (break ridge) and hinge line; True
interarea is found only in a strophic shell.
PALAEONTOLOGY
126
erior portion of either val ve; interarea
Palintrope : Recurved or turne d- back post· . .
. 1 pe "aces opposite val ve m strophic shells.
1
deve Iope d on pa m ro 1, •
Card.inal (lateral) : Lateral terminus of hinge line; may be angular and ~harp ~r rounded ;
may be provided with ear-like extension s on either side called
extremity
auricles.
Alate : Shell bearing auricles.
Intera~ ngle
(Fag. I0-5a) : Angle subtended by the lines joining the two beaks and the trace of
anterior line of commissure. This angle becomes variable ranging
from acute, right angle, obtuse, straight ( 180°) and reclined (> 180°).
Pedicle opening : The opening through which the flesh y stalk pedicle protrudes out for
attachment t)f shell with substrah.'. In inarticulates it is simpl y a
posterior gap between two val ves, but in articulate it is a restricted
zone shared by both the valves but mostly situated only on pedicle
valve.
Foramen =.~A circular or subcircular opening, large or small, restricted to pedicle
valve for the passage of pedicle; generally a feature of non-strophic
form (e.g. Terebratula) where hinge is poorly developed. The position
of foramen is variable (Fig. 10-5c).
Delthyrium : Be~eath the be~ of pedicle valve located on the pedicle hinge are~
a tnan~lar ~penmg/~otch for the passage of pedicle, often extending
upto hmge lme, mechally besecting pedicle interarea.
Notothyrium : A simil~ but smaller structure like delthyrium but situated on brachi a)
valve hmge area. ?pening for pedicle is often shared by both the se
structures (delthynum and notothyrium) .
Deltidium and : ~ingle plate partially or completely closino .
Chilidium num respectively These o delthynum and nototh y-
. I
Pates are formed whe th pec1· ·
or non-functional. n ·e 1cle 1s absent
Deltidial and : Each of the two areas (dethyrium and .
Chilidial plates by a pair of plates. For partial I notothynum) may be closed
c osure pedicle
as a small foramen situated t
delthyrium. Accordingly t: ' .
.
openmg may occur
any place_ along the mid way of
amphithyrid (basal), hypotiiyri~\ a~~ designated by the term .
permesothyrid (top of delth . mt die), mesothyrid (terminal)
. ynum) and ·1 · '
(F ag. I0-5b-c). epi1 iyr,d (above delthyrium
C. Features on shell surface (Fig. J0-5e)

Growth lines : Markings on sheJI surfa
. . ce parallel to val ve, . .
successive growth stages st . s antenor margin indicating
. anang from u b -
concentnc growth lines a II m ona l region. Very coarse
re ca ed rugae.
~
. ' . ,._ ,,.,
·-·--
BRACHIOPODA 127
Radial lines : Markings of some ridges and furrows radially disposed from umbo
towards anterior margin; they may be of following types; capillae or
fila (very fine), costae (moderately coarse) and ribs (very coarse
ridges and furrows).
Reticulate structure: Equally prominent growth lines and costae sometimes produce a mesh
or reticulate or checkered surface on some brachiopod shells (e.g.
Productus).
Tubercles/Spines : Raised spherical/angular body often developed at junction point of
growth line and costa; spines may be erect or oblique.
Squamose : Very coarse growth lines, may become raised at point of intersection
Ombricate structure) with costae.
Median sulcus : A major rounded median depression on ventral shell surface along
longitudinal mid-line.
Median ridge : A major rounded elevation on shell along longitudinal midlinc of
brachia! valve and generally correspond to the median sulcus of the
pedicle valve.
The median sulcus and ridge in many genera seem to be instrumental
in separating the two lateral inhalants from a median exhalant stream.
D. Features of anterior margin (Fig. I 0-5d)

Line of Commissure: Line along which the two valves are joined anteriorly.
Anterior margin : Anterior margin may show various patterns when viewed keeping
dorsal valve above the ventral. These are as follows :
(i) Rectimarginate : A smoothly curved or straight anterior margin (produced when the
shell grows uniformly along radial, lateral and vertical direction).
(ii) Unisulcate : Produced by a single sulcus on pedicle valve with/without a
corresponding ridge on brachia! valve (e.g. Productus). In bisulcate
margin pedicle valve has a sulcus on both side of a central plic~.
(iii) Uniplicate : A single plica (ridge) at the anterior margin with two troughs on
either side (e.g. ~pirifer). Purpose of such a large uniplication is to
separate inhalant and exhalant current, sometimes exhibited by some
t rhynchonellids and spirifeij9s.
(iv) Biplicate : Uniplicate margin broken by a median sulcus. (e.g. Terebratula) .
(v) Multiplicate : Margin with numerous plications and troughs. (e.g. Rhynchonella). '
i.. .
(vi) Trail : Anterior prolongation of some brachiopod shells, generally at high
angles to the posterior (e.g. Rhynchone/la).
- i
Palae(Geo)WP-17
l'ALA EONTOLO<.iY
128
- - - Dent11l Phttc
- - - Tooth
Spondylimn ---oWJ-" , __--,.....-Dclthyriol

Median Scptum.--~-.-..;-f!!r..i ·avity
Adductor
Mu1clc Scar Median Septum
f;nl/1/e.,
----..'lo---- Median Septum L'!Jlh..-w,p,r;:-

----s1111rr
Brachidium
(Forming Loop)
C1mlinul Process
Myophorc
(b) INTERIOR OF BRACHIAL VAL VE
Jugum (d) BRACHIAL rNSIDE OF HEBERTELLA
(e) INTERIOR OF BRACHIAL

VALVE SHOWING
G) (@@) (!;~ 0
A trypo1'd s·ptr11cro1
· ..,. 'd .
Alhyru1d
Tc:rcbrn1ulo1d
(Non. oiled)
BRACHIDIURM
(f) VARIATION OF LOPHOPHORE
FIG. JO - 6: INTERNAL MORPHOLOGY OF BRACHIOPOD SHELL
Dm
(i) LongjtudinaJ section
Dm
(b)
BRACHIOPODA 129
E. Dimensions and Orientation (Fig. 10-4a)
Anterior : Part of shell adjacent to midline at extremity opposite to beak.
Posterior : Part of shell along midline at umbonal side.
Length : Line joining anterior and posteri<:>r end.
Width : Maximum distance of two lateral margins of the shell.
Thickness : Maximum distance across the two valve between dorsal and ventral
surface.
10.4 INTERNAL MORPHOLOGIC FEATURES (Fig. I0-6a-f)

Features inside the two valves are quite different. Morphologic features present within the
brachial valve are important especially in the classification and recognition of different genera
within a homeomorphic group. However, fossil brachiopod valves are mostly found in closed
condition and fossils showing interior of valves are very rare. The important internal features
are as follows :
A. Features of pedicle valve associated with articulation (Fig. 10-6a)

Hinge teeth : Projections on hinge line of pedicle valve, one on either side below
the delthyrium.
Dental plates/ : Vertical/oblique plates extending from the floor of the pedicle valve
lamellae to the edge of delthyrium supporting hinge teeth.
Delthyrial cavity : Dental plates divide the space below the umbo of pedicJe valve into
three compartments, the middle one is called delthyrial cavity.
· Spondylium : Curved platform for muscle attachment inside the beak region of
pedicle valve usually formed by union of two dental plates. ·
Median septum : Calcareous wall, built up along mid-line of each valve. In pedicle
valve it supports the spondylium.
B. Features associated with brachial valve (Fig. I0-6b-t)

Cardinalia : Collective terms for all structures inside the beak region of the
brachial valve functioning for articulation, muscle attachment and
brachia] support. It includes the following elements.
(i) Dental sockets : Two depressions on hinge plate of brachia! valve for articulation with
the two corresponding hinge teeth of the pedicle valve.
(ii) Hinge plate : Divided or undivided platform inside beak region of brachia! valve
joining dental socket\ n11d crural bases; may be divided into outer
,·
and inner part. '
(iii) Crus (pl: crura): Proximal part of calcified brachia! support, attached to hinge plate.
(iv) Crural base : Projection from hinge plate on either side of notothyrium for attach-
ment of crus.
··- I'
PALAEONTOI OG Y
130
(v) Crural process : Inward oblique projection of crus facing opposite valve supporting
brachia.
(vi) Crural Iamellae : .One or two short plates (analogous to dental lamellae of t~e pedicle
valve) adjoining the cavity below notothyrium in beak region of the
brachial valve.
(vii) Ala : Lateral flange outside of each crural lamella.
(viii) Cruralium : A spoon-shaped structure produced by union of two crural lamellae

at the centre of beak region of the brachial valve (analogous to
spondylium of the pedicle valve).
(ix) Cardinal process: Projection at or near the beak region of the prachial valve for
attachment of deductor muscles; axis of cardinal process is called
shaft.
(x) Myophore : Zone of muscle attachment at the top of cardinal process.
(xi) Brachiophore : Short and stout projection from beak inside the brachial interior of
some primitive forms for attachment of ]ophophore. In advanced form
it is replaced by crural base.
(xii) Brachidium : Simple or complex calcareous support for fleshy lophophore or
brachia. Different brachiopod groups exhibit different types of
brachidium. Crus is the most simple form of brachial support found
i~ rhynchonellids. Spiriferids exhibit brachidium as spiralJy coiled
ribbon laterally place~. _These are called spiralia which may occur
separ~tely or may be J~me~ ~y a rod-like process along midway of
brachial va_lve_ whe_n this pair 1s calledjugum. Usually there are four
types of sp1ral1a .(Fig. I0-6f) : atrypoid ·· the tips of two sp1ra· 1·ia +,acmg·
eac h· other
·· on either side of bilateral symmetry plan e, spiriJero, .
. · ;r. 'd
the m1t1a.1 part of each spiralium goes mid-way o f va Ive an d from·
there spirally grows towards posterior-lateral s · d k·
almost parallel to the hinge line· atlzyroid . g . i ~· .":1a mg them
·d · . ' · rowmg initially towards
m1 way, each spirahum coiled laterally backward· t b .
. I b h. . , ere ratulo,d . non-
sp1ra rac idmm, looked like a calcified loo .h . .
attached to cardinalia by crura. P Wit the open sides
C. Muscular structures and_ valve attachment (Fig. I0-7a, b)

Two. valves. are mostly activated. (open and closed , by som e muse 1es, areas of h. h 1
some 1mpress10ns or scar-marks m the valve interior. Three t f · w ic eave
adductor muscle working for closing the two valves· deducto ypes ° muscles are present :
' r muse1e helping in . h
two valves; adjustor musele helping to adjust the shell when it is . tt h _openmg t e
near the base of P-valve) joining it with pedicle. a ac ed by ped1cle (present
In normal condition, the two valves of a brachiopod shell are clos d b

. e Y two adductor . I
which cross the two valves more or 1ess perpend1cularly on inner sid Of muse es
e two deductor muscles.
BRACHIOPODA 131
The latter run obliquely below the umbo to the inner posterior margin of pedicle valve. By
contraction of deductor muscles the animal can open the two valves along the anterior margin
(Fig. 10-7a, b). As muscles become inactive after the death of the animal, the shell of the
brachiopods are usually preserved in its normal closed condition. The edge of the entire mantle
or part of it due to folding leaves some sinuous markings in the interior of valves called pallial
markings.
D. Functional mechanism of lophophore

The only major organ within the mantle cavity of a brachiopod is the loplwpliore which is
principally a food-gathering and respiratory apparatus of the animal. The lophophore has a
hydrostatically supported fluid-filled canal (brac/iial canal) as its axis. This canal is again
supported by a loop-shaped calcareous ribbon (bracl,idium). From the brachidium-supponed
axis of lophophore springs a large number of slender parallel filaments which are sticky and
lined with numerous cilia. The movement of the cilia generates current in water. The inhalant
current enters the shell along the shell margin from two laterals sides inhalent currents. The
filaments are so arranged as to provide an effective net for trapping food particles while water
is passing through them. The filtered water is exhaled anteriorly as a single current exhalant
current. The caught food particles are allowed to pass downward in a mucus belt to a food
groove leading to mouth situated at the anterior body cavity.
10.5 CLASSWICATION
The universally accepted two subdivisions of the phylum Brachiopoda are based on the mode
of attachment of the two valves. In inarticulates, valves are joined only by muscles whereas in
articulates the two valves are articulated by teeth in the pedicle valve and corresponding sockets
in the brachia} valve. Funher subdivisions are based on (a) ontogenic development (b) variation
of pedicle opening (c) shell microstructure (d) internal morphologic features (e) ontogenic change
of brachidia and such other characters. ·
The classification, upto the level of order, given here mainly follows that of Williams and
Rowell in the 'Treatise on Invertebrate Palaeontology' (Moore er al., 1965).
Phylum : Brachiopoda
Class 1 : Inarticulata (Cam.-Rec.) Chitino-phosphatic shell, lacking teeth, valves attached by
muscles.
Order 1 : Lingulida (Cam.-Rec.) e.g. Lingula.
Order 2 : Acrotretida (Cam.-Rec.) e.g. Acrotreta.
Order 3 : Paterinida (Cam.-Ordo.) e.g. Paterina.
Order 4 : Obolellida (Cam.) e.g. Obollela.
Order 5 : Kutorginida (Cam.) e.g. Kutorgina.
Class 2 : Articulata (Cam.-Rec.) Shell calcareous, endopunctate; valves hinged by teeth and
sockets.
Order 1 : Orthida (Cam.-Rec.) e.g. Orthis.
Order 2 : Strophomenida (Ord.-Jur.) e.g. Strophomena.
Order 3 : Pe~tamerida (Cam.-Dev.) e.g. Pentamerus.
,,
,•
v.>
N
A:::1
-- ' ~- I
(b) SEMI-INFAUNAL (Clitambonites)

(c) PEDICALLY AITACHED (Magellania)
'I (a) ENCRUSTED/CEMENTED (Ricrhofenid)
,I
~
"',~::.?·.t=
... ,..·, .
:,\""'
.--...... ..,. ..
~~~~::.
~~:: ftt;-
~.:;
~ ~~ . .:;:1··
~,.
•·..,.._h .....
·11.~~ ...·--.., ...__.....
''.;9:aa:._
' ~:~:.
r.~
:?l'°?._;J
.1 ... '•'-.
;;·.:=:;·
....... ,
·.r.>~-
~';-":·;,:\
•' ,.~;,' }f (t) ENCRUSTED lCrania)
·!·:.... "'°ir:,
:,-;...._~.,
.•.. ··i
.: /}
(~: .: : .
·:.~..'!.-:· (e) SPINE AITACHED (Prod11ct11s)
(d) BURROWING (Ligu/ia)

~
(g) STRONGLY UNJPLICATED FORM -,,- (h) SEPARATION OF COARSE DETRITUS FROM ~
INHALENT CURRENT (Kochiproductus) ~
FIG. IO - 8 : LIFE HABITS OF SOME BRACHIOPODS ~
r---
a
Cl
• "-:::
ii :.. .., - ~~
•
BRACHIOPODA 133
Order 4 : Rhynchonellida (Ord.-Jur.) e.g. Rhynchonella.
Order 5 : Spiriferida (Ord.-Jur.) e.g. Spirifer.
Order 6 : Terebratulida (Dev.-Rec.) e.g. Terebratula.
10.6 ECOLOGY (Fig. l 0-8)

Ecologically brachiopods may be classified into four groups :
A. Pedically attached
Brachiopods are normally attached on the sea floor by means of their pedicles mainly on
dead shells of other organisms, rocky bottom, or rarely on soft mud. From fossils, it appears
that brachiopods had a tendency to grow in nests of clusters; their early member anchored
themselves to some dead shells of other organisms and later to their own dead shells. Growth
of these nests may come to a sudden end by being over~helmed by sediments which preserved
them in situ. These shells are often found in distorted forms due to over crowding of individuals.
Recent brachiopods have also ability to orient themselves in a preferred position with respect
to water current si()ce inward and outward movement of water within shell (inhalant and exhalant
wa.ter current) would be assisted if the animal keeps the axis perpendicular to current direction.
To achieve such a position, a pedically attached brachiopod may swing through a substantial
arc by the activity of adjustor muscle.
An unusual adaptation of the pedicle has been found in some recent brachiopods (Magadina).
They live in a high energy environment in drifting mobile shelf sand and their shells are partially
buried with their anterior edges only protruding out. If their shells are further buried by
sediments they push themselves upward to the surface using their finger like pedicle as elevator.
The pedicle of a brachiopod has a remarkable attachment strength and it can withstand even a
subtidal current. Functional pedicles are found in modern terebratulids, rhynchonellids and in
most of the fossil forms like spiriferids, pentamerids and strophomenids.
B. Encrusted
Some modern calcareous articulate forms cement their pedicle valves to rocks; the
commissure of the brachia) valve being shaped to fit it. Cementation is also known from some
fossil articulate richthofenids and strophomenids. A scar on the umbo of P-valve _of the latter
marks the place of attachment at juvenile stage, though in adult they were freelying .
C. Semi-infaunal / Semi-burrowing
Inarticulate lingulids live in partially buried condition in sediments. Fossil articulates like
productids and some strophomenids seem to have spent their adult life in a similar fashion with
sediment-cover on their concave brachia! valve. Productids were anchored by their strong spines
projected out from their P-valve surface. Only the flanged margin of the two valves would lie
above the general level of the surface. A few productids (Waag~11oconcha) had delicate spines
on their B-valve also, which prevented settled sediments from being winnowed away. In
Kochiproductus, the marginal flange internally forms a wide horizont~l extens_ion which
probably acted as a settling table for larger extraneous and inedible particles whale smalle~
particles were trapped inside by some taleolae extended like rows of stakes at the entrance of
the mantle cavity (Fig. 10-8h).
-
';.>)
~
lNARTlCUl.ATA ARTICULATA
-
R
T
-
C
~
l I ~
~
p
-
C
-
D
-s
-
0
~
LINGULA •
ACROTHELE
HESPERORTIIIS
LEPTAENA
~ RHYNCHONELLA
PENTAMERUS
SPIRIFER TEREBATULA i
!;
tr,
0
<'
FIG. 10 - 9 GEOLOGICAL RANGE AND RELATIVE ABUNDANCE OF SOME MAJOR GROUPS OF BRACHIOPODS 6
t-
o
C)
"-<:
'J
BRACHIOPODA
D. Free lying
Most brachiopods with closed pedicle openings (such as strophomenids, spiri fe rids, or nrthids)
must have lain on the sea-floor freely. Whereas, in case there is a small pedicl • ope ning, a
slender cord-like pedicle emerging from it could have a tethering function but woul d not huv ..
been supportive. Sediments settling in such cases around the commissure could have been easily
cleared by rapid clapping of the valves and in some cases such clapping might have all owed
a limited amount of swimming movement. These brachiopods possibly had functional pcdicl ·
at early stage of life.
E. Homemorphism
For adaptation to an almost similar mode of life (sessile benthic) brachiopod shells belonging
to related or unrelated stocks sometimes exhibit identical or near identica l external shape and
morphology. This phenomenon, called homeomorphism is the re sult of convergent evolution.
Homeomorphism in brachiopods may be observed within the groups appeared and pr ·sent in
same geological time, called isochro,wus homeomorphism or may be found within genera of
different ages called heterochronous homeomorphism. Well-known forms o f Pe rm o-
Carboniferous, all of which were formerly identified as the genus Producru.,· are later foun d to
be composed of several genera as appeared from their internal morphologic features and
'Productus' at present is considered as a 'form genus'. There are numerous such exal'nples
within brachiopods. The striking example of heterochronou s homeomorphy is the Middle
Ordovician genus Productorthis belonging to the order Orthida and the Carboniferous genus
Dityoclostus belonging to the order Strophomenida. Morphologically, both exhibit an orthid-
like outline and shape.
010.7 STRATIGRAPHIC 1-MPORTANCE

Brachiopods are the most dominant group of invertebrates in Palaeozonic. But most of them
failed to cross the Permo-Triassic boundary. Many of their Palaeozoic species exhibit a short
vertical range making them excellent zone fossils. Many of these zone fo ssils al so attained a
wide geographif ~ispersal through their meroplanktic larvae (although, sessile at adult stage).
These are successfully used as index fossils. Some important inarticul ate index fo ss il s of
Cambrian are : Obolella (L. Cam.), Acrothele (Mid. Cam.). Dicellomus (U. Cam .), Kutorgina
(L. Cam.), Rustella (L. Cam.) etc. Among the articulate brachiopods, Nanorthis (L. Ord.)
Dinorthis (Mid to Up Ord.), Pentamerus (Mid. Sil.), laptaena rhomboidali.\· (Ord.), Marginifera
(Perm.), Linoproductus (Up. Carb.), Neospirifer (Perm.), Syringothyris (Carb.) are the important
forms.

The earliest brachiopods are known from Lower Cambrian beds of many parts of th.e world
and on their earliest appearance they were quite diverse. They mostly be long to ina11iculates
with phosphatic shells. Well-known articulate groups ~a~e their first .appearance i~1 Ordovic'.an .
Inarticulates remained as a diverse group upto Ordov1c1an after which th ey declmed. Earliest
Ordovocian articulates were orthids and pentamerids which soon became diversified and the
former were probably the root of all other articulates. Other.group~ ap~ared were rhynchor~ell:ds
and strophom 'd I S ' lurian there was an abrupt reduction of 111art1culates and most ot them
em s. n 1 , . . h ll 'd
became extinct. There was also reduction of orthids and strophomends while rhync on~ 1 s
Palae(Geo)WP-l S
-- -·. ..
136 PALAEONTOLOGY
and spiriferids continued, to expund. Ti rebratulids appeared in the Silurian. During Devonian
spiriferids attain d their peak but end of Silurian saw a general fall of different groups with
extinction of several forms including pentamerids. From Carboniferous onwards there was again
a burst of evolution with diversification of productids. Permian was a time of great diversity of
strophomenids which in addition to productids included several other groups such as lyttonids
and richthofenids, which all colonized within reef-environment. A drastic fall of brachiopods
came after Permain and only a very few groups such as terebratulids, rhynchonel1ids,
thecospiriferids were able to cross the Palaeozoic-Mesozoic boundary. By the end of Jurassic
the last spirifierid also had gone.
Inarticulates seem to have changed a little since their decline at the end of Ordovician. A
single group, lingulid is persisting till the date without showing any distinctive change of their
morphology since their appearance (living fossil).
Chapter 11
BRYOZOA
11.1 GENERAL FEATURES

Phylum bryozoa includes a group of smaJl marine (except one or two fresh water forms)
colonial invertebrates, often forming reef-structures, ranging in age from Orodvician to Recent.
They produce hard laminated calcareous (rarely chitinous) colonies which are looked like small
shrubs or mosses (Gk.-bryon : moss), hence popularity are called 'sea mosses'. Sometimes
they are also cat1ed polyzoa as numerous of these animals together constitute the colony. There
are many records of bryozoan fossils in the geological column.
11.2 MORPHOLOGY (Fig. 11-1)

The entire colony is called zoarium which is composed of a number of individuals called
z.ooids. Zooicls are polymorphic and the most common forms are called autozooids and the less
common forms are known as heterozooids. Zooids are very small, basically cylindrical, fusiform,
or bottle-shaped, bearing number of tentacles. Zooids usually secrete tube or boxes of lime
partially encasing the soft parts. Each of them is called cystid or zooecium.
A colony of bryozoa grows from an initial zooid called ancestrula (Fig. I 1-1 b) by the growth
of stolons which are creeping tubular branched structures. This growth talces place by asexual
budding. However, a new colony is produced by sexual reproduction.
A matured cystid (Fig. 11- lc) shows a distal ovicell, a swollen, spherical structure in which
eggs develop into larvae. Directly behind the ovicell is the orifice or the opening of a zooid
which is key-hole shaped and often closed by an operculum. It is hinged at its rear end by a
narrow neck called cardelles. The distal and proximal end of operculum is called anter and
poster respectively. In cheilostomata groups zooids show two different organs named as
avicularia and vibracula. The latter bears a whip like bristle or setae projecting out from the
sessile basal chamber (Fig. 11-ld-f)
The more complex colony is exhibited by trepostomata group (Fig. 11-1 g) which are
commonly known as stony bryozoa. They form massive zoaria with elongated autozooecia
(initially thin walled but later become thick walled). Two such adjacent autozooecia may be
separated by a small thin-walled mesozooecial c~vity. Cavities of zooecium is partitioned by
closely spaced diaphragms or septa.
11.3 ECOLOGY
Bryozoans are abundant throughout ocean ranging from shore line to abyssal zone with
diverse number. Ecologically, their growth is controlled by various conditions _such as :
(I) Temperature : Most of the forms are restricted in warm water.
(ii) Wave action : They avoid a zone of strong current and wave act.ion that can damage
their colony.
137
,i J ..,. ~ -- '
'. ,. . , -~ ....
138
Zooid
Stolon
(a) A COLONY
Vibracula
(b) ·YOUNG ZOARIUM GROWING

Frontal Wall
FROM ANCESTRULA
(c) ZOARIUM OF A CHEILOSTOMATA
(operculum missing)
~:__-+ Operadum
CradcUe Pos\Cr
(d) AVICULARIUM (enlarged) '-11..,_~~~-Avicularium
(e) AN OVICELL (enlarged)
) ~kw .woe , ,I
Ca1- i
Seta
Septum
(f) VIBRACULUM (mlargedj
(g) A TREPOSTOME COLONY IN SECTION
FIG. 11 - 1 : MORPHOLOGY OF BRYOZOANS
·• ·.---·.
BRYOZOA 139
(iii) Hard substrate : They prefer hard bottom on which larvae settle and hence are common
in shelf zone.
(iv) Salinity : Mostly they are stenohaline except a few which are euryhaline.
11.4 GEOLOGICAL HISTORY

The geological history of bryozoa is known rather completely. They appeared at first in
Ordovician and the most important groups trepostoma and cryptostoma reached their climax at
Upper Palaeozoic. Cyclostoma, though appeared in Palaeozoic, were reduced in number but it
expanded vastly with the extinction of former two groups after Permian. They continued to be
dominant upto Upper Cretaceous when cheilostomata arose to become the dominant forms of
bryozoa in Cenozoic.
Chapter 12
MOLLUSCA
12.1 INTRODUCTION
Phylum mollusca (Latin- molluscus : Soft) is the second diverse group of invertebrates after
arthropoda. This includes snails, mussels, cuttle fishes, slugs, cockles, _limpets, octopus and a
host of other forms which arc so different in appearance that superficially they appear to be
unrelated. They, like their external forms also exhibit a wide range of mode of life; some ~re
marine, benthic sessile or mobile, some are nektic; many are freshwater and others terrestnal.
All types of food habits such as carnivorous, vegetarians, filter-feeders, deposit feeders are fo~nd
in them. Size of these invertebrates also exhibits a wide range from few millimeter to the giant
squid of 16m or more. In spit~ of this great variety of forms, size and mode of life, the animals
exhibit a common anatomical blue print and that is why they can be easily placed within a
single phylum. All of them belong to triploblastic coelomate grade with oligomerous body plan.
The fundamental organization or basic features of all molluscs can be visualized with
reference to the hypothetical form often described as 'Archimollusc' (Fig. 12-1 a). However,
no living mollusc is known to possess such a model, but such a form might have been present
during Cambrian-Precambrian times. This primitive root-mollusc has a cap-like shell of calcium
carbonate, secreted by a fleshy inside layer, the mantle which covers a soft, apparently
unsegmented body of the animal. There is an empty cavity towards posterior between the soft
animal and the mantle called mantle cavity in which lie gills and the anus. Mantle cavity is
produced by infolding of mantle layer. In front and above the mantle cavity lies the viseral
mass which includes digestive canal (including stomach), heart, kidney, gonad etc. Mouth and
anus lie at anterior and posterior end of the digestive tract. Mouth possesses a rasp-like tongue
called radula bearing a few sharp inward pointing teeth. In some cases, tentacles are found
surrounding the mouth. In the ventral side occurs a fleshy foot upon which the animal rests
~nd moves. Not all, but many molluscs have_ tubular siphonal system inside used to carry
inhalant and exhalant current of water and this has contributed greatly to the success of the
animal in the marine habitat.
12.2 SUBDIVISIONS
An outline of subdivisions of the phylum are as follows :
Class-I : Monoplacophora (Cam.-Rec.) : Primitive marine molluscs, univalved· a circular
~entral mantle cav~ty in which lie the gills; they are the only mollus~ with true
mternal segmentation e.g. Neopiliha.
Class-2 : Amphi?eura (Cam.-Rec.) : This includes two groups : aplacophorans, a group
of ~arme small ~orm-like molluscs that lack shell and h~'lce any record of
fossils; the other mcludes the marine polyplacophorans having a symmetrical
shell, made up of seven to eight calcareous plates e.g. Chiron.
140
1• . ·1 •
; ~ ; , .;l -
MOUUSCA
141
Sh---
S--- Sh
Dc-----l~~~'W'fft1r.r4illt~ Mc
Si
R--~ A -.,....__sc
M .--...;::a'li: lllH----Dc
ILMi-.--R
F
G-----..J
b. AMPHINEURA
~----A H
M
T--M··~,lll"f""--F
re.SCAPHOPODA Gd
y d. CEPHALOPODA
S---7/llE_,~~~~
Sh---_,,.R'
Cg~~~~ite~~~~~
F M-~~~~;;;::.:J:===~~~~~~
R---,~'-3 . ·-Al....--F
c. PELECYPODA NI H
H De
T
M
R
f. MONOPLACOPHORA g. GASTROPODA
FIG. 12-1 BASIC STRUCTURAL PLANS OF DIFFERENT LIVING MOLLUSC CLASSES

WITH REFERENCE TO HYPOTHETICAL ARCHIMOLLUSC
A : Anus, Cg : Cerebral ganglion, De : Digestive canal, F : Foot, G : Gill, Gd : Gonad, H : Hearl.
K : Kidney, M : Mouth, Mc : Mantle cavity, Ms : Muscle scar, NI : Neural loop, R : Radula, S : Stomach.
Sh : Shell , Si : Siphon, T : Tentacle.
PALAEONTOLocy
142
Hinge AIU
Umbo (Protoconch)
--lnneBide
~1 ...
(CoocaVC)
(a) Rapid Growth Of Shell

Rapid Growth Of In Widda l)irec:tion (W > L) UncQual Orowth Of Two
Shell In Lcngtb Sides Of A Cone Of Two Coiled Valves
OiRction (L > W) Pdeypod Making The Valve Of A Pdccypod
UNCOlLED CONE Making The Shell
rl3 Axis of, Coiling Axis of Coiling
I
I
I
I
I
I
CcphaJopod With Rapid Rate (d) Cephalopod With Slow Rate

Of Increase Of Height' Of lnCl'Ctie Of Height'
Producing rnflated Shen Producing Discoidal Shell
rll .
!.J._ = !J_ = r JO
= 1.5
r2 r6 r 12
Coiling Of Cqmq,od Shell Rcprescnting An
F4ppiengnw Spire Sbowin8 COOltlllt Ratio Of
hs .Rati.i After Each Revolution
(c.onicll Sbdl Planispcrllly Coiled In Cq,halopoda)
; Axis or Coiling
{f) Gamopod Shell With

Decreasing Rate Of
Whorl Translation
GIStlopod Shell With
(e) Rapid Increase or Rale
Of Whort Translation (T)
Conilpinlly CoiW G1llbOpOd Shell Sbowin
Cbmp Of 'Jbttic Parametas (D,W And T) I
fg 10 mm
W . - .. - - - 2 : D •aha:Smm;cf • IO mm ; T=ac
bd S mm
FIG. 12 • 2 : GROWTH AND RELATED SHELL MORPHOLOGY OF MOLLUSCA
MOLLUSCA 143
Class-3 : Scap~opoda (Ord.-Rec.) : They are marine with tapering and curving shells
openmg at both ends. They feed on small organisms using their specially adapted
tentacles; no gill. e.g. Dentalium.
Class-4 : Rostroconchia (Cam.-Perm.) : Bivalved, but dorsal commissure is lacking.
Juvenile shell is univalved and coiled e.g. Macroscenella.
Class-5 : Bivalvia (Lamellibranchia or Pelecypoda) (Ord.-Rec.) : Bivalved shells with no
definite head; valves dorsally hinged; opened and closed by muscles; a distinct
muscular foot; siphonal system may be present e.g. Clams.
Class-6 : Gastropoda (Cam.-Rec.) : Marine, fresh-water or terrestrial; shell with a single
valve, mostly coiled helically, internal organs twisted by a 180° torsion so that
the mantle cavity faces anteriorly; well defined mouth and foot. e.g. Snails of
all kinds.
Class-7 : Cephalopoda (Cam.-Rec.) : Advanced molluscs having chambered shells; coiled
or uncoiled; chambers linked by a siphuncle giving buoyancy in marine nektic
life; well defined head with number of tentacles and several sense organs e.g.
Nautilus, Octopus, etc.
12.3 SOME ASPECTS OF GROWTH RELATED TO SHELL MORPHOLOGY AND

SHELL SHAPE (Fig. 12-2)
Shell is the main structure in molluscan body normally fossilized and thus study of shell-
growth would be significant to understand the shell morphology and its shape.
In mollusca and brachiopoda growth of the· shell takes places through accretion of materials
around the shell aperture. The simplest form or shape of the shell resulting from such growths
would be a cone, the initial part of which is called protoconch. (Fig. 12.2a)
If the growth is rapid and uniform on all sides, the diameter of this conical shell would
expand in a short distance (length < width). Conversely, if the addition of shell material does
not cause any sufficient increase of diameter, the cone would be long, tubular type (longer than
width) (Fig. 12-2a). If the rate of forward growth of shell be slightly/highly greater on one
side than on the other, the shell would be coiled giving a spiral form. This coiling may take
place in one plane around the axis (planispiral) or in different pl'!nes around an axis (helicoid
or conispiral) (Fig. l 2.2c-f). It is interesting to note that the bivalved brachiopod and pelecypod
shell, in fact exhibit a type of coiled shell (Fig. 12.2b) where each valve represents a cone and
its open side being the aperture of the cone. Here the outer side of the cone grows much rapidly ·
than the inner side, thus giving a convex and curved external surface and a small concave inner
side (forming the cardinal area of the valve); umbo of each valve being the protoconch-part of
the shell. Here the two cones, representing the two valves, are joined along their apertural
margin. Raup (1966) used a computer-based graphical method to produce various kinds of
hypothetical coiled cones. Theoretically, he indicated the presence of a large number of shapes
of coiled shells based on four parameters. (Fig. I 2-2e).
(a) Shape of the cone/tube in cross section, otherwise known as shape of generating curve
(or apertural outline).
(b) Rate of increase of apertural diameter (W) after each revolution. This is the ratio of
the two diameters after one complete revolution.
Palae(Geo)WP-19
PALAEONTOLOG y
144
. d with respect to axis. It is the horizontal
(c)_ Position and orientation of generatmg curve t.
. f T ng after each revo Iu ion.
distance of aperture (D) from axis o cm •
(d) Rate of whorl translation (T) along the axis. It is the measure of vertical fall of aperture
after each revolution in a conispirally coiled shell.
ds) shells are actually rolled up cones
In most coiled molluscas (cephalopods and gast~opo four arameters metioned above may
whic~ grow at apertural end only. In _a cephalopod shell, rith;nic or equiangular spire. The
remain almost constant and hence 1t represents a longa . E h .
· · It 1'ts shape as 1t grows. ac increment
peculiarity of these spiral shells 1s that it does not a er . d h d .
is identical to its predecessor (Fig. l 2-2c). In nature however, many cot 1e s e II s may eviate
from this ideal mathematical form.
·
However the four parameters mentioned ear1·1er may vary in different groups
. of
. coiled
, , forms.
. .
' · · d
Thus gastropods are usually hehcally cmled an ten o av d t h e low 'W' with variable
. , , . T g1vmg
.
long or short spiral cones (Fig. I 2-2f). In a bivalve 'T' is very l~w _while_ W ts very high,
whereas in a brachiopod T = 0. Cephlopod shells are normally plamsp1ral with ~nd T = 0, thus
producing highly globular (high W) to highly discoidal sheJls (very low W) (Fig. I 2-2d).
12.4 ORIGIN AND PHYLOGENY OF MOLLUSCA

The most common approach is that the origin of mollusc tooks place prior to Cambrian from
some worm-like creatures with a hard shells. The Proterozoic origin of the mollusc may be
confirmed as most of the groups of mollusc appeared by Lower-Middle Cambrian. The simple
morphology of hypothetical ancestral mollusc (called Archimollusc) is reconstructed which has
probably a close resemblance to modern chitons (amphineura) and/or Neopilina, a modern
monoplacophoran. It is assumed that all other groups of mollusc were diverged from this ancestral
form and the differenciation is bound up with their mode of nutrition (Projeta, 1980). In fact,
their structural differenciation is closely related to their mode of feeding. Monoplaco-phorans,
amphineuras and most gastropods are slow-moving animals with a radula, a belt of serial teeth
near mouth. These make them herbivores, carnivores or scavangers and necessarily they have head
and sense organ. Bivalves are mostly suspension feeders or deposit feeders. They have no head,
no radula, and have poor movement capacity. Cephalopods are active hunters with strong and
large head, sense organ, radula and additional jaw for crushing of food materials.
Regrading the origin and nature of ancestral mollusc there is another hypothesis. It is found
that molluscs and annelids h~ve some embryological similarities which may indicate their origin
from a common ancestor like a turbellarian flat worm. From such an ancestor annelids
developed_with segmentation of body while molluscs remain unsegmented throughout like their
ancestor.
The ~ise of molluscs fr?m turbella~ian-li~e ancestor was the result of a single evolutionary
accomphsh~ent; the. se~~et10n of muco1~ cuticle over their dorsal surface. This was subsequently
developed mto a pnm1ttve mantle. Spicules of aragonite later became embedded within the
cuticle forming the outer layer o~ the shell (periostracum). Ostracum was secreted later below
the periostracum as layered calcium carbonate. From the presence of spicular turbellarians, it
is reasonable to say that ancestral mollusc was similarly a spicular worm-like animal with
cuticle. In this respect, the ancestral mollusc was very similar to the living worm-like
f - ~ -r · . ....
"') ' ·':--: ; .
• .a ~
. '
!,{OUUSCA
145
iacopborans, as the latter possess various moll .
~ nn.) uscan anatomic features but without a sh II
(hke a wo · . e
Aplacophorans have the most simple mo holo
bUt they have no fossil record and it is oft: deb!ie;ndhanthatomhy ~m~ng the living molluscs
. w e er t e1r simple morphology is a
Primitive· feature or second anly derived. If they are accepted . .. & • • :
·· II t . . as a pnm1t1ve 1orms exhibiting
many ~nnutive ~o use~ eatures hke radula, restricted coelom, mantle cavity and gill there
is no ~1fficulty 10 acceptmg them as the earliest forms having a spiculose cuticle prior 'to the
formation of first shelled-mollusc.
Polyplacophorams and monoplacophorans both evolved from such a spicular ancestor

(aplacopho~). ~olyplacopho~s developed eight-valved shells secreted under spicular cuticle.
This fonn 1s believed to be deviated from the rest of the molluscs and never shows three fold
division of mantle margin like other molluscs. Monoplacophorans were also derived directly
from aplacophoran, having a dorsal univalved shell. The remaining classes of mollusc are all
e,rolve.d from this monoplacophoran ancestry. Gastropods probably evolved from
monoplacophorans with their dorsally placed shells helically coiled and exhibiting a torsion of
their viseral parts. There are some living monoplacophorans which can rotate their helical shell
90°. with respect to their head-foot mass with spire facing away from head causing torson of
their soft parts. Cephalopods probably derived from some dorsoventrally elongated monoplaco-
pborans; some of which are also known to possess primitive septa. Such forms often show shell
curvature away from the head and a tube or snorkel on the dorsal side which might have been
transformed later into the siphuncle of a cephalopod.
Pelecypods probably evolved in early Cambrian from rostrochoncha (an extinct Palaeozoic
group). The latter, however were descendant from some groups of monoplac~ph~rans by
differencial calcification of the dorsal margin. This could have produced the elastic ligaments
and development of adductor muscle of a pelecypod. The broad phylogenic pattern of molluscs
may be shown below (Projecta & Tulane, 1980).
Geological ranges of three major groups of mollusc are also shown in fig. 12.3.
Pelee~ /opoda Gastr~ /lopoda

Rostroconcha (Ext.) · ~
~~.
(~~I::::~,'' /acophora
', Aplacoph<?ra '
', l
Turbellarian-Jike worm
(Ancestral form)
- . - _,,,.,, ...
.,· ./
146 PALAEONTOLOGY
:r. Geological
~
J .i :;
I•
"- "' Time C
0~
j :I u
c:.i
JI
'=
uU
·....; E
E ~
tz 0-=
:E
Name
of Orders ~
·I
i J ~ l .... u
'....ii :.i
;;
::,
0
..,
'.,
::ii::
Nuculoida ,...
~
Solcmyoida
•
~ida ...,
Arcoida ,'
Mytiloida ..
Plcroida ,...
t
l
Modiomorpboida
Unionoida
.'
.'
...,
Trigonoida
Vencroida ...
'
Myoida ...,
Hippuritoida
~
Pholadomyoida ...,
Arcbacopstropod
',
Mesogastropoda ...
,
'
I
~
Ncopstropoda
C>pisthol,rmdua
(Subclass)
....,
...,
Pulmonata ...,
(Subclass)
Orthoccratoidea ....
7
Endoceratoidea ,....
Actinoceratoidea ...
' ...
J Bactritoidea
Nautiloidea
Ammonoidea
-,
...,
'r
Celeoidea ....,
FIG. 12- 3 GEOLOGICAL RANGE OF MAJOR GROUPS (orders) OF PELECYPOD

GASTROPOD AND CEPHALOPOD
-·-- ' ·- · . · .: . 9.~-
~~
MOUUSCA
147
12.5 EFFECT OF PREDATION ON LIFE-HABIT OF MOLLUSCS
The soft ~es~y and muscula.r body of most of the mollucas is taken as food by many
predators. This danger of predation has had a great impact upon the types of adaption taken
by these animals. Different kinds of molluscs have responded to this problem in different
manners. Their effect can be assessed even iR fossil specimens. (Vermeij, 1983).
Cephalopods depends upon escape and avoidance. For this, they have developed quick
movement by jet-propulsion system. They have also developed a poisonous liquid within an
ink-sac which when disturbed is injected on ememy's body. These have been proved successful
devices for their survival. Gastropods possessing comparatively thick shells, narrow apertures
have ability of a quick and complete retreat of their soft parts within the shells. But bivalves
have intrinsic limitations imposed upon them by their shell construction which render them
more vulnerable than gastropods. For this, they exhibit a variable types of sessile forms having
very thick shells. Burrowing is another obvious response. Some develop active swimming mode
of life depending upon avoidance. Sheltering within sea-grass communities, invasion in deep
water, high rate of fertility are some other adaptations commonly found in them. All these
have been proved successful strategies, as evident from the present diversity of molluscs.
Chapter 13
PELECYPODA (BIVALVIA)

The name 'pelecypoda' is derived from an anteriorly placed mascular ha~chet-s~aped f~o!
(Gk.-pelekys : hatchet; pous/podas : foot) of the animal. It is also known as lamellibranchia
for its possession of internal lamellar gills (Lat.-lamella : leaf; branchia : gill). However, the
original name 'bivalvia' of this group was given by Linnaeus in 1758.
The animal possesses a calcerous exoskeleton called shell covering its soft pa~s. Th.is shell
consists of a pair of valves which are generally identical, equal-sized but mostly meqmlateral.
Valves are placed laterally on either side of the animal and referred to as right valve and left
valve (Fig. 13-lb). The two valves are mostly articulated along their dorsal margin usually by
means of several teeth and sockets borne by each valve. As the two valves are identical, a
bilateral symmetry is shown by the shell where the symmetry plane is passing in between the
two identical valves parallel to dorsal-ventral margin of the shell. There are however, a few
bivalve shells showing unequal valves thus lack any symmetry (Fig.13-1 h). Animals also exhibit
a wide range of life habits. Most] , the are marine and benth· . Within them there are both
sessile an vagile groups. ~an o(.tbem-,are...bun:ower~ Qf s..and~(infaunal). There are also fresl't
water bivalves ILving_ia.p.~ · · - ---- - - · - - -
The pelecypod shell is composed of three layers, the outer periostracum layer of hormy
conchioline (chitin-like substance), a middle prismatic layer of calcium carbonate (calcite or
aragonites) called ostracum, and an inner hypostracum layer composed of closely spaced
alternating lamellae of conchioline and calcite. (Fig. 13-1 a)
The shape of the shell (Fig. 13-2a) is defined by the outline of valve which shows a wide
~ at~on ran~mg fr~pi ctr~_ular subcircuJar e li12tical oval _triangu ar, rectangular e tc-:-Tateral
shell margin may be rounded/protruded or truncated. The shell may sfiow a orwar O 1quity
(prosocline), backward obliquity (opisthocline) or may be neutral (acline or orthocline) (Fig.
13-2b).
13.2 EXTERNAL FEATURES (Fig. 13.1-3)

Umbo (Fig. 13-3b) Elevated or protruded part the dorsal margin of each valve· may be
erect, suberect or incurved. '
Beak (Fig. 13-3a) Sharp and pointed end of umbo marking the initial shell-growth. Beak
may face three ways.
(i) Prosogyral : Beak pointing anterior direction.
(ii) Opisthogyral : Beak directing towards posterior.
(iii) Orthogyral : Beak points neutrally i.e., neither anteriorly nor
posteriorly directed.
148
...... ..
149
p£LECYP0DA (B/VALVIA)
Left Valve--111i1.-.
----........a igamcnt
()stnc,uaa----,. .
Pcriostracum
PcriostraCUlll,-....,., ~
~
----E=:..
.
Hyposb'ICUffl
Gill--...
(a) SHELL-WALL
MICROSTRUCTURE
Free Line Madcing
Mantle Mantle Attachment
(Pallial Linc)
(b) A SECTION ACROSS Hinge Line
PLANE OF SYMMETRY ----Ligamcntal Arca
---Umbo
I)orSal
Posterior· Adductor ""'!!!!!!!!!~lllllii-=::--.t Bc:alt
Muscle Scar --~~~ HingcTccth
&Socket
C
Posterior --t
Anterior Adductor
Growth _ _..
Linc Ventral
Pallial Sinus Muscle Scar
- - - Pallial Line ',.
..)
(c) EXTERNAL VIEW
Lunule - - - '
Hinge Linc
Anterior
Umbo--• Space between valves
Escutcheon
Posterior view
(g) GAPING ALONG

(0 ALATE & BYSSATE FROM
POSTERIOR MARGIN (h) INEQUIVALVED
SHELL
FIG. 13 - 1 : BASIC MORPHOLOGICAL FEATIJRES AND ORIENTATION OF BIVALVIA

SHELL
'
PALAEONTOLOGY
so
CirCWI'
('' .::"':' "" ·:::;:;:.-

Mytilifonll
c:::::J R~
Etongl&Cd Or projc=d
(a) VALVE OUTLINE

Posterior
Prosoclin~
<>pisthoclioc
Ac\ine
(b) OBLlQUITY Of VALVE
Carina Tubere\es
Growth Lines Tut,cn:ulate

And Carinate
• Plicacd
. Spinale And Cos1ate
Divariate
Anterior ~ Posterior
0~ Roundecl Angular
Truncaacd
(d) NATURE OF ANTERIOR AND POSTERIOR MARGIN

sl4£LL
FIG. 13 - 2 : GENEllAL MORPHOLOOICAL FBA11JltES OF PELECVP0D (DIVALVIA)
PELECYPODA (BIVALVIA) 151
Hinge line Dorsal margin (umbonal side) along which the two valves are
(Fig. 13-3c) articulated. Hinge line may be straight or curved; long or short.
Hinge area or Plane or concave areas between hinge line and beak of each valve;
Cardinal area usually smooth. In some forms cardinal area is divisible into two
(Fig. I 3-3d) portions.
(i) Lunule : Small heart-shaped depressed portion of cardinal area
anterior to beak.
(ii) Escutcheon : Elongated, depressed portion of cardinal area
posterior to beak.
Hinge area may be very poorly developed in many forms.
Auricle (Fig. 13-1 f) Forward and/or backward ear-like projections of each valve along
hinge, found in pectinid pelecypods; shell possessing auricles are called
auriculate or alate. Two sides of the ear may be equal or unequal.
Byssal notch Indentation below anterior auricle of pectinid shells for attachment of
(Fig. 13-1 f) byssus (some thread-like processes derived from foot, used to attach
the shell to substratum).
Ligament Elastic tissue on cardinal area attaching the two valves along hinge
(Fig. 13-3e) line serving to open the valves. Structurally, ligame·nt may be of three
types : alivincular (consisting of a single coral-like strands);
multivincular (consisting of a bundle of strands) and perivincular
(consisting of a hemicylindrical band). As regards the position with
respect to beak, ligament may be prosodetic (anterior to beak),
opisthodetic (posterior to beak) and amphidetic (on either side of
beak).
Ligamental pits Cjrcular/linear depressions on hinge area marking the lines/zones of
or grooves attachment of ligaments, usually posterior to beak but may also be
central. Inverted V-shaped ligamental grooves are found in Arca.
Line of commissure Junction of two valves along ventral side, which may be smooth,
crenulated or plicated.
Gape (Fig. 13-1 g) Permanent open space rarely found along anterior and or posterior
margin for the passage of foot and/or siphon e.g. Schizotherus.
13.3 INTERNAL FEATURES (Fig. 13-1 d)

Hinge plate A plate projected downward from . the hinge line bearing teeth and
sockets.
Teeth and Sockets Projected ridges and corresponding grooves on the hinge plate; tooth
(Fig. 13-1 d) of one valve fits into the socket of the opposite valve.
Cardinal teeth Teeth on hinge plate projecting from below the umbonal part whose
Jong axes are nearly perpendicular or oblique to the hinge line.
Pa!ae(Geo)WP-20
PALAEONTOLOGY
\52
--,. Posterior
Opisthogyral
Qr1hogyral
(a) BEAK
~
t1\-~~
Sidcvicw Upright Slightly tncurvcd
StronslY Incurved
Side View
(b) UMBO
H.L
Suaight
Curved
Undivided Hinge Arca
Ligamcntal Groove Resilium Ligament Slit-Like
~ Rcsilifer
Chonddrophorc
~Resilila
I~ ~ Buttress (viii)
e·
LigtlDClll Anterior
A)ivincul• ' Ligament Posterior .
Anterior ~ .... iri ~
PrOIOdctic
-:<9!:,
Opis1hodctic
l Posterior
Amphidctii.:
(c) LIGAMENT, Rl:SILIH::R AND ASSOCIATED STUCTURES
FIG. t 3 - 3 : MORPHOLOGICAL FEATIJRES OF PELECYPOD (BIV ALVIA) SHELL
PELECYPODA (BIVALVIA) 153
Lateral teeth Teeth anteriorly and or posteriorly projected from hinge plate whose
long axes are nearly parallel to hinge line.
Teeth pattern Teeth pattern of bivalvia is found quite variable and the fossil bivalves
(Fig. l 3-4a, b) are sometimes classified on their nature of teeth. Variable types of teeth
found in them are as follows.
(i) Taxodont : Hinge plate characterised by numerous identical
serreted teeth and sockets e.g. Arca, Nucitla.
(ii) Schizodont : Sharply diverging teeth (cardinal or lateral) often
bifurcating e.g. Trigonia, Unio.
(iii) Isodont : Two large subequal hinge-teeth and sockets
cardinal in position e.g. Spondylus.
(iv) Desmodont : Teeth highly reduced or even absent but acces-
sary structures present for articulation of valves
e.g. Mya.
(v) Dysodont : Two identical lateral teeth e.g. Pecten .
. (vi) Pachydont : Dentition in sessile rudistid bivalves where teeth
are large heavy and blunt e.g. Hippurites.
( vii) Edentulus/
Palaeconcha : Bivalves lacking teeth of any kind e.g. Lima.
(viii) Heterodont
(Teleodont) : Most Tertiary and Recent bivalves have advanced
type of teeth and sockets composed of both cardinal
' or lateral, although, both may not be equally
prominent and their modification of some kinds
may be present. Typical heterodont teeth may be
of two types: Cyrenoid with three cardinal in each
valve and more lateral teeth in left valve; Lucinoid,
with two cardinal teeth in each valve and more
lateral teeth in right valve (Fig. 13-4b). Usually
there are 2/3 cardinals and 1/2 lateral teeth on the
hinge plate. These teeth are often symbolized
according to the system devised by Bernard and
Munier-Chalmas. Odd number are used for right
valve and even numbers for left valve.
Internal ligamental features (Fig. 13-3e) :
Resilium : Portion of ligament below the hinge line on
hinge plate or below it; compressed by hinge
plate when valves are closed.
PAL4.EOr
154
So)ieo-• (Tri&O*)
~
Pac*,._.
(Hippw m)
"V*'?
O'jsudoala (.Pa:tal)
D: I .... (.W,..)
(a) PRINCrPAL TYPES OF DENTITION IN PELE:CYPOD SHELL
(b) NUMBERING Of KETERODONT TEETH AS DEVICED BY BERARD A D M lER-CHALMA , RO\ \A:-,l :\X .
ARABIAN NUMERALS ARE USED R)R LATERAL AND CARDINAL TEETH RESPECTIVELY. D NLI MFRALS AR
USED FOR TEETH OF RIGHT VALVE AND EVEN NUMERALS FOR LEFT AL\ E. · • DISlGNAT ..\NTc.Rl( R.
·b' RJR POSTERIOR CARDrNAL TEETH (VARIX). ·A' STANDS FOR ANTERIOR AN ·p' FOR POSTER! R.
FIG. 13 • 4 : TYPES OF DENTITION IN PELECYPOD SHELLS
• 1 • • , .
·-•=· :..• , .. . 't_J. :--~ - · A..
PELECYPODA ( BIVALVIA) 155
Resilifer : A pit or depressed area on hinge plate bearing

resilium. e.g. Pecten.
Chondrophore : A prominent spoon-shaped structure sometimes .
r .
present hanging below the hinge plate holding
the resilium e.g. Mya. . .
Myophore : Plate or rod-like structure below hinge plate for
muscle attachment e.g. Pho/as.
Buttress : Two ridge/rod-like projections from below the
hinge, supporting part of hinge.
Muscle Scar Depressed circular/subcircular areas placed anteriorly and posteriorly
(Fig. l 3-5a) inside each valve, marking attachment of adductor muscles (serving
to close the valve). There may be two types of muscle scar.
Dimayarian : Shell with two adductor muscle scars in each
valve placed anteriorly and posteriorly. If the two
muscle scars are of equal-sized the shell is called
isomayarian, if unequal, it is called
•
a11isomayarian (posterior one generally larger).
Monomayarian : Shell with only one adductor muscle scar
(posterior one).
Valve attachment A bivalved pelecypod is living with its valves opened along the ventral
(Fig. 13-5b) margin and hinged or attached along its dorsal margin in normal
condition. It can close its two valves along the ventral margin as
and when necessary by contraction of the two adductor muscles
transversely running across the two valves. As there would be no
function of muscles after the death of the animal, the two valves remain
opened along the ventral margin and soon become disarticulated along
the dorsal margin and are usually preserved separately.
Pallial line A linear depression inside each valve near the ventral margin and
nearly parallel to it marking the inner margin of thickened mantle edge.
J
Pallial sinus Inward deflection/in~entation of posterior part of pallial line defining
the space for retracted posteriorly placed siphon. (Fig. 13-1 d).
13.4 EXTERNAL SURFACE ORNAMENTATION (Fig. 13-2c)

Growth lines More or less uniform concentric lines parallel to the shell margin
marking successive growth stages starting from umbonal protoconch.
Growth line may be very much prominent in some bivalves e.g.
Lucina.
Costae Radial lines diverging from umbo towards the ventral margin; these
lines may be faint (e.g. Cyrena) or very coarse (e.g. Pecten); presence
of coarse costae result crenulation of ventral margin.
PALAEONTOLOGY
156
Ms
~~~r---Shcll---61'
AdductOr
Muscle
Anisomayarian
Monomayari111
(a) TYPE OF MUSCLE SCARS
(Ms: Muscle St!ar)
(b) ACTION AND EFFECT OF MUSCLES
FIG. 13 - 5 : MUSCLE SCAR AND MECHANISM OF OPENING AND CLOSING THE VALVES
BY ACTION OF ADDUCTOR MUSCLE AND LIGAMENT SHOWN IN CROSS
SECTIONS
BRACHIOPOD SHELL PELECYPOD SHELL
Posterior Anterior
C Ventral
SIDE VIEW
a-b & e-f: Length
c-d Thickness Of Shell g-h Hcight:ABCD Symmctry Plan
Posterior
IC
I
v' :f
Dorsal View
Dorsal View
a-b & e-f length '"" Trace Of symmeotry Plane
c-d; Maximwn Width
Ix
Ventral View
Anterior View
x-y=TraccOf~--
-:1 ......, ... u,
Plane
FIG. 13 - 6
:~~~·sC:~A~ON AND DIMENSIONS OF BRACJDOPOD AND
PELECYPODA ( BIVALVIA) 157
Multicostatc Surface with costae which increase in number through bifurcation
towards ventral margin e.g. Volsella .
Plicate Shell radially folded to form coarse ridges and furrows. e.g. Pecten.
Cancellate Shell surface marked by prominently growing growth lines and costae
intersecting each other. e.g. Chione.
Di variate Shell surface marked by two sets of costae meeting along midway at
an angle, e.g. Acila.
Carinate Shell often showing an angular ridge extending outward from beak
in the posterior side e.g. Trigonia.
Foliaceous Shell surface with very irregularly space lamellae marking growth lines
e.g. Ostrea.
Rostrate A tongue like posterior projection of shell e.g. Cardiomya.
Truncated Lateral shell margin abruptly ending.
Spines Sharp tubular structures emerging at points of intersections of growth
lines and costae in some bivalves e.g. Spondylus, Plicatula.
13.S DIMENSIONS AND ORIENTATION (Fig. 13- lc, e)

Dorsal Direction of shell where two valves are joined/articulated towards
umbonal side.
Ventral Opposite side of dorsal; direction towards which the valves can be
opened and closed.
Anterior Direction of shell towards the mouth.
Posterior Direction pointing anal-opening or siphon.
Length Maximum distance between anterior and posterior.
HeighUWidth Maximum distance between ventral and dorsal margin.
For recognition of right and left valve in fossil pelecypods one has to recognise at first
anterior and posterior direction of the valve. After this, the valve has to be place in such a
manner that anterior would be away from observer and posterior toward the observer, ventral
side downward and dorsal side upward. In such a position the external surface of the valve, if
it would lie on the left side of the observer the valve would be left valve, in reverse case it is
right valve.
To recognise anterior and posterior direction of a bivalved-shell we can look for the following
positive features :
(i) Byssal notch in a pectinicl shell lies anteriorly.
(ii) Pallial sinus, if present, is always found toward the posterior of the valve.
(iii) If the shell, is monomayarian, the single muscle scar is the posterior on~ and its position
may indicate the posterior direction of the valve.
158 PALAEONTOLOGY
TABLE-10
Distinction between Brachiopod and Pelecypod shell (Fig. 13-6)
Brachiopods Pelecypods
I. Valves unequal, generally larger valve Valves generally equal sized, aligned on !wo
occurs on the ventnd side called ventral lateral sides of the body, hence called nght
or pedicle valves, smaller valve on dorsal valve and left valve.
side called dorsal or brachia( valve.
2. Valves equilateral. Valves generally inequilateral.
3. Bilateral sy.mmetry plane passing . across Bilateral symmetry plane passing in between
the two valves. the two valves.
4. Two valves articulated along posterior Two valves articulated along dorsal margin.
margin.
5. Inarticulates have no teeth; in articulate, Teeth present in most of the cases and each
teeth occur in pedicle valve and valve bears both teeth and sockets.
corresponding sockets occur in brachia(
valve.
6. No ligament; muscle controls the opening Valves are opened by ligaments/resilium and
and closing of the valves. closed by adductor muscles.
7. Shell wall composed of three layer. Shell wall mostly composed of more than
three layers.
In case the inequilateral shell is devoid of all the above features, one can apply a simple
method. The valve may be held with dorsal margin upward. Then, an imaginary vertical line
should be drawn from umbo to the ventral margin, dividing the valve into two unequal halves;
the direction of smaller half would be the anterior. It should be remembered that this is not a
positive criterion as there are many shells with anterior half larger than posterior.
13.6 CLASSIFICATION
Pelecypods includes a large group of molluscs with variable life habits and morphology.
Classification of such a group is not easy. However, recognition of smaller taxonomic groups
of fossil bivalvia such as families, genera and species is based on shell-forms and structures,
the presence or absence of pallial sinus, dentition and such other characters. But recognition
and grouping of higher categories of fossil bivalves into a meaningful phylogenetic subdivisions
is difficult. This is because soft part morphology and shell-microstructures actually bearing most
of the useful informations about the phylogeny of the group are rarely preserved in fossil
specimens.
The classification given below mainly follows the scheme proposed in Treatise on
Invertebrate Palaeontology (Moore et. al., 1969) based on shell-microstructures, dentition and
other hinge structures, gill types and such other features.
pELECYPODA ( 8/VALVIA) 159
Cl-: Bivalvia (Pelecypoda) (Cam.-Rec.). Shell mostly bivalved, hinged dorsally, bilateral
symmetrical with ligaments, head lacking. e.g. Calms.
Subclass 1. Palaeotaxodonta (Ord.-Rec.)
Order : Nuculoida (Ord.-Rec.). Small, protobranchia, taxodonta e.g.

Nucula.
Subclass 2. Cryptodonta (Cam.-Rec.).
Order : Solemyoida (Cam.-Rec.). Largely toothless, infauna! e.g.

Solemya.
Order : Precardioda (Cam.-Rec.). e.g. Cardiola.
Subclass 3. Pteriomorphia (Ord.-Rec.). Normally byssate bivalves with variable
musculature and dentition.
Order : 1. Arcoida (Ord.-Rec.). e.g. Arca.
Order : 2. Mytiloida (Ord.-Rec.) e.g. Mytilus.
Order : 3. Pteroida (Ord.-Rec.) e.g. Pteria.
Subclass 4. Palaeoheterodonta (Ord.-Rec.). Variable imperfect heterodont dentition.
Order : 1. Modiomorphoida (Ord.-Re~.). heterodont imperfect, e.g.
Redonia.
Order : 2. Unionoida (Trias.-Rec.). e.g. Unio.
Order : 3. Trigonoida (Sil.-Rec.). e.g. Trigonia.
Subclass S. Heterodonta (Trias.-Rec.). Heterodont, eulamellibranchs, infauna!, siphon
feeding.
.
Order : 1. Veneroida (Jura.-Rec.) e.g. Venus, Cyrena.
Order : 2. Myoida (Trias.-Rec.) e.g. Mya.
Order : 3. Hippuritoida (Cret.) e.g. Hippurites.
Subclass 6. Anomalodesmata (Ord.-Rec.). Anomalous teeth.
Order : Pholadomayoida. Burrowing or boring, desmodont, e.g.
Pholadomya.
13.7. ECOLOGY (Fig. 13-7)

The mode of life of bivalvia is quite variable which is sometimes reflected by their shell-
morphology. Understanding the ways in which modern pelecypods are adapted to different modes
of life enables reasonable inferences to be made as to how their extinct counterparts lived. As
regards the nature of the life habit, the modern and extinct bivalves can be grouped into
following ecologic subdivisions (Stanley, 1970).
Palae(Geo)WP-21
~
.,._ 0
[sEA WATER l B
- ,. ,. + T'
. -. .,.-. . .. - _ • ;:.-.:, ..
-+
~
..
_._
+
+
1"
'T
•. ,.
T
~
+
"+
1
1 ~
-
~
+
. , .:... •to"~ + ... T
.....~
+ -+
... +
T
-r
~
.+
...'"'t
~+ +
T
~ ~
+
-4,
+
'P' T" ... , . .,. 1"
+
-" ... . ~
+
.. ...- . .
+
+
-t'
T
-t'
+
....
~
...
+
~
•
T
t
1"...--++ + ,ti' + -+ "t" +
- • -t_._ -t't,t-T + + • t-T .._
I -+' ~
t .. ~
.,. -t ~ + ~ t +1-t+.,. + + + ' t + t- + ++- .. +
t t -r+ t -+ + +
t-t+ ·t-;-t t i
-t t
t
+ t'
"t
~+"" ,_,-
4, t-" t t • t T ~1",+ + + -t-+,1-
t t t t - + + T T-t 1 + +-~ t -t T't" + T + +t t + t+
+t -t+ ~ '\-.,.-tt + + ...
1"
t -tt--tt t++ 'r,.. t t + , . , +- ~+ 1
~
T
.,... 1-
. 't
+- t ·t- + -t + ~ + ,t 1' t"t 1"1"~ t .+ 1'-t + ++
1
1- ... r . ~ .,...,..++ -t +'" .l,,,t + +-t 4
,.t- t -t..,.t ~ t t- ~ .,.. t- -t + -t"f t -t ;- 't-+ "T-t ~ t++ t -t +
.. ~ ~ t ~ 't t t t t ;. /" t + ~ + * -+ -+
t'
~ t.,.. + f- ~ -t -t
-t- t -t -t- ~ t' -\" -t 't . t
~ .... t +- ..... t ... t .,.. -t i" -+ t +"t it' 't T ,t t "t-t-t-t
-1' f' +
T
+ -t . 1-tt t .,.-t .+ t . t -t ++-+
1'w 1" + + - t " t -f "t .,-+-t t

... t +- + t +- ·t' t t t f t -t 1' 1"'" -t++ + ++~
+T 1' - , . t t ' ~
-t +
~ ~ + ·t- \- t t t t t t + -t
t t -t t t t ./t
, t t t - + ; t t~
-\' ·t t t + -t -+ 1' *' + -t- + +
+ +- + t t t- -r t t -t "t
1'
~
t
t,. t
-t i" ·t-
..,.. +~ + r ~+
t
-""+.~_t-~-+ !
~
FIG. 13 -7: LIFE HABITS OF SOME MARINE PELECYPODS
~
A : Rock Borer• B: Biyssally Attached• C: Free Lying Sessile, D: Scmiinfaunal (shallow burrower), E: Deep Burrower, F: Nl,rnlal <:
Vagilc Bcnthic I H: Cemented Gregarious I Cemented Isolated• G: Swimmer.
23
r--
0
C)
"'<
161
PELECYPODA ( 8/VALVIA)
(a) Normal epifaunal benthic vagile forms . .

Many pelecypods live on the sea bottom as normal vragnants. sluggis.hly ~ovmg by their
muscular foot. lying at the anterior. They have normal shell-forms, meqmlateral valves.
anisomayarian muscle scar and a simple pallial line. Cardita, Arca are such forms.
(b) Infaunal shallow and deep burrowing forms .
Many pelecypods live within sea bottom by burrowing the substrate (sandy) by thetr shells
activated by the foot and muscles. Most shallow burrowers are equivalved or nearly so,
with nearly isomayarian muscle scars and with a small pallial sinus. The anterior-posterior
line is approximately parallel to the hinge line. The shells usually show a small gapping
in both interior and posterior direction for the passage of foot and siphon. Some burrowers
have a peculiar external surface sculpture or ridges helping them in easy burrowing.
Cyrena, Divericella are such bivalves.
There are however a number of deep burrowers which have usually very elongated,
strongly inequilateral (posterior very large) shells. They possess a small anterior but a
very big posterior gape and a deep pallial sinus (for the passage of a vuy long siphon).
Muscle scars are very much unequal. Mya, E11Sis. Solen are the examples.
(c) Bys.sate epifaunal forms
Some bivalve groups possess some thread like structures attached below the anterior
auricle called byssus threads, by which they fix themselves temporarily with the
substratum. Pectinid pelecypods are such forms. They are mostly attached in an upright
position with dorsal upward. Very often the zone of attachment of ventral margin of the
shell becomes almost flattened by high reduction of anterior side of the valves and so
also the anterior adductor muscle, so that in many cases they become almost
monomayarian. These forms may be epibyssate when they attach themselves to some
sea-weeds or rocks or may be epifaunal and endobyssate when byssus threads anchor
within the ~ediment like plant roots.
(d) Epifaunal cemented forms
Among the pelecypods which are attached by fixing themselves with hard substrate.
oysters (Ostrea) are the most successful forms. They are mostly found in their original
life-form as fossils and are also found profusely in the modern sea. They attach
themselves to the sea-floor by their left valves cemented to some hard substrate and this
encrusting valve becomes usually thicker, larger and more irregular than the other valve.
The shell becomes in general very thick and monomayarian.
Another group of cemented bivalves are the extinct rudistids which show a great
modification of their shell-forms. The two valves become very much unequal. The larger
right valve by which it was attached to the substrate becomes conical and it was fixed
by the pointed end of the cone. The left valve becomes much smaller, nearly flat lying
on the right valve as a lid. They are very much alike the skeletons of some Palaeozoic
solitary corals and Permian richthofenid brachiopods due to their adaptation to a similar
mode of life.
Gryphaea was another sessile form otherwise free lying. It has a very large left valve
with pointed inwardly curved umbo by which it possibly was attached to the substratum
PALAEONTOLOGY
at least at their juvenile stage. This gave them stability and prevented overturning of the
shell by the action of current.
(e) Swimming forms
Byssate pectinids have very thin valves. This indicates that thei~ attac_hment by bys~us
threads is rather a temporary one. More often they are able to swim qu1~kly by clappmg
of thei.r valves together so as to expel water in successive jets on both side~ of the ear~.
As such activity was exhaustive the animal cannot sustain it for a long time, when tt
settles to the bottom and attaches itself with their byssus threads.
(f) Boring forms
Some bivalves .are adapted to a life within some hard substrate like stones or woods
through boring the substrate. Pho/a (rock borer), Lithophega and Pteredo (wood borer)
are such forms. They have also very elongated shells and usually they live with their
long axis vertical. They have also a very long siphons. The shells are however very much
resistant as they excavate the hard substrate by the edges of their shells. Frequently, the
shell edges are pointed with spines (used as scrappers for excavation).
Synthesizing the above discussion the following inferences may be drawn :
(i) Vagile bivalves are mostly inequilateral, equivalved isomayarian or nearly so with
normal pallial line.
(ii) Shallow sand burrowers are also like the above forms but normally with smooth
surface, a streamlined body and a small pallial sinus.
(iii) Deep burrowers usually possess inequilateral, but elongated shells (posterior much
larger) with _gapping in both anterior and posterior direction and with a large pallial
sinus.
(iv) Hard rock-borers also possess similar shells but the shells are much harder and
thicker, often cylindrical.
(v) Byssate forms are alates, mostly inequilateral with larger posterior, strongly
anisomayian and they normally bear byssus notch.
(vi) Cemented forms have inequivalved thick shells, the attached valve becomes much
larger with a very rugged external surface; the other valve becomes smaller and
flat. The shell becomes monomayarian.
(vii) Free lying sessile forms also exhibit unequal valves, the underlying valve becomes
much large~ and thicker often with spines and crenulations. They also become
monomayanan.
13.8 GEOLOGICAL HISTORY AND STRATIGRAPHIC VALUE (Fig. 12.3)

Bivalves are thought to have arisen from an extinct minor group of mollusca, the
~ostroconcha, w~ich also po~sessed a two-valved sh.ell, but u.n~inge~. The first bivalve appeared
m Lower Cambnan. Expansion of the group began m Ordovician with the development of more
forms adapted to several other life habits like deposits feeders, deep burrower and forms attached
by byssus. Most of the bivalves of that time were living in rnarine saline water and many of
.....
p£LECYP0DA ( BIVALVJA) 163
them became extinct after Permian. With the beginning of Triassic, appearance and
diversification of many other groups took place and most of them are still living. Some genera
appeared in Mesozoic times are Trigonia, Gryphaea, Alectryonia, Exogyra, /noceramus Ostrea,
Hippurites, etc. Of these, Gryphaea, Exogyra, Alectryonia and Hippurites became extinct after
Cretaceous. Pelecypods become very much abundant during Cenozoic with the appearance of
most of the modem forms of today.
As most of the pelecypods are long ranging forms, they rarely serve as good index fossils.
A few short-ranged forms of Palaeozoic and Mesozoic however are used as index fossils such
as Eurydesma (Lower Permian), Hippurites (Cretaceous), Alectryonia (Cretaceous) etc.
'
Chapter 14
GASTROPODA
14.l GENERAL FEATURES . . . . , . . .

. I d II sna1·1s and slugs llvmg m a wide range of environments, manne
d
Gastropo s me u e a • . · ed II I · '
. Th name refers to its fleshy foot satuat a a ong its ventral side
freshwater an d terrestna1. e . .
which also bears internally the stomach of the animal (Gk-~astros . stomach;_ podos : foot).
hich are almost always coiled and usually m a conispiral
They secrete exos ke Ieton Shells W .
manner. InternaII y, they Possess a well-defined
. head, bearing tentacles and sensory organs .
Mouth contains a rasping jaw' radula at its lower p~rt. The mantle lobe encloses digestive
canal, gills, nervous system and reproductive system (Fig. 14-la). Internally these organs shows
a peculiar anterior twist (Fig. 14-lb). This internal a~ymm~try, often call~ 'torsion' is a
fundamental features of gastropod morphology. The curious displacement of mtemal organs is
no way connected with the asymmetrical helicoid coiling of the shell. Garstang (1951) suggested
that such torsion takes place in veliger stage (second larval stage following the initial
trochophore) to protect the larva, and is retained in the adult form. The two main effects of
this torsion are, firstly, the change of orientation of the shell relative to head-foot mass which
is sited posteriorly instead of anteriorly above the head. This becomes a convenient alteration
for the animal. But the second alteration caused by to~ion by which the mantle cavity originally
lying posteriorly now opens anteriorly. This results the exhalant current, bearing metabolic waste
and faeces have to be expelled through anus located over the mouth. To overcome this unwanted
consequence many gastropods have developed some devices to separate and divert the inhalent
and exhalant stream. In many forms an indentation (slit band) developes along midway of the
sheJI margin forming a channel carrying exhalant stream dorsally away--from head region (Fig.
14-lc).
14.2 SHELL FORMS AND COMPOSITION

Gastropod shells are made of calcium carbonate usually in the form of aragonite. Living
forms are found with sheJI coated externally by a thin organic film called periostracum. During
fossilization process this is mostly lost. Below this, occurs the shell proper which is also
composed of aragonite but divisible into two zones, an inner layer of thin aragonite lying parallel
to shell surface (hypostracum) and an outer layer where aragonite lamellae are arranged normal
to the shell surface (ostracum) (Fig. 14-ld).
The sheJJ, except a fe~ (~.g. Clio), is a coiled one. Helically spiral shells (asymmetric) ~re
more com?1on t~~n plamsp1ral froms (bilaterally symmetrical). The imaginary central lane
around which cmlmg taJces place is called the axis of coiling.
14.3 A. MAJOR MORPHOLOGIC FEATURES ASSOCIATED WITH SHAPE OF THE

SHELL (Fig. 14-2)
Apex : Point of beginning of shell growth (protoconch).
164
GASTROPODA 165
Ee
---A.
.---Sh.
lc---..1
(a) OORSOLATERAL VIEW OF A MODERN GASTROPOD SHOWING SOFT PARTS
Alimentary
canal straight
Vi.seral mas------
(Postaior) Anus (Anterior)
(c) REMOVAL OF WASTE mROUOH PERIPHERAL SLIT-BAND

(Sclenizone) lN BELLEROPHON
Hypostncum---
(cl} SHELL-WALL
STRUCTURE
FIG. 14 - I : GENERAL FEATURES OF GASTROPODS

A : anus, Ee : cxhalant siphon, F : foot, Ft : food tube• 0 : gill, le : inhalant siphon, M : mouth, Mc : mllltlc cavity, 0 :
operculwn, Sh : shell , SI : slit band , T : tentacle • · ·
166 PALAEONTOLOGY
--Tuberelc
Axis Of Coiling (Spine/Node)
---Ramp
I
I
Q
1
Antaior
Anterior Siphonal
' (ii) Canal .
A,r-shoutder
~
(a) MEASUREMENTS, ORIENTATION &
COllJNG OF SHELL Spire= Body
Gutter
Whorl
(ii)
Relative Length Of Spire
&BodvWhorl
T- ~ ~ ~2Jeocoo4ru~~
p-
!m ~ ~ ~ A '!? ~-
... - - • Ortustaopbic --1 (c) SHELL FORMS
~~4, ~$ ~e L1 & Cooispiral Myperstrophic Form

Uncoiled Hetaostiophic
Shell
! ®~ @2J~
Advolute
T i ~ Of Coiling
(d) COILING OF SHELL
FIG. _14 - 2 :- MORPHOLOOY OF GASTROPOD SHELLS
GASTROPODA
If{/
Base : Part of shell-surface furthest away from apex.
Whorl : A complete revolution (360°) of shell around its ax b,.
Whorl profile : Whorl outline as appeared in the axial plane; it may be flut, convex or
concave outward.
Periphery : Part of a whorl furthest away from shell axis.
Spire : All the whorls of a shell except the last one. It may be larger, smaller or
equal to the last whorl. A shell with very few spiral who rls is called
paucispiral; a shell with numerous spiral whorls is called multiapiral.
Body whorl : The last whorl (usually the largest one) of the shell, bearing the soft animal.
Length of spire may be larger, smaller or equal to that of body whorl.
Shell shape : Usually defined by peripheral outline of the shell as appeared in axial plane
(Fig. 14-2c) which becomes quite variable and some common forms are as follows.
(i) Turreted : Elongated conical; spire much larger than body whorl e.g.
Turritella.
(ii) Conical : Spire and body whorl are nearly equal; spire with pointed
apex, base flat e.g. Trochus.
(iii) Biconical : Spire and body whorl both form moderately elevated cones
in two opposite directions (apex and base both pointed) e.g. Physa.
(iv) Conoidal : Like the biconical form but basal cone becomes steeper e.g.
Levifusus.
(v) Extraconical : Like biconical but just opposite of conoidal i.e. apical
cone is more steep and sharp.
(vi) Fusiform : Biconical forms, widest at the middle e.g. Fusus.
(vii) Obconical : Reverse of conical form, base conical, apex flat e.g. Conus.
(viii) Ovoid : Egg-shaped, moderately elongated with rounded apex and base
e.g. Oliva.
(iv) Turbinate : Top-shaped, rounded base, pointed apex; shortened conical
e.g. Turbo.
(x) Globular : Almost spherical, highly inflated last whorl, spire negligible
e.g. Nerita.
(xi) Pupaeform : Slightly elevated, ovoid, like pupa of insect e.g. Vertigo.
(xii) Patelliform : Low cup-shaped shell like that of Patella.
(xiii) Discoidal : Highly flattened apex and base; more common among
planispiral shells e.g. Planorbis.
(xiv) Vermicular : Irregularly coiled like a worm e.g. Vermitus.
Palae(Geo)WP-22
. .,......,,..
PALA EONTOWCY
\68
M~
~tiooal : __ -Umbilicus I v~
VIC\V
Pcrphorated shell
{a) COLUMELLA AND UMBIL1Cl1S

Slit Band1Selcni1.ooe
lncurrcnt Excurrmt
~/
CDOil<ur (ii) ~
Holostomatout
Anterior
Cana.I (i)
Anterior Peripheral ApcrturaJ Sick v~· Aperture
Canat View View
Siphooostomatous
Aperture
(b) SIPHON AND SLIT BAND·
Reticulate
Structure
Costac
( Spiral
Granules
(Varix)
tlnffl~- Axial
Granules
Apcrtural
Digitation
(c) SURFACE SCULPTURES
FIG. 14 - 3 : MORPHOLOOY OF GASTROPOD SHELLS
GASTROPODA 169
8. Colling Patterns nnd Asoclatcd Features (Fig. 14-2d)

Conlspiral : Coiling around an axis like a screw, i.e. coiling of each whorl takes place
('Irochosplral) in different plune.
PlanJspiral : Coiling around an uxis in a single plane.
Orthrostrophic : Common conispiral coiling in which whorls are coiled in an erect cone so ·
that the apex pointing upward.
Hyperstrophic : Uncommon consipirnl coiling in which shell is coiled in an inverted cone
so that the apex points downward.
: Holding the apex pointing upward (for orthostrophics) and downward (for
hyperstrophics) and keeping aperture towards the observer, if the coiling is
found clockwise so that the aperture would lie on right-hand side of the
observer e.g. Voluta.
~al : In the above case, if coiling is anti-clockwise; aperture would be to the left
hand side of the observer e.g. Phyl·a.
HeterostrophJc : Shell showing abrupt change of type of coiling between initial and later
formed part of shell.
Evolute : Loosely coiled, whorls not in contact e.g. Ecculiomphalus.
Advolute : Whorls just in contact but not embracing e.g. Planorbis.
Involute : Outer whorls slightly or strongly covering the earlier whorls e.g. Natica.
Convolute : Outermost whorl covers all the preceding whorls that are almost non-visible
externally e.g. Cyprea.
Umbilicus : A central gap along the axis of coiling from base to apex in some conispiral
[Fig. 14-3a(ii)] shells enclosed by the inner whorl wall when the latter does not reach the
central axis; umbilical cavity opens at the base of inner lip of aperture; such
shell is called peforated (e.g. Natica). Sometimes it may be closed by
secondary shell growth.
Phaneromphalus : Shell with a wide open umbilicus (phanero : evident; omphalus, cavity).
Hemlomphalus : Shell with partially open umbilicus.
Cryptomphalus : Shell where umbilical opening is closed by shell materials (crypto : hidden).
Anomphalus : Shell lacking umbilicus (thus bearing a columella).
Columella : A pillar or rod-like structure along the axis of coiling extending from apex
[Fig. 14-3a(i)] to base formed by fusion of inner whorl wall when it reaches the axis of
coiling; columella when present, shell would be obviously devoid of
umbilicus and is called imper/orated e.g. Turritella.
PALAEONTOLO t
170
:-- -
-------: I
I I
I I
I I
I
I
I
I
I
I
'- ------
Shell with radial apc:rturc
"i
I
I
/
/
/
I / /
. / ... - -
··- -,t:' ~
;
Plane of aperture
Shell with tangential aperture
Law of shell balance
Posterior
Exhalant
current
om
Slit band
FIG. t4 _ 4 : MORPHOLOGICAL ANALYSIS OF SHELL-FORMS (aft.er Linsley, 1977)
'
hrt",
.· , II
l
'I
I .
(.
i
GASTROPODA 171
C. Features Associated with Spire (Fig. J4.2a-b)

Apical or : Angle subtended at the apical point of shell by two lines, tangents to the
Spiral angle spiral whorls if they touch all of them. If all the whorl are not touching the
lines, the apical angle may be determined by the two lines, tangents to the
last whorl of the spire.
Spiral suture : Siprally coiled junction among the succesive whorls. This junction may be
indistinct (flushed), depressed or elevated.
Dip
(sutural angle) : Angle of deviation of spiral suture from the plane normal to the shell-axis.
Ramp and Shelf : In many forms, surface of each spire may be divisible into two distinct areali :
a vertical portion (ramp) and a horizontal part (shelf).
Shoulder : Salient angulation of a whorl periphery parallel to coiling generally formed
at junction of ramp and shelf.
D. Features Associated with Body Whorl (Fig. 14-2a, b, 14-3b)

Aperture : Opening of the shell for the outlet of the animal. This may be variable in
outlines, e.g. circular, subcircular, rhombic, rectangular, elliptical, cresentic,
slit-like etc. Aperture is sometimes closed by a circular plate called
operculum.
Peristome : Margin of the aperture; may be smooth, crenulated, denate, spinose or
digit~d; Peristome has two sides : outer lip or outer margin away from shell
and inner lip or inner margin towards the axis.
Parietal Lip : Portion of inner lip towards posterior side.
Columellar Lip : Portion of inner lip towards columella (towards anterior).
Siphonal notch : Notch on anterior end of aperture occupied by inhalant siphon; when present,
(Fig.14-3b) it lies between columellar lip and outer lip junction; may be small or deep,
often extended as a long anterior canal. A similar notch may be present
toward posterior exhalant current.
Holostomatous : A shell without siphonal notch on the apertural face; apertural margin entire.
Siphono-
stomatous : A shell with a siphonal notch on the apertural face usually placed anteriorly.
Sinus : Groove or re-entrant in lateral margin of aperture with non-para1lel sides.
Selenizone : A sharply defined band found in some shells parallel to coiling of whorls
(Slit band) bearing fine cresentic growth lines (lunule), denoting a slit on outer lip for
(Fig. 14-1 C, 14-4) the passage of exhalant stream. ·
Gutter : A notch or groove at posterior extremity of aperture in some gastropods
[Fig. 14-2b(iii)] marking anal outlet.
Neck : Constricted anterior part of body whorl.
[Fig. 14-2b(i)]
- '
PALAEONTOLocy
172
E. Surface Ornamentations (Fig. 14-3c)

Van·.x :
R'd
1 ges,
fl anges
. or rows of spines , tubercles or granules, parallel to spiral
whorls.
Axials : Similar structures running axially.
Costae : Coarse thickened lines running spirally or axially.
Carina : Spiral keel (ridge) at edge of shelf.
Callus : Thi~kened shell-deposit on anterior part of inner lip partly or wholly covering
[Fig. 14-2bi] umbilicus.
Inductura : Layers of lamellar shell materials all along inner lip often extending beyond
the outer lip devoid of any ornamental features. This also includes the callus.
Digitation : Finger-like outgrowths from margin of outer lip.
Spines : Rows of sharp angular projection parallel to axis or spiral whorls.
14.4 CLASSIFICATIONS
Classification of gastropods is largely based on soft parts like gill, osphradial morphology
(osphradia is a specialised organ inside whose function is to sample the water entering the
cavity). The scheme below mainly follows that given in Treatise on lnvetebrate Palaeontology
(Moore et. al., 1969).
Cl~ : Gastropoda.
Subclass 1. Prosobranchia (Cam.-Rec.) : mostly marine,
Order : 1. . ~rchaeogastropoda (Cam.-Rec.) : Gill filaments arranged
m a double comb on either side of axis; marine. e.g.
Beller.ophon, Eomphalus.
Order : .2. Mesogastr~poda (Ord.-Rec.) : Mainly have pectinibranch gills,
more efficient than earlier types e.g. Cypraea, Natica.
Order : 3· ~eogastropoda (Cret.-Rec.): Pectinibranch gill, long inhalant
siphon, e.g. Murex, Voluta.
Subclass 2. Opisthobranchia (Carb.-Rec.) M
: arine, have lost the shell partially or
completely e.g. Sea slugs.
Subclass 3. Pulmonata (Perm.-Rec )· Land d 1 ·
by modified lungs e · ~ . we h~g slugs and snails, gills replaced
.g. a11oma, Vertigo, Physa.
14.5 ECOWGY
Gastropods chiefly live on the shallow sea botto A
depth of more than 3 miles. Most of them are "\ few ~orms are found to live on ocean
swimming or passively floating near surface wate vagfihe benthic. A very few forms are found
. . r o t e open & M
gastropods are found burrowmi m shallow sea 'th' ocean aar away from land. any
exhalant and inhalant siphons. Terrestrial gastro;~
O
~n sand and they are mostly provided w.ith
on land, climb on trees or ascend hills to an el . ave lungs for respiration and they can hve
evatton of 18,000 ft. above MSL .
,,,. • ,.,,. r
, . 1
( ,'AS'l'N( )l•(Jl)A
173
Most uf th· ~nnils urc livin~ on various sorts or vegetation like phytoplanktons, leaves of
plants le. Som nr' cnrnivorous predutors und kill some invertebrates (bivalves) by boring their
shells and ·uting lh ir 11csh. A few live 11yrnbiotically with other invertebrates. Association of
l:rahs nnd 'forritt'/la, ph11'1c.:erutid gustropods with crinoids are few examples. Attached or
hurrnw rs nrc mostly filter feeders. Development of a long inhalant siphon found in some
gaslropnds muy have preudapted such forms for other modes of life. The burrowers employ it
ns a snnrk I und carnivorous forms tend to use it as a sense organ. There are however many
id ·ntifiabl" nnd r curring shapes of shells within gastropod in different geological times. This
111uy suggest that there muy be more functional significances of the overall shell-shape. Careful
study has shown that marine gastropods living in hard bottom and soft bottom tend to develop
two distinctive types of shell shapes. With change of environment and condition many species
b ·came extinct and replaced by new species and the later developed same shell morphology
suitable for such condition and this strongly suggest that some type of shapes were very much
favoured by certain environmental conditions.
14.6 MORPHOLOGICAL ANALYSIS OF GASTROPOD SHELL FORMS (Fig. 14-4)

Important analysis has been made on the shell-form of gastropods and five laws have been
postulated (Linsley, 1977) as a foundation of functional morphologic analysis of prosobranch
gastropods.
(a) Law of radial aperture
Gastropods with more than one whorl with radial aperture (planispiral or near so) live with
their plane of aperture perpendicular to the surface. The plane of radial aperture lies along
a radius from the axis of coiling to the shell-periphery as found in Architectonica. These
types of apertures are rare in modern gastropod but are common within Palaeozoic forms.
(b) Laws of tangential aperture .
Gastropod shells with more than one whorl and with tangential aperture (plane of aperture
forms a tangent to the earlier formed whorls) live with plane of aperture approximately
parallel to substrate. Most living prosobranchs have tangential aperture.
(c) Law of shell balance . . .. .

If the shell of a gastropod is supported above the body, 1t will be pos1t1oned m such a manner
so that the centre of mass of the sheH is just over the mid-line of head-foot mass. Most of
the living species carry their shell mass with the centre of gravity above the centre line of
body mass.
(d) Law of re-entrant .
Angulation or re-entrant on the apertural face usually indicates presence of inhalant and
exhalant siphons; the former is directed most anteriorly and the latter posteriorly. This feature
is found in many mesogastropods and neogastropods where a single current enters the mantle
cavity anteriorly through inhalant siphon, passes over the single gill and leaves the cavity
posteriorly by exhalant siphon. However, in archaeogastropoda there is a pair of gills within
mantle cavity where two inhalant currents enter the body along two anterior-lateral directions
and the exhalant water is expulsed peripherally along the selenizone (e.g. Bellerophon).
174 PALAEONTOWGY
(e) Law of elongation

Th ~ . .l· the entire length of shell (Cyprea)
astr pod havtn . ,. an e lon gal ,t1 aperture a ong .
po · s • , u.··" ,. , s, 1·11 ,. Ic I. 11 ms1
. .dt.~. nn d I-1 deve lopes. •,1 ·L
"·,·,,g· le current of water .from antenor . to
po t ri r direction through mantk cavity. They usually posses small anterior and posterior
notch s on the apertural margin.
l4.7 E OLUTION AND GEOLOGICAL HISTORY (Fig. 12-3)

It is more or le .. s accept~d that gastropods evolved in Cambrian _from a 'monoplaco~horan'
ancestry which for th~ first time exhibited the phenomenon of torsion of shell. ~e~e 1s some
deb~te about it~ nature of coiling whether conispiral or planispiral. One hypothesis _is that the
earh t gastropod s are planispiral bellerophonids, a descendant from monoplacophonds. Others
consider heller phonid. as an intermediate forms between true gastropods and monoplacophorids.
A.pan fr m the planispiral forms, a few conispiral forms are also found. At least nine families
are recorded in Cambrian, all belonging to archaeogastropods. A great enlargement in number
of gastropod genera is found from Ordovician when apart from the nine existing families
fourteen additional families made their first appearance. Eight additional families appeared in
Silurian, two in Devonian, one in Carboniferous and three in Permian. Most of them were
belonging to archaeogastropods and mesogastropods. Opisthobranchia, appeared for the first
time in Carboniferous and is continuing till now Neogastropods including most of the modem
gastropods appeared in Late Mesozoic times but became abundant after Cretaceous. Palmonates,
although, appeared in Late Palaeozoic become abundant only after Tertiary.
Chapter 15
CEPHALOPODA
Cephalopods are a group of highly advanced marine molluscs. It appears that the animal is
moving by using its head as foot (Gk.-cephalon : heads; podos : foot). They include extinct
ammonoids and living forms such as nautiloids and coeloids (octopus and sepia). In most of
the cases, the animal bears a calcareous shell surrounding the soft parts which may be coiled
or uncoiled.
Cephalopod's anatomy (Fig. 15-1 a) is conspicuously different from the basic molluscan plan.
There is almost a 90° shift of standard orientation forming a ventral head-foot pointing anteriorly
with a shell placed posteriorly. The head bears a number of strong tentacles, used as functional
feet; a mouth with a strong radula, a pair of highly efficient eyes and also advanced type of
gills. The mantle cavity migrates on all the way from back to point anteriorly bearing one or
two pairs of gi1ls below the head-foot mass. Its opening is surrounded by a mascular organ
called funnel or hyponome which is able to direct jets of water in forward direction controlling
the characteristic jet propulsion mode of locomotion that enables the animal escaping quickly
away from its enemy within water. Unlike other molluscs (mostly bisexual), sexes are separated
within cephalopoda. As they are swimmers, there is no larval stage and the young animals grow
directly from eggs. Most of the living cephalopods have no she]] but have some internal skeletons
in some modified forms. As regards food, most of them are carnivorous living on actively
moving sma11 to large sized preys. Cephalopods were dominant marine forms from Late
Palaeozonic upto Mesozoic, but at present they are declining in number.
15.2 MORPHOLOGICAL FEATURES OF CEPHALOPOD SHELL (Fig. 15-1-3)

A shell of cephalopod is in most of the cases, an external skeleton except in coeloids where
it is internal. Shell is altogether absent in octopus group. Shell in a living nautilid is mainly
composed of two layers (Fig. 15-lb) both aragonitic; an outer porcellaneous layer cal1ed
ostracum made up of aragonite prisms and an inner 11acreolls layer (made up of aragonite and
organic matter. In addition, the entire inner surface of the shell is coated with an opaque
calcareous film often called ann11l11s layer. A septum is composed of elements of nacreous layers
(aragonite and organic matter) which is not usually preserved in fossils. The shell may be coiled
mostly in a planispiral pattern. But there are many fossil cephalopods which exhibit uncoiled
or even partly coiled shells. Coiled shells exhibit bilateral and some uncoiled forms show radial
symmetery. Unlike gastropod, the cephalopod shell is multichambered and the animal lives
within the last chamber adjacent to the aperture, also called living chamber. Excluding livino
chamber, the remaining part of the sheJJ is caJJed pliragmoco11e. The initial chamber is called
protoconch and its pointed end is the apex of shell. The direction towards aperture is called
adoral end.
Palae(Geo)WP-23 175
__........_._lllfc-~~~~~-----shel1 -..J
c,,..
Dorsal M11ttlc Lobc----- ~~:::::=~~-----Septum
111,.___ _ _ _ _ _ Siplwncle
Hood--------
Tonguc~------- 4~ Chamber
Tcntaclc - - - - -
I Liver
---~~~Stomach
,
;
ariurn
Mouth
- - - - - - - - Heart
Jaw
~---------Anu.'1
Radula -_. ~ /~~
Gill
H~apCJosing 1t": ~t . MmkCavity
Hyponome~~~~~~
q_iif3 Ventral Mantle Lobe
Mantle Cavity
(a) SAGI'IT AL SECTION
~ Sepwm
Annulus Layer (Aragonitic)

Nacreous Layer (Aragonite & Urgaanic Matter)
Porcelaneous Layer (Prismatic Aragonites)
(b) WALL STRUC11JRE AND SEPTUM

:ti
~
~
<:
FIG. 15 - I C~IIEF VISERAL ORGANS AND WALL MICROSTRUCTURE OF AN IDEAL LIVING CEPHALOPOD (Nautilus)
d
t""-
0
C)
0-.::
Connecting Rings • ~
Suture
Septal Neck
~
:i:,.;
Saddle t""'
0
""O
Lobe Siphuncle 0
Plngmooone __.,
Camera
Protoconch ~
f../ l .Wl"~J (Chamber)
Whorl
Umbilicus
,tum
Living or
Body Chamber :I ) Aperture ff )
Hypooomic
Sinus
(a) AN INfERNAL MOULD OF NAUf/LUS (h) TRANSVERSE SECTION OF NAUT/1.US
Apertwc
Ahoral end
~·~~ Hyponomic Sinus
Body Chamber
Umbilical Plug
i
=-~~
Septum
Ownber
Protoconch
:1f&J
&--!H:.-...~M----Siph1D1cle
Phragmocone - ··
....::::~~~~~i--~--Do1num
I
1...----iff---Whorl Height
------iJ..,_-----Venter
Pm1oconch
Ap.:,
tel AXIAL SECTION ( Na111i/11.r) (d) LONGITUDINAL SECTION

UNCOILED NAUTILOID (Ortlto«ra.r)
FIG. 15 - 2 : MORPHOLOGY OF CEPHALOPOD SHELL

--..J
-...J
178 PALAEONTOLOGY
Gyrocone
Orthoc-one Cyrtocone Lituiticone (Open Coiled)
(Straight) (Slender Curved) (Initial Part Lossely
Coiled Later Part Straight)
(a) UNCOILD/PARTLY COILED SHELLS
Evolute
(Coiled ·w horls
Not Touching)
Advolute
(Coiled Whorls Barely
Touching Each Other)
V Involute
(Earlier Whorls
Party Enclosed)
V
Convolute
(Earlier Whorls
Fully Concealed)
(b) PLANISPIRALLY COILED SHELLS (Side Views)
(Type of Forms Based On Tightness of Coiling)
~n.nQno~
~~ ~ ~ ~ tw IQl Circular Subci~cular Ell·i~~ical Pe!t~::nal Rec~~~gular ~ ..
Sphacrocone (Cresentic Aperture) (Cadicone) (Platicone) (Discoid) Rhombic
(c) COILED SHELLS IN APERTUREAL VIEW (Oxycone)
Turrilicone
f
Hook-Shaped
(Criocone)
Boat-Shaped
Baculicone
(Initial Part Tightly
(Conispiral) (d) OTHER UNUSUAL TYPES (Heteromorphs) Coiled Later Part Smight) -
Venual Saddle I st Lateral Saddle Aperture
Dorsal Saddle
Goniatitic Suture
Ccratitic Suture
Saddle
\ Ammonitic Suture
Nautilitic Suture
Aperture
(f) SUTURAL PATTERN ON SHELL
FIG. t 5-3 MORPHOLOGY OF CEPHALOPOD SHF.T .T
CEPHALOPODA
179
The diffe~e~t morp~ol~gical features of a cephalopod shell given below are based on the
shell of the hvmg nautI101d Nautilus and also on numerous fossils of nautiloid and ammonoid
group found in Palaeozoic and Mesozoic times. Morphologically, nautiloid and ammonoid shells
exhibit many features in common.
A. Shape of shell (Fig. 15-3a, c, d)

Basically an uncoiled cephalopod exhibit a conical shell with the posterior part tapering away
and a board anterior with a large opening for the exit of the animal, called aperture. From
this basic form cephalopod shells show various modifications starting from a curved, partly
coiled to tightly coiled cones and accordingly their shapes also change. Terms used for shape
are mostly derived from the name of the genus ideally exhibiting the particular ·shape. The
different terms commonly ,used are as follows :
(i) Uncoiled/partly coiled shells (Fig. 15-3a)
Orthocone Uncoi1ed and straight cone, e.g. Orthoceras.
Cyrtocone Shell slightly curved. e.g. Cyrtoceras.
Brevicone Shortened cyrtocone, e.g. Breviceras.
Lituiticone Initial part coiled loosely, later part straight uncoiled. e.g. Lituiticeras.
Gyrocone Loosely coiled evolute type of shell; whorls not touching. e.g.
Gyroceras.
(ii) Coiled shells (as seen in apertural view) (Fig. 15-3c)

Sphaerocone Globular shell. e.g. Macrocephalites.
Cadicone Globular, but pentagonal in outline, e.g., Phlycticeras.
Oxycone Rhombic in outline, e.g. Oxyceras.
Platycone Rectangular in outline. e.g. Acamhoceras.
(iii) Unusual types (Fig. l 5-3d)

Turrilicone Conispiral, turreted form, e.g. Turrilites.
Hook shaped Apical and apertural end both curved towards each other. e.g. Hamites.
Boat-Shaped Earlier part tightly coiled later part straight, uncoiled. e.g. Scaphites.
Baculicone Initial part tightly coiled, rest portion straight, uncoiled. e.g. Baculites.
B. 'fypes of coiling (Fig. 15-3b)

Planispiral Coiling around an axis in a single planes; usually found in cephalopods.
e.g. Perisphinctes.
Conispiral Coiling around the axis at different planes rarely found in cephalopod.
e.g. Turrilites.
180 PALAEONTOLOGY
Evolute Coiled shell but whorls not touching. e.g. Gyroceras.

Advolute . h' nd alJ whorls visible, e.g.
Coiled shell, whorls JUSt touc rng a
Centroceras.
Involute Coiled shell in which earlier whorls seen partly enveloped by later
whorls. e.g. Macrocephalites.
Convolute . th preceding whorls. e.g.
Coiled shell in which last whorl concea.Img e
Nautilus.
C. Other shell-features (Fig. 15-2, a-d)

Whorl A complete revolution of shell around the axis.
Venter Whorl-margin furthest away from the axis of coiling.
Dorsum Whorl-margin towards the axis of coiling.
Whorl height Maximum distance between dorsum and venter.
Umbilicus Open or depressed zone in position of axis of coiling which is left
when successive whorls do not reach the axis and whorl-height of
successive, whorls increases towards periphery; umbilicus may be deep
or shallow depending on rate of increase of whorl-height.
Umbilical plug : Calcareous shell-deposit filling the umbilicus.
Umbilical shoulder : Portion of shell bordering the umbilicus.
Aperture Opening of the last living chamber; . may be variable in outline
(Fig. 15-4a) depending upon the shape of shell; it may be circular, subcircular, sub-
elliptical, rectangular, triangular, crescentic etc. Aperture sometimes is
found closed by one or two calcereous plates called anaphycus and
aptychus respectively.
Hyponomic sinus : Embayment in ventral margin of aperture for holding the funnel or
hyponome.
Chamber/Camera : Compartments within the shell; initial chambers filled up by gases.
Protoconch First or initial chamber; conical for nautiloids, barrel-shaped in
ammonoids. .
Pbragmocone Portion of shell with all chambers except the last one.
Body chamber Last or living chamber with aperture.
D. Septum and Suture (Fig. 15-2a, b, 15-3e-t)

Septum Partition wall in between two adjacent chambers, us~ally curved
concave towards the aperture. '
Suture Junction line of a septum with the inner wall of th 8 h II·
wavy, straight or frilled. e e , may be
CEPHALOPODA 181
Lobe In case of undulating suture, the portion of suture, concave towards

the aperture.
Saddle In case of wavy suture the portion convex towards the aperture.
E. Sutural types (Fig. 15-3f, 15-7b)

(i) Orthoceratitic suture : Straight suture line, characteristic of uncoiled nautiloids like
Orthoceras.
(ii) Nautilitic suture : Undulating suture line with development of rounded lobes and
saddles, characteristic of Nautilus.
(iii) Goniatitic suture : Suture undulating but lobes become angular and saddles remain
rounded; In some cases both of them may be angular, e.g. Goniatites.
(iv) Ceratitic suture : Sutures forming lobes and saddles, where lobes are further crenulated
or fluted but saddles remain rounded. e.g. Ceratites.
(v) Ammonitic suture : Sutures where both lobes and saddles show complex fluting;
sometimes, this becomes so much crumpled that it is locked like a tree, when it is called
dendritic suture. e.g., Macrocephalites.
F. Siphuncle and associated features (Fig. 15-2b, .15-4b, c)

Siphuncle A tube which leads from protoconch to the living chamber piercing
each septum; may be central (in nautloids) or peripheral or ventral
(marginal) as in ammonoids.
Connecting rings Delicate calcareous rings forming the wall of siphuncle.
Septa) neck Tubular extension from each septum at the junction of siphuncle
covering the latter.
Prosiphonate Septal neck projecting towards the aperture ·as found in many
ammonoids (Fig. 15-4b).
Retrosiphonate Septal neck projecting away from aperture as found in nautiloids (Fig.
15-4b).
Siphuncle tube exhibits three layers; an ·outer thin pellicle, a middle chalky layer (CaC0 )
3
and an inn~r harder horny layer (Fig. l 5-4c). Depending on nature of septal necks and
connecting rings siphuncle may be of three types (Fig. 15-Sc) : orthochoanitic where tubular
siphuncle is composed of short septal necks and intervening connecting rings; cyrtochoanitic
where siphuncle is expanded within chambers but constricted at the crossing of septa;
· holochoanitic where tubular si~huncle essenti~lly is composed of connecting rings and septa\
n~ck.s e~tend back to ~he previous septum. S1phonal deposits are usually calcareous formed
~·t~m s1phuncle and this ~ay be .of three types : annulosiphonate where the calcareous deposit
ms1~~ form~ an annular rmg thicker at ventral margin; actinosiplionate where the deposit
el xh1b1t(sF~ad1al51 a4rrangement; and endocones where deposit forms as apically pointed conical
ayers 1g. - c).
PALAEONTOLOGY
Subcircular Semi.Circular Semi·

~ ~
Crcsentic Elongated Rectangular
.. Rectangular
Triangular
Elliptical
(II) APBRTURAL OUTUNE (A : Aperture)
(b) POSITION OF SIPHUNCLE

Aperture
- - · - - - - Pellicle
~l~<:11----- Chalky Layer
i l.t'11.tl\---- Horny Layer

,:;i;:~~---Siphuncular Veins And Artery
Scptal Neck
A Longitudinal Section Through Siphon Tube

Prosiphonatc
Retrosiphonatc
Direction Of Septa! Neck
-m. ~ Annulosiphonatc
Septal Neck
§ 1 f-~
~~ ~ Aonnos;phooate
e:_tf:c ~
Orthochoanitic Cyrtochoanitic
; Type Of Siphuncles Based On Septa! Neck And

Connecting Rings
Types Of Siphonal ~ i t s
(c) Morphology Of Siphuncle
Sulcatc Carina1e
Carinate-Sulcatc
Cll1 PERIPHERAL 1--'EATURES
FIG: 15 - 4 : MORPHOLOGY OF CEPHALOPOD SHELL
CEPHALOPODA
183
G. Shell ornamentation (Fig. I 5-4d, 15-7c )
Capillae Fine raised lines radiating from dorsum to Yc!nter at right anglc!S to
whorls. ..,
Costae Similar to above but the lines are Yery coa.~: co·tae r capil!Je may
be concave or convex towards aperture, r it may be - traight at right
angles to whorl margin ( rectiradiate , s-shaped (jalcoid ; often they
become bifurcated. trifurcated or looped.
Tubercles Angular or spherical projections mostly present at the points of
and spines bifurcation of costae/capillae.
Peripheral A raised band all along the periphery of the _hell: often rope or ribbon
keel/carina like as found in Amalrhe us.
Peripheral sulcus A depression along the perphery of shell.
Carinate-sukate A central sulcus with two keels on either side of it. or ma be re, erse
periphery in some cases.
15.3 MORPHOLOGY OF BELEl\lNOIDS (Fig. I 5-5)

Belemnoids belonging to subclass coeloidea are an extinct group of cephaJopods of Mesozoic
times. Among them Belenuzites is an important fossil of Jurassic rocks.
In Belemnites the shell has two parts. The largest and hardest part of the shetl located
posteriorly, called guard or rostrum is a massive bullet-shaped or cigar-shaped body tappering
posteriorly, made up of solid calcite. It is indented anteriorly by a conical cavity. ab·eolus. A
transverse section of guard exhibits radially oriented calcite needles with some concentric growth
ring but the aixs of growth is shifted towards the ventral margin. The surface of guard is usually
smooth but rarely pitted or granulose. An additional skeletal elements, epirostrum is a tapering
rod extending backward from rostrum with a hollow anterior part (enclosing the tip of rostrum)
having a massive internal structure. Withing the alveolus cavity is embedded the second part of
the animal, phragmocone. This is a conical thin-walled aragoniric shell that projects anteriorly
outside the guard. The cavity of phargmocone is partitioned by number of septa, concave anteriorly
with a ventrally placed narrow siphuncle. This septed shell along with the bulbous initial
protoconch undoubtedly puts them within the cephalopod group. However, guard of this animal
has no direct homologue in the other living coeloids. The guard of Belemnites probably acted as
a necessary counter-weight to maintain equilibrium at the time of swimming, a function analogous
to that performed by camera) fluid of other cephalopods. A long flat tongue-like structure
pro-ostracum is found projecting forward from the base of dorsal pan of phragmocone possibly
protecting the anterior soft part of animal. It seems to be homologous to the 'pen· of a squid.
This part is rarely preserved in fossil. It has been assumed that belemnoids were squid-like animals
with different hardpart-construction and with a different mechanism of buoyancy control.
15.4 CLASSIFICATION
There has been a considerable debate about the method of subdivision of cephalopods. Many
authors have divided the entire group into two divisions based on number of gills. Others like
Palac(Gco)WP-24
c,:.
.:-
--------.Proosl111Cum Guard
A
-·
B
f>hragmoconc
----- Siphunclc (, .:ntral)
(c) SECTION THROUGH PHRAGMOl'ONl:
r-=7 I 11 I ~ Phragmoconc ( I 'i-
&Lt Concentric Gro"th Lines

~ Radial Calcite NttdJcs
\Vh' : : .t "' Protoconch ( 1 u __ ., D c- • D
C
L-----Alvcolus
/~• "fuuard Or Rostrum ( 1'i >
(d) SECTION THROUGH GUARD
...-----+ Epirostrum ~------4

(b) A VENTRAL VIEW EXPOSING PHRAGMOCONE ~

s:
ti,
(a) LONGITUDINAL SECTION SHOWING SIDE VIEW ~
FIG. 15-5: MORPHOLOGY OF BELEMNOID SHELL c3t""'
0
C)
"s::
CEPHALOPODA 185
TABLE-11
Distinction between Gastropod and Cephalopod shell
Gastropoda Cephalopoda
1. Univalved, mostly asymmetric.
Univalved, but mostly bilateraJly symmetri-
cal.
2. Shell usually coiled conispirally; uncoiled Shell usually coiled planispirally; uncoiled,
forms very rare. partly coiled forms are also found .
3. Interior of shell is unsegmented; no septa, Interior cavity of shell is divided into a
no suture. number of chambers (camerae) by partition
walls called septa which at junction with
inner wall produce suture lines.
4. Externally the shell has two parts, the last Shell of cephalopod internally shows two
whorls or body whorl and the rest part portions, living chamber (space between last
called spire. septum and aperture) and rest portion with
other chambers cal led phragmocone.
5. No structure like siphuncle or septa) neck. Siphuncle and septa! necks are associated
with inner septa.
6. Apertural face very often interrupted by Apertural face usually entire, uninterrupted.
siphonal notch.
TABLE-12
Distinction between Nautiloid and Ammonoid shell
Nautiloid Ammonoid
1. Shell coiled planispirally, uncoiled or Shell mostly coiled planispirally.
partly coiled.
2. Both radially and bilaterally symmetrical. Mostly bilaterally symmetrical.
3. Protoconch conical. Protoconch barrel-shaped.
4. Suture mostly orthoceratitic or nautilitic. Suture goniatitic, ceratitic, or ammonitic.
5. Siphuncle mostly centrally placed; septa) Siphuncle mostly dorsally placed; septa) neck
neck retrosiphonate. in most of the cases prosiphon~te.
6. Shell usually smooth without any Shell mostly sculputured by various types of
significant ornamental features. surface features like, costae, tubercles and
spines, peripheral keel or sinus.
to subdivide it into a number of subclasses based on some hand-part morphology seen in fossils
which obviously becomes more suitable for plaeontologists. The proposed classification of
Tiechert ( 1967) is given here.
PALAEONTOWGY
186
Class : Cephalopoda . . .
. - d bitaterat/radiat mner cavity
Shelled/unshelled; univalved; co1led/unco1 1e ; symmetry
septed; marine, e.g. squids, ammonites.
. . h ht to be ancestral of all other
Subclass 1. Orthoceratoiden (Cam.-Tnas) : A group 1 oug · 1 well developed·
cephalopods; straight or slightly curved shells; protoconch comca • •
. h ·t· ture e g Orthoceras.
cameral deposits present; ort oceratt 1c su · · · ..
. · hell· suture orthoceratit1c·
Subclass 2. Endoceratoidea (Ord.-Sil.) : Orthocomc1cyrtocomc s ' . h '
. tral septal neck retros1p onate;
siphuncle large with funnel-shaped endocones, ven _,
cameral deposit absent, e.g. Piloceras, Endoceras.
. t .d (Ord -Carb) : Shell orthoconic; ~uture c1thoceratitic; septa.I neck
Subclass 3. Act mocera 01 ea . · ·h d · huncular
retrosiphonate; cameral deposit present; siphuncle large wit en osip
canals. e.g. Actinoceras.
Subclass 4. Bactritoidea (Ord.-Perm.) : Shell orthoconic/cyrtoconic; suture orth~eratitic;
siphuncle small; cameral deposit absent, bulbous protoconch e.g. Bactntes.
Subclass S. Nautiloidea (Cam.-Rec.) : Shell orthoconic/nautiliconic; mostly with nautiliti~
suture; siphuncle small, subcentral; septal neck retrosiphonate cameral deposit
present e.g. Nautilus.
Subclass 6. Ammonoidea (Dev.-Cret.) : Shell small/large, coiled; ammoniticone; suture
ceratitic/goniatitic/ammonitic; septal neck prosiphonate, e.g. Goniatites, Ceratites,
Perisphictes, Baculites.
Subclass 7. Coeloida (Dev.-Rec.) : Small to large cephalopods, shell may be present/ absent;
straight. e.g. Octopus, Belemnites.
15.S FUNCTION OF SOME FEATURES FOUND IN CEPHALOPOD SHELL

A. Siphuncle
Since the success of cephalopods is lying on their possession of buoyant shells, 1t 1s
appropriate to consider how modem cephalopods (Nautilus) and their fossils forms (ammonoids)
actually achieve such buoyancy. Researches in these lines by Denton ( 1974), Ward and Martin
(1978) and other have shown that buoyancy of cephalopod works on a quite different principle
from that of fishes. In fishes, the swim-bladder contains gas at equal pressure to that of the
surrounding water; naturally in shallow water fishes the air pressure within bladder is much
less than those of deep water fishes. That is they can change the. specific gravity by increasing
or decreasing the gas pressure within bladder. But recent cephalopods (Nautilus) contain gas
r
I pressure in their empty chamber much less than 1 atm; (0.3 atm in younger chambers and
0.8 atm in older chambers). For this reason they achieve neutral bouyancy at different depth
not by pressure equalization but by density control. It is found that some chambers of the shell
also contain a liquid called cameral fluid which can be extracted out from the body through
siphuncle. Removal of such liquid from body will lighten the shell and it may rise in water.
The only disadvantage of cephalopod shells is that their non pressurised shells will implode
under high water pressure below a certain depth. In the living Nautilus cameral liquid is present
in considerable quantity in recently formed chambers and its amount diminishes from newer
to older chambers and the earliest most chambers are practically empty only with gas.
CEPHALOPODA 187
It is the activity of siphuncl e that extracts the liquid from the shell. The siphuncle consists
of two parts. The impermeable septa! neck and a permeable siphonal tube between them . This
tube has an inner core of living material with arteries and veins running along the whole length
of it with a cylinder of epithelial cells. Outside this, is horny tube of conchoilin fibre and
surrounding this again, there is a tube of irregularly arranged aragonite crystals. Both the
aragonitic and horny layers are very porous and permit the passage of the liquid through them.
The soft body of Nautilus is in contact with the septum during the time the latter is formed.
Eventually, the body moves away from the last septum and is separated from it by a cushion
of camera! liquid. Then a new septum is gradually formed at the base of soft part through
secretion of CaC0 3. Initially the chamber is filled up with the liquid but when the new septum
becomes, sufficiently strong to withstand the pressure of the sea-water, the siphuncle begins to
pump out the liquid to adjacent chambers leaving only a small residual amount. This pumping
process is worked by osmosis. Gas very slowly diffuses into the space left by the liquid which
can explain why gas pressure in a newly formed chamber is very low. Due to slow removal of
liquid from chambers, a cephalopod cannot adjust its buoyancy quickly but takes several week
to achieve full adjustment.
It is generally held that ammonoid achieved buoyancy in much the same way as does the
recent Nautilus and moreover its fluted septa might have increased the strength of the septum
so that it was able to resist implosion at depth as their 'Relative Strength Index' (R.S.I.) is, in
most of the cases weaker than ti1at of Nautilus.
There is good evidence from some rare unaltered fossils that Ca-phosphate is the primary
constituent of ammonoid siphuncle, though juvenile forms have calcium carbonate. R.S.I of
two Mesozoic orders of ammonoid, phylloceratids and lytoceratids is like that of Nautilus.
Calculation has shown that Nautilus can withstand a water pressure of 450m deep, and this is
also applicable for those Mesozoic ammoniods. But the descendant forms from them mostly
could not withstand a pressure more than I OOm deep water. Some ceratid ammonoids
occasionally show overgrowth of oysters on their shells. Considering the weight and size of
these oysters, it has been suggested that _in order to retain neutral bouyancy such shell might
have initially a large quantity of camera) liquid within their shell which the animal was able to
remove to counteract the weight of the oysters, and this may indicate that the animal might
have functioned in the same manner like the recent Nautilus.
B. Septal fluting
What could be the extra-advantage of strong septa) fluting found in many ammonoid shells?
Unfortunately, we have no living cephalopod possessing fluted septa. Thus our interpretation
on their probable function is somewhat conjectural.
A seeming advantage of crumpling of septa) margin is their buttressing effect which might
have increased the shell-strength so that it was able to prevent explosion at depth. Calculation
of 'Relative Stre11gtl, J11dex' of siphuncle tube (h/r x 100, where h = wall thickness and r =
radius of siphuncle tube) in living Nautilus and fossil ammonoids has revealed that an
ammonoid shell was in general, weaker than that of a Nawilus and it is difficult for such shells
00
Lytl.'Ki-ratiJ11 Ammomtida i>h,•llocc_mt ida
-
Danian
Maastrichtian
J
Campanian
-
Santonian
-
Coniacian .
"':,0
-Turonian --
w
~
8
~
Cenomanian
- Albian
--
II I tlj ·) :I Lt!U
Aptian
- ~-,LU ~l-~ ~f. -.··-- . -u~.J
. --~
Berrcmillll ~
HnutcrivitUl 1, -- .........
Valanttian
·--------.._... _.,,,_._..... __ ._, .........
\
Rcrriasian
. . 4-. '·l
.t~ .....
Portlandian
Kimmcridginn
Oxfordian
Callovi•_n I
UBI§
--u_. ····.
:· ::;; ····-• •.
UnthQflilll\
uuu,, - I •• -·· ••
t' ~~ .!~~~:::·.:=·-·:_·::.-~-~-~- - _.ti.I~==

~ -
.._ ,uocilUl UU\.
2 - --~-
- ..air--,--
a111i:1uan
1oarci~
1>Bcnsliac1an
Sincmurian
,,.~m~ . I
Hcltan1ian
-
ij~\L~.1~~ ~~ti:}~;1~~~J.. . ---~J
Rhactian
Norian
~
Cw
LadiIi~~·
-- 11 s:
~
~
-
Ani
Scythian
~
FIG. 15. 6: EVOLUTIONARY PATTERN OF AMMONOID IN MESOZOIC (after Moore in Treatise
5
C"l
part-LI
"<
CEPHALOPODA
189
to withstand a hydrostatic pressure at a depth more than I OOm. It is reasonable to say that
Mesozoic ammonoids probably compensated their relative shell weakness by developing fluted
septa. Moreover ammonoids were able to develop variable shell-forms in addition to tht! normal
ovate type (as that of Nautilus). This model has been accepted by Raup and Standy ( 1978).
But more and more fluting of septa beyond a certain limit might have decreased the angle
between the septum and the shell causing decrease of shell strength and such shells became
more prone to damage than others. Many other ideas are put forward as regard the function of
fluted septa but most of them are difficult to prove.
Bayer ( 1977) maintained the view that fluting of septa arose through geometric necessary.
A septum was initially formed as an organic membrane by accretion of materials from apical
mantle and even pressure from either side caused fluting of septal margin. When this septum
became mineralized it retained this fluting. If this is correct, then strengthening of shell and
other possible functions of septa become secondary.
15.6 MORPHOLOGICAL SPECIALIZATION IN RESPONSE TO LIFE HABIT

Though the major groups of cephalopods are now extinct one can have an idea of their life
habit from the observation of the same in those forms living today including the Nautilus which
ha s a long geological range. Cephalopods are all marine and nektic animals. The two
environmental stresses with which such animals have to deal are hydrodynamic effect (resistance
given by the fluid on the moving animals) and hydrostatic effect (i.e. enormous water pressure
borne by the animal at depth). Hydrodynamic effect of water has been dealt by the animal by
developing a streamlined shell-structure as those of belemnoids, nautiloids and many ammonoids
for ease of movement through water. They tackled the hydrostatic effect by increasing the
strength of the shell developing fluted septa and changing the specific gravity in water by
controlling the amount of camera! liquid within the chambers, just like a submarine is changing
its weight in water by flooding and emptying the water tanks. In ornamented shells of
ammonoids ribbing may also have had a role in strengthening the shell as well it might have
reduced the water resistance. Some ribs and spines probably acted like ship stabilizers to prevent
pinching and rolling. Like many other marin~ animals, .some cep~alopod shells are c~mouflag~d.
Surface of a Nautilus shell exhibit colour-strips on their dorsal side but the ventral side remains
uncoloured. This has been present to diffuse the shell outline. Many knob-like tubercles or
extended spines of some ammonoid shells mig.ht have the s~":1e function. ~nother problem
faced by the swimmers is to maintain the body m a stable position: F~r uncoiled ~eph~lopods
the load of soft part at anterior is counter-weighted by camera! fluid m the opposite side and
thi s is also true for coiled cephalopods where there is always a tug-of-war between the force
of gravity and force of buoyancy, both acting on shell at different .cen~r~s but in two o~~osi~e
directions. This is a very successful solution to the problem of mamtammg correct pos1t1on m
water in condition of movement or rest.
1.5.7 EVOLUTION OF AMMONOIDEA (Fig. 15-6, 15-7)

A. Ancestry
The earliest ammonoids belong to ancestral stock anarcestids which are of Lower Devonian
age. Many palaeontologists have considered that they were evolved from bactritids, a group
of ~traight shelled nautilids with bulbous protoconch, small marginal siphuncle and a long living
PALAEONTOLOGY
110
Ophioconc Arnmoniticone
(lyroucruticonc
(Kyniwnc)
( · ·rn111hl\ •1rhl1;ori •
l\nw 111 ·011 -·
(1111h '-' our) .
( CVflllCIIIIC)
(ll) '11/\NOE OF COILING PATTERN
;\11 ·nu,c Orthoccratitic (nautiloid) Palaeozoic
Nnulililic (nautiloid) Ord-Rec

'
Goniatitic (mnmonoicl) Carb-Pcrm
Ccrntitic (ammonoill) Trias
Anuuonilic (nmmonoid) Jura-Cret
(h) CHANGE OF SUTURAL PATTERN
Snmolh Costntc Tubcrculatc Spinose
(c) CHANGE Qr, ORNAMENTATION
Anc_,·locerus
8uc111/tc1 Ham1t11s Scaphites
Turrlllttu
(d) COMMON HETEROMORPHS Or: CRETACEOUS
FIG. 15 - 7 : EVOLUTION OF AMMONnrn~
CEPHALOPODA 191
chamber similar to early goniatids. The sutures of both the forms are also slightly wavy. Many
palaeontologists thus like to put 'bactrit ids' within earliest ammonoids.
B. Phylogeny
After origin, ammonoids got diversified, underwent various types of modification. Besides
the primitive group anarcestids, appeared a second minor group in Late Palaeozoic namely
clymenids with marginal siphuncle and goniatitic suture. Anarcestids soon led to goniatids with
typical coiled shells, zigzag septa and prosiphonate siphuncle. They attained their climax at
Carboniferous-Permian but became extinct at the end of Palaeozoic.
With the beginning of Triassic, the few Permian surviors gave rise to vastly successful
ceratids exhibiting complex shell ornamentation and sutural pattern. They remained dominant
throughout the Triassic but ultimately failed to cross the Triassic-Jurassic boundary.
Two ancestral ammonoid stocks phylloceratids and lytoceratids arose in Early Triassic, but
remained almost in a subdued stage so long ceratids were dominant. With the end of ceratid at
the Triassic-Jurassic boundary, they took the position and ultimately became the most dominating
marine invertabrates of the Mesozoic sea. These two groups are the ancestors to all other
Jurassic-Cretaceous ammonites. The history of these two orders is peculiar on account of their
stratigraphical persistence (since both continued throughout the Mesozoic) and their remarkable
conservatism. Within the two groups there was little evolution but each of their radiating lineages
or evolutionary offshoots became very much diverse. The phylogeny of these forms are so
complex that in many cases it is difficult to reconstruct them. Thus many palaeontologists like
to retain the name 'ammonitida' as a polypliyletic order (considering their origin from more
than one ancestral stocks (phylloceratida and lytoceratida).
The pattern of evolution illustrated by ammonoids especially in Mesozoic times has been
described as an example of iterative evolution (Fig. 15-6) in which the two long-ranged
ancestral stocks (here phylloceratids and lytoceratids) from time to time gave rise to short-
lived superfamilies and families which replaced each other in time. Each of the offshoots
dominated the scene for some time and .during this small period it constitute a miniature
adaptive radiation, producing several short-lived families so that each exhibiting a phylogenetic
tree like the prongs of a toasting fork. After this they became extinct and the space vacated
by them was occupied by the members of a new offshoot rising from the same ancestral stock.
Such a pattern of evolution is also described by the term 'palaeontological relay'. Because of
the rapidity of turnover, wide dispersal, abudance and ease of recognition, most ammonoids
become outstanding index fossils.
C. Phenotypic trends
d
''
A careful study of ammonoid phylogeny clearly indicates that within the entire group some
II
I I
morphological characters of the shell gradu_ally change with the time for various reasons. The
: l pattern of such morphological modifications are as follows :
' '
i: a. Change of shells from simple conical, uncoiled forms to more tightly coiled forms.
i I
I\ b. Change of sutural pattern from nautilitic to highly complex ammonitic type.
[~
t ' c. Gradual complexity of shell ornamentation.
'. t
: 1. d. Gradual increase of average size.
Palae(Gco)WP-25
192
a. Change of coiling nnd shell form (Fig. 1.~-711) l

Earliest bactritid ammonoids like th• ;arlicsl 111111lilldi. •,d1ihil d 11 Ill i1dy 11111 1' 1•I 1
11111 111
'"
· to ortlwcoue.,· of· 11aut1·1·tl1s 111· 'N' N· 11 • II ~., •.,,.L, •11llrd /Jm'll'lfl,•,mt
shell. Correspondmg ·
1
ti , '" tl11 11 I
, • ' k ' •l':ff"/,(/t fl;lf/l'IJl/fl ((II mll/ltJ/'flll#)
'I 1
stage appeared a slightly curved shcll ~lu1111 11ow11 !IN u, & 11
· · ·· .·, , ' · I1 ,l'i 1,ti lhl1 111111111111111 1 I c1 l1111,1t1u ltt

Ultimately m Late Devonian, H1s a11rI ·111t.1ss1
Carho111fcn:. 1 ·
goniatids and ceratids exhibited cuil ·d shells. At th· b ·~lt111h1~ 111' ud, 1111 'l tlf'l. '. '. 1111 11 1' With
1
loose and the whorls were nol tuuching. Su ·h f11n11 ·· 111·1· ·idled /,()ll'm;,,,wtlt'O IINt , I 1'111 1rlill'd
forms of ammonoids are known as ammouiticoue.,· (both illVllhtlc 111111 L'11l1vc,l111,•) 11!111 div,' lli 11il
in Mesozoic. It is seen that at certain periods of gcologi ·al hi llll'Y , 11111,· f!l'Ollp • of li1tt11101111 d 1
evolved shells of highly aberrant forms. Such . ~hells arc k11ow11 11 ' lwh.1t,m1,1111lt,t <I 11 , 11 ,d,
7d). Some of them were separated in Late Triassic among · •,uf idN., ll f •w f 01'111 11 f1Pl'llt d ''
Jurassic and more extensively they appeared in Late Crctuc 0111-1 Pcn11d .
Heteromorphs may be loosely coiled with later whui"ls compl '1 "IY 01 pml ly , ·11111'111 ·d, , '11111'
are almost straight-shelled. But unlike straight or partly <.:oiled 111111tlluid th p1'c1to •111H·h ·fr
remained always tightly coiled. Some common heter11101·phs 11l'c S, ·a1,1tl1,,,,. (i11l1l11lly '111 1•d, 111 I
whorl uncoiled), Spiroceras (coiled but whorl not touching), /Jan,/it,•.,· (1qn·l1d11 lll1 ·o I cl ,11,~fl,,
Hamites (hook-shaped) etc. The Triassic Coc:hlocera.\·, und lhc ·,· 'li.H.! cou O.v1//11,,rm·,1rrM,
Turri/ires and a few other genera were helically coiled like a gusft'or,od . Th ('1·,·uh.!1•1111
Heteroceras and Hypantoceras exhibited a mixed type of coiling.
Existence of such heteromorphs has given rise to very much evolul i1111ury spcc.:ulul iuu . I or
a long time it was thought that heteromorphs were relrogressive forms. Thal L"I lightly ~ol l •J
ammonoids in Late Cretaceous due lo reverse or retrogressive evolulion dcv ·lop ·d purlly t.:ol fcd
and uncoiled forms again like their ancestors. This idea has been disl.:unJ ~d Inter r-ii 11 • • •volu1im1
is considered as an irreversible process (Dollo's law). Many have sugg ·sl ~d I.hat h •1crom111·ph
were degenerated, biologically inadaptive forms. Furthermore, because ~rl'iussi<.: 11t1d ·r~,u~ ·ouN
episodes of heteromorphy took place shortly before two majnr periods of cxti11ctio11 fof' lh
ammonoids (ceratids in Triassic and ammonites in Cretaceous) it seems tn muny rulu ·011tolOBiNtH
that there was definite some relationship between heteromorphy and extinction of u111tnonoidN.
The view was that the rapid evolution of ammonoids eventually culminated in a kind of 'racial
senescence' during which bizarre and overspecialized forms developed us last and final pn>uuclt'
before evitable extinction of the entire group.
First of all one cannot considered these heteromorphs us <legcncrutcd unu ovcrspccinlizcd
forms, because in the history of ammonoids they were present ulmost throughout with lcs 11er
or greater abundance. Extinction of ceratids at the end of Triassic or ammonites nl the end of'
Cretaceous was not selective. That is if these hctcromorphs would he 1101rnduptivc dc!(encnuing
forms, then there is no reasons of extinction of normal coiled forms ut the same titnc, ft is
also noted that some Late Cretaceous heterornorphs again produced descendants with normul
or near normal coiled sheJls and hence they cannot be considered non-adaptive or ov -'rNpccinfizcd
forms. Their ov~rall ~~rphol?gy might have indicated that most of them were sp--cializ •J for
a b?ttom dwell mg .life h.ab1t. It appears that they probably cvol ved ;11 Ulh•i>tntion lo u11
env~ronmental domain which was vacuted in Triussic times hy the cxtiuctiou of the sirnilM
straight-shelled Palaeozoic nautiloids.
·1; nu1. )l>Ot >A 193
h. Suturul pnttcrn {Fig. 15~7b)
Earli~st nmmnnoids ·volvl·d from nautilids also exhibited a wavy 11autilitic suture. But soon
thcs' sutlm~ li1r•s h"came romprcss"ct and both lobes and saddles became v-shaped. This is
th' <..'haral'I •risfr·s of Lat' Pnh1 '07.oic goniatids and is known as go11iatitic suture.
In Triassi · gt>niatitic suturl' was r •placed by ceratitic suture where lobes became crenulated
m~d _the s:~ddks r •nminlld round~d. In Jurnssic-Cretaceous a more complex suture appeared
Wtthm typ ical ammonit ·s wh ·r·~ both lobes and saddle became highly frilled. Such a suture is
call •ct ammo11itic s11t11re.
c. Complexity of ornumcntution Fig. 15-7c)

Pr!mitivc gt>n •ra of ammonoids were usually with smooth unornamented shells. This stage
was tollowcd by a capil/ate stage showing a pattern of fine raised radial lines on the shell
surfoc . This was successively followed by subcostate and costate stage in which more coarse
radial stria · called ribs or costae developed. Towards Jurassic and Cretaceous some costate shells
produc •d tub -•rcles and spines. Shell became progressively stronger with more complexity of
ornament at ion.
d. Increase of a,1cruge size

It is also noted that the average size of ammonoids increases with time. The Palaeozoic forms
mostly had the size of a golf ball . Saucer-sized ammonoids were common during Triassic.
Jurassic ammonoid's average size was still larger. Gigantic ammonoids were present in
Cretaceous including some discoidal forms greater than 2m diameter. Many smaller forms
however continued upto the end of Cretaceous.
15.8 PROBABLE CAUSES OF EXTINCTION OF AMMONOIDS

Extinction of ammonoids at the end of Cretaceous was rather an abrupt event because they
became extinct at their climax. Various reasons were put forward to explain such event. Some
call up intrinsic causes; others solicit change in the environment caused by one or more physical
events. The former workers have invoked essentially a sort of adaptational potentialities. They
have referred to many aberrant forms within Cretaceous ammonoids such as uncoiled, partly
coiled, helically coiled and gigantic forms as degenerating condition of the ammonoids before
their fine extinction. But the point is that such aberrant forms within ammonoids were found
throughout the Mesozoic, right from Triassic and none of the features had affected the whole
group at a time and their extinction was also non-selective. Moreover, Cretaceous was a period
of mass extinction when many other animal/plant groups including phytoplanktons, many
gymnosperms, all gigantic reptiles also became extinct and this strongly suggest some external
causes which might have had affected in some way the physicochemical or biotic environment.
The probable events could be as follows :
(a) Large scale volcanism or meteorite collision resulted in the accumulut.ion of huge
amount of dust in atmosphere preventing sunlight to penetrate and reach the ennh-
surface. For want of sunlight phytoplanktons and many other plant groups becam •
extinct. Phytoplanktons are linked with many animals in the marine food chain. So the
marine ecosystem was shuttert!d leading to cx1inction of ummonoids and other related
forms.
PALAEONTOLOcy
194
Cretaceous caused rise of mountains
·
(b) L arge scale orogenests towa rds the en °
d f ·
. regression and transgress10n an d many'
. 'b . ·
re d 1stn ut1on of continents an d oceans ' marine
other catastrophic events.
15.9 GEOLOGICAL HISTORY (Fig. 12-3) f ephalopods appeared first in Late

0
N;mtiloids are considered the most oldest form~ c d mostly exhibited straight or partly
Ca.mbrian. By Ordovician-Devonian they became abun ant~~ families in Palaeozoic period but
cmled shells. Altogether, they were represen~ed ?Y about Their radual decline continued
most of them failed to cross the Permo-Tnass1c boundary. g d b . 1
throughout the Mesozoic and at present, the whole group is represent.e T ~ a. smg e genus
· ·t ppearance m nass1c
Nautilus which however, the only successful form since l s a ·
The appearance of first ammonoid was found in Ordovician represent~d ?Y Eobactrites w~ich
however exhibited somewhats intermediate characters between nauttlmds and ammonmds.
Ammon~ids began to increase in number in Devonian with the appearance of goniatids. But
actual outburst took place after Triassic with the appearance of ceratids. Triassic is sometimes
called age of ceratites. After Triassic with the falls of ceratids, ammonites took their position
and became the most dominant marine invertebrate of Jurassic-Cretaceous sea. They exhibited
variable types of shell-forms and size during this period which may be sometimes referred to
as age of ammonites. Surprisingly , such an abundant group ultimately failed to cross
Cretaceous-Tertiary boundary and all forms abruptly became extinct at the end of Cretaceous.
Within coeloid group, belemnoids appeared for the first time in Carboniferous, attained the
climax in Jurassic-Cretaceous times and became extinct at the end of Cretaceous. Other forms
mostly appeared in Mesozoic and are still continuing.
15.10 STRATIGRAPHIC USE

Among the cephalop~ds, am~onoids_ are the prime stratigraphic tools used for the Mesozoic
rocks of the world. Rapid evolut10n of iterative type and quick h" d'
their nektonic life-habit together have made this fossil group 1.d gle~grab~ ic •_spersa~ he1pe~ by
· h. · ea ior 1ostrat1graph1c zonation
stratlgrap 1c corre1at1on and age determination of marine M · '
ammonoid genera and species are widely used index foss"l eto:oic rocks of the world. Many
0
index fossils of Carboniferous are Gastrioceras liste . c's . ti e world. Some of the common
n, onw oboceras go · L b X ·
carbonaria, which are mostly goniatids. The Marine T . . moo us, enasp1s
the Himalaya has been zonated mainly on the bas· fnassic r~ck-succession of Spiti basin of
. . . f . . is o ammomte fos ·1 f d .
Tnass1c mdex oss1ls belonging mostly to cerafd , . s1 s oun m them. Some
• ,,. . l s are . Otocera O h.
Hungantes, , ropltes subbulatus etc. The importa t M"ddl I
. s, P .,ceras, Meckoceras,
macrocepha Lus, Aspu· Joceras, Rhineckia etc The n . e Jurass1
· c forms are Macrocephalites
. · .. · se a1e mostly · . .
Cretaceous ammono1ds are Cnoceras, Ptyclw . , . . . ammonites. The mdex fossils of
. S c1;1as, Hanutes Lt
Bacuiltes, capIutes
.
etc. a ematus, Turri lites costatus,
Chapter 16
ARTHROPODA
16.1 INTRODUCTION
Arthr~poda (Gk:-arthron : joint; podus : limbs) includes an enormous group of invertebrate~
charactenzed by b1lateral symmetry, segmented body, paired and jointed appe·ndages used for
locomotion and feeding. Commonly they possess chitinous exoskeletons which is moulted
peroidically to allow the growth of animals. The animals are of tripJoblasticcoelomate grade
with metamerous body plan having well developed internal systems (digestive, circulatory,
reproductive and nervous system). They include a vast group of insects, crustaceans, and spiders
as well as some extinct groups like trilobites -and eurypteriods (giant Palaeozoic water-
scorpions). (Fig. 16-1 ).
Arthopods resemble annelids in terms of their segmented body with which they possibly
share a common ancestry. However, in the possession of hard skeleton and in terms of
mechanism of movement, growth and feeding, arthropods have an overriding superiority over
annelids. In structural complexity, adaptation almost to all sorts environments and development
of remarkable social organization among some groups, the arthopods seem to represent their
climax of evolutionary advancement. Most of them possess well equipped defensive organs.
They are also habituated to all sorts of animals and plants as their food.
16.2 SUBDIVISIONS
Problems related to classification of arthropods remain unsolved. Many different taxonomic
schemes have been erected for arthropods. Some consider the group as polypliyletic (arose from
multiple ancestors). Manton (I 973, 77) considered 'Arthropoda' as a superphylum and
subdivided it into a number of phyla of which one it 'Trilobita'. The author has followed the
scheme proposed by Moore & others (1959}. The broad subdivisions are as follows :
Phylum Arthropoda : True jointed-limbs bearing invertebrates.
Subphylum Trilobitomorpha (Cam-Perm.) : Trilobites and related forms; mostly aquatic,
marine; exclusively Palaeozoic; e.g. trilobites.
Subphylum Chelicerata (Cam.-Rec.) : Lacks antennae; presence of some appendages
bearing pincers (chelae); includes eurypterid (merostomes), arachnids,
scorpions, kingcrabs etc.
Subphylum Crustacea (Ord.-Rec.) : Mostly aquatic; possession of antennae and some
biramous limbs; includes crabs, lobsters, shrimps. etc.
Subphylum Myriapoda (Sil.-Rec.) : Elongated, annelid-like; numerous segments, each
with a pair of locomotive appendanges; includes centipedes, multipedes etc.
Subphylum Insecta (Dev.-Rec.) : 3 pairs of limbs and a pair of wings; terrestrial or
aquatic e.g. Insects of different types.
195
-
,C
::Jo.
~WJmulm~
e
i
f s:
t'?']
FIG. 16-1: SOME REPRESENTATIVES OF ARTHROPODS 0
<:
(a) TRILOBITES (b) SHRJMP (c) SPIDER
~
l
(d) EURYPTERJD (e) CENTIPEDES (t) INSECT
§
~-.~- ··~"" ·i·~·- ,. • ,,.___ - - - - ---~- - - -•L __... _ . ___ - ---• ---• --L-- · - - - -- - - --- -- -
ARTHROPODA 197
- - - - - - - - - - - A r t i c u l a t i n g Furrow
::..---Waxy Layer
" - - - - Rigid Calcified
======~ Chitinous Layer
Flexible
Chitinous Layer
" - - - Epidermis
(a) LAYERED STRUCI'URE OF ARTHROPOD EXOSKELETON
Swimming Leg Of
Aquatic Arthropod Biramous Appendage
Walking Leg Of Of Trilobites
Land Arthropod Chda Of Lobster
(b) APENDAGES OF ARTHROPOD
FIG. 16 - 2 : SKELETAL STRUCTURES AND APPENDAGES

Width
Anterior .
I
I
Cephalon , ~ - - - - Axial Lobe
Length-+ Thorax
- - - - - - Trace Of Bilateral Symmetry Plaie
L--.\--+-i-/~1-::::::::a--- Axial rurniw
Pygidium 'r
I
------~------~~~~~
' (a)
r··
5 , GlabelJa
Facial Suture--,;----..61 ''*'""--Anterio-Lateral 1..: Eye
Cheek---r-'71 Border Ir I
U ! Cheek
Glabclla,--~Q~~'--.-t~ Eye I ....
Thorax
198 PALAEONTOLOGY
16.3 GENERAL FEATURES OF ARTHROPODS (Fig. 16-2a, b)
Exoskeleton of the arthopod is generally called carapace. The segmented body is bilaterall
symmetrical and the segments are called somites or metameres. Exoskeleton, secreted by th~
epidermis (outer skin) exhibits a layered structure : an outer most waxy. waterproof layer and
a thick inner layers of calcified chitin whose outer part ?eco~es tough but •~ner part is somewhat
tlexible or elastic. The rigid outer layer is absent in articulating zones makmg the hard segments
movable over each other along the joints. This rigid covering not only serves for protection
but also provides places of attachment of muscles, the activity of which causes the movement
of appendages.
The hardened part of the skeleton of each segment, called sclerite is consisting of a dorsal
tergite and a ventral sternite that make a hard ring for each somite. In some groups there are
also two lateral parts of segment called pleurites. The somites may be grouped according to
their placement as head, thorax and abdomen or tail. In many groups, head and thorax are fused
together forming a cephalothorax.
Normally, each segment bears, a pair of jointed appendages, attached to the side of each
somite at junction of sclerite and tergite by muscJes. The number, shape and relative size of
appe~ndages vary greatly. The more common types attached to head are : a pair of slender and
long sensory structures, antennae; short jaw like structures, mandibles; and long or short .limb-
like structures, maxillae used for capturing food and passing it to mouth. Thoracic segments
are mostly provided with legs used for locomotion. The number of appendages is varying from
numerous (centipedes) to a minimum of six for insects. Some arthropods developed a pair of
pincer bearing appendage called chela as found in lobster and crab. Swimming and aquatic
arthropods, develop bristles on their legs. In trilobites and crustaceans each leg appendage
becomes two branched; an outer exopodite and an inner endopodite, both attached with a basal
segment coxa that remains articulated with the somite.
Arthropods usually adopt a sexual means for reproduction. They develop from eggs. In some
groups after the egg hatches, it passes through a number of stages of development before attaining
the adult form. This phenomenon is called metamorphosis. This is well seen in case of insects.
But after attaining maturity it continues to increase in size, so long the life lasts. The increase of
size and/or change of shape require successive renovation of the exoskeleton because the old
skeleton becomes too rigid to accommodate the growing animal. Hence the older skeleton is
discarded, whenever the animal finds it unsuitable to accommodate the body. This growth pattern
in called moultillx or ecdy.vi,v. This is soon followed by secretion of a new skeleton. for
palaeontologists this mechanism of growt.h creates a serious problem as a single animal throughout
its life-time, in this way, trwy give rii,c to a large number of skeletal-remains each may be
morphologically different from other und may he fossilised separately.
'16.4 'J'IULODl'fK ' Gfi:NJ•:HAL CIIAHAC'l'RRS (Fig. 16-3a-c) ·

. Trilohit ·s ar · ·xti11 ·t Pula ·cw iic .,.rou p ol' urthoprn..ls exclusively marine (shallow). in ~a~t
The num · re f ·r t< , it , P<1 c · iou of' 1111· ·e di Ii net lubes in the body placed both lon~itud•:~
and tn111sv ·n ·ly. Tiu; f hn: ' tra 11 1v ·r w lolw ar. : nntcriorly placed cephaloq, median thaxial
und pontcr'iorlY plu,, ·d /IYP. t/111111 , <JI' t 111~ th1·1·t· l1111~it udinal lobes median one is called ted
und otl, ·r tw<> o11 ni 1J1cr . ti .. ,f' s11d11I ,., • lawml ur /Jlcmral lobes. Lateral lobes are separa
ARTHROPODA 199.
from ~he axial lo~s by .two ~rrows (axial furrows). Each lobe contains one to many segments.
The size 0 .f the animal as v.an.abte ranging from small (10 mm) t.o as large as 60 cm. Although,
totally extinct, close assoc1at1on of fossil trilobites with other known marine invertebrate such
as brachiopods and corals definitely indicates their marine habitat. ·
16.4.l Morphological features of Trilobites

A. Head shield or Cephalon (Fig. 16-4-16-5).
Cephalon or head shield Anterior most part of the three transverse lobes;
Border Raised peripheral part _of cephalon and pygidium.
Marginal furrow Groove or abrupt inflection of surface a'long .the inne~ edge
of border. ·
Glabella Cephalic portion of axial lobe bounded an.cl separated from
(Fig. 16-4a,b; 16-5a,b) the rest head by axial furrow; It may be unsegmented-or
fully or partly segmented (5-6) (glabellar segments).
separated from one another by transverse furro'_V~
(glabellar furrows); glabbella may touch the anterior
border of cephalon or may be short of it; it may extend
beyond the anterior margin of head; may be anteriorly
narrow or wide or abruptly truncated.
Occipital segment and furrow : Posteriormost glabellar segment in front of tJ)or~~~11d the
transverse glabellar (urrow in front of it . .:,.· · .-_. .''~·...~ .
• .. . .". !'./
Occipital node A tubercle on mid-portion of occiptal segrrie.rii. : .-

Cheek Portion of cephalon minus glabella occurring on either
side of it; may be divisible into two parts by a suture
(facial suture)
Fixed cheeks or Fixigena The inner portion of each check fixed with glabella.
Free cheek or Librigena Outer part of each cheek including its extension on ventral
side; this portion is movable on fixed cheek along suture .
line that separating it fr:om fixed cheek.
Cranidium Glabella and _fixed cheek together form the cranidium.
Genal angle Angle subtended by anterior-lateral and posterior margin
of cephalon often extended posteriorly as genal spines.
Neck furrow Transverse posterior groove of cephalon separating it from
thorax.
Eyes (Fig. 16-4b, 16-5c) Visual area, usually a pair, located on either side of
glabella on fixigene near the suture line between free
cheek and fixed cheek; eyes may be single or multiple-
lensed; small or large; sometimes lying on a stalk-like
1. , body; eyes may be totally absent in some groups.
Palae(Geo)WP-26
., ' PALAEONTOLOGY
.----Occipital Funow
1...----0ccipital Segment
. - - - Gcnal Angle
(a) DORSAL VIEW

Rostral Suture
.---Rostram
- - - Free Cheek
Hypos10111i.:
Facial Suture
~-+."""",n-- Hyposlomal Sutwe
Cnmidium
(b) DORSAL VIEW
(c) CEPHALIC SUTURES IN VENTRAL VIEW
~~~
~~~
Protoparian Proparian Gonatoparian Opisthoparian Hypoparian
(d) FACIAL SIJfURE
1------ Ccphal ic Spine
,___ _ Genal Spine
(e) SPINES IN CEPLALON
FIG. 16 - 4 : MORPHOLOGY OF CEPHALON
ARTHROPODA
201
Palpebral lobe Raised lobe-like feature on fixed cheek along inner edge
of each eye surrounded by a depression (palpebral
fu"ows).
Doublure Reflexed portion of carapace along ventral margin of
cephalon.
Hypostome A plate on ventral side of cephalon in front of mouth.
Rostrum A small shield-like elevated part in front of hypostome.
B. Cephalic sutures These are some weak lines on the cephalic lobe of
(Fig. l 6-4b-d) trilobites marking the areas of fracturing of skeleton
during ecdysis process. These are of several types and
except facial suture, all are ventrally located. Those are
as follows.
(a) Facial sutures The two depressed lines on either side of glabella
separating the free cheek and fixed cheek on dorsal side
of cephalon. This may be again of five types.
(i) Protoparian Suture marginal to cephalon; cheek undivided forming the
entire cranidium; eyes present e.g. Olene/lus.
(ii) Proparian Suture originating from anterior and ending in front of
genal angle e.g. Dalmanites, Phacops.
(iii) Opisthoparian Suture like the proparian but ending posterior to genal
angle at posterior margin e.g. Jsotelus.
(iv) Gonatoparian Facial suture ending at genal angle e.g. Calymene.
(v) Hypoparian Marginal but on ventral side, without eyes e.g. Paraharpes.
(b) Rostral suture Ventral line separating the anteriorly located rostral plate
e.g. Calymene.
(c) Hypostomal suture Ventral line separating hypostome from rostrum or from
anterior part of cephalon. (in case rostrum absent) e.g.
Paradoxides.
(d) Connective sutures 1\vo ventrally situated depressed lines isolating hypostome
and rostrum; they may represent ventral extension of facial
sutures e.g. Calymene.
(e) Median suture Sometime the two connective sutures meet in front of
ventral side forming a single depressed line e.g. /sol<:1us.
C. Thorax (Fig. 16-6a-d) Midd~e portion of the three transverse lobes.
Thoracic segments Transverse divisions of thorax, each composed of a centr.\l

axial segment and two lateral pleural segments (plturom)
on either side of each axial segment. Thoracic and pleum\
segments are movable along the joints. .
202 PALAEONTOLOGY
Partiallty Segmented Unsegmented
A cerill B
L.S. through Glabella
(a) SEGMENTATION OF GLABELLA
Anteriorly Wide Touching

®·
::,: '• ',
·.··:...
Behind
Anteriorly Narrow Anteriorly Lobed
Anterior Marain Anterior Margin
(b) SHAPE & SIZE
~ - - Facial Suture
~ - - Eye
11.-Wl-+t:----- Palpcbral Lobe
Eyes Very
Large Compound Eyes On
Small (Simple) Stalk
Eyes
(c) MORPHOLOGY OF EYES
FIG. 16-5 : MORPHOLOGY OF GLABELLA AND EYE
ARTHROPODA 203
Articulating furrow Transverse groove between two adjacent thoracic segments
which also marks the location of movable joints.
Pleural furrow A depression on side of each pleuron which curves
backward and outward from the edge of each pleuron. It
is interpreted as trace of primary segmentation.
Articulating half
segmenUring An arched semicircular area in front of each axial thoracic/
pygidial segment, covered under the reflexed border of the
preceding segment; the edges of these two elements are
connected by flexible unmineralized chitin so that one
segment becomes movable over the next segment.
Articulating facet Sharply downbent area along outer anterior edge of each
pleuron to allow for relative movement of segments.
Apodeme Inward ventral projection of each axial segment to which
attach the muscles causing articular movement of segments.
D. Biramous appendages On the venral side, the pair of appendages attached to each
(Fig. 16-6e) thoracic and pygidial segment functioning limbs; four pair
of similar appendages also present at ventral side of
cephalon (functioning as post-antenna} limbs); these are
all moved by muscles attached to a ventrally directed
pointed zone below each thoracic segment called
appendifer; each of these appendages in attached to
apodeme and has the following parts.
(i) Coxa The basal segment of each appendage which is articulated
with apodeme.
(ii) Endopodite Inside branch emerging from coxa acting as a limb;
composed of nearly six/seven segments (podomeres), the
last one forming one or more claws.
(iii) Exopodite Outer branch emerging from coxa," often called gill-
branch, telopodite, or outer ramus is composed of many
short segments; It bears a flattened shaft with closely
spaced bristles or setae along one margins; it is believed
to have been used for both respi_ration and swimming.
E. Pygidium (Fig. 16-7a-c) Posterior-most lobe of the body; may be segmented/

unsegmented or partially segmented; axial part may reach
the posterior border or may not. Size of pygidium is
variable and compared to the cephalon it may be of
following types.
(a) Micropygus Very small-sized pygidium, e.g. Paradoxides.
204 PALAEONTOLOGY
Articulating _ _ _ _.... Articulating Half-Ring

FurrC1w
---Articulaling Furrow
Apoderne
~:~:.,~l',Wi, ~
A Detached Thoracic Segment (Ventral View)
Articulating Facet
(a) DETACHED THORACIC SEGMEN/ ldorsal view)
Articulating Half-Ring
(b) ENROLLED SPECIMEN
Axial Spine
Pleural Spine - - - Thorax
(d) THORAX WITH TWO SEGMENTS

(c) MULTISEGMENTED THORAX
Dorsal Shield Appcndifer

Endopodile
Apodcmc
Exopoditc
At111chmcn1 Of Birnmous Appendage

- Endon~d ·
,..., tic
On Vcntrnl Side
(c) GENERALIZED DIAGRAM OF BIRAMOUS APPENDAGE
FIG . 16 - 6 : MORPHOLOGY OF THORAX
ARTHROPODA 205
(b) Heteropygus Relatively large but smaller than cephalon. e.g. Ca/ymene.
' (c) lsopygus Pygidium and Cephalon are almost of equal size. e.g.
l.wte/u.'i.
(d) Macropygus Pygidium larger than cephalon. e.g. Anisopyge.
Pygidial axial lobes
and furrows Axial segments of pygidium and the grooves occurring in
between them.
Pygidial pleural lobes
and furrows Segments of pleural part of pygidium and the furrows in
between the segments.
Articulating half ring
& Articulating facet Similar to those found in thoracic lobes.
F. Ornamental features The chief ornamental features of trilobites are the spines
(Fig. I 6-4e, I 6-6c, 16-7b) and in few cases also tubercles; spines may be present at
various parts of the skeleton. The chief spines are :
(a) Genal spine Posterior extension of genal angle.
(b) Cephalic spine Anterior spinose extension of cephalon along the long axis
(e.g. Ampyx). There may be a number of spines emerging
from cephalic margin e.g. Oloenelloides.
(c) Axial spines Rows of spines emerging from mid-line of each axial
segment of thorax and pygidium.
(d) Pleural spine Margin of each pleuron may be smooth, rounded or may
be spinose; each pleural spine may be short/large; even/
uneven-sized and generally curved backward. e.g.
Paradoxides.
(e) Marginal spines Sharp projections of margin of pygidium, may be two/
more in number. e.g. Cybele.
(f) Axial or Caudal spines Posterior extension of pygidial axis. e.g. Paedeumias.
(g) Telson One of the pygidial segment often extended posteriorly in
the form of a small or large spine. e.g. Redlichia.
16.4.2 Classifications
The classification given here mainly foHows the one in Treatise on lnvetrebrate Palaeontology
(Moore et. al. 1959). Former classifications _were mostly based on facial sutural pattern. But
this classification is based on the whole complex axial and other characters. But as this group
is comprising entirely of fossils, subdivisions are recognized based on characters visible in fossils
and whether these _phenetic groups actually reflect phylogenetic or genetically related categories
will remain a matter of debate. The board groups are as follows :
PALAEONTOLOGY
206
Unsegmented Pygidium
Segmented Pygidum
(a) SEGMENTATION
~!!~--Maginal
Spine
C . 11111.,1
(b) PYGIDIAL SPINES
Micropygus
lsopygus Macropygus
(c) RELATIVE SIZE OF PYGIDIUM & CEPHALON
FIG. 16 - 7 : MORPHOLOGY-OF PYGIDIUM (dorsal view)
.~:
@.~
I
I 2
(a) PROTASPID
(b) MERASPJD
(c) HOLASPID
FIG. 16 - 8 : ONTOGENIC STAGES OF TRILOBITES
ARTHROPODA
2m
Order l
Agnostida : (Cam.-Ord.) lsopygus, marginal suture, a few thoracic segment , blind.
e.g. Agnostus, Eodiscus.
Order 2 Redlichiida ·· .(L-M Cam) ce p ha Ion, very i,;ma ti pyg1'd'1um ( m1cropygus),
·
. · L·1rge
•
strong genal spmes, opisthoparian suture. e .g. Redlichia.
Order 3 Corynexochida ·· (Cum ·) He terogeneous
.. .
. . group, glabella variable Nhapcd, fiuture
op1sthopanan
. • thorax w'th . . . . h . .
• seven to e,g t segments, often m1cropygu s. e .g.
Ollenotdes.
Order 4
~ychopariida : (Cam.-Ord.) Largest order, large cephalon and pygidium, most ly
isopygus or heteropygus, suture mostly opisthoparian, thoracic segments variable.
It has five suborders. e.g. Ptychoparia, A:,aphus, /llaenus, Harpes.
Order 5 Phacopida: (_Ord.-Dev.) Large cephalon, heteropygus, eyes large, suture proparian
to gonatopanan, many thoracic segments. e.g. Phacops, Calymene.
Order 6 L~chida : (Ord.-Dev.) Opisthoparian suture, macropygus; leaf like pleurons. e.g.
Ltcas.
Order 7 Odontopleurida : (Cam.-Dev .) Highly spinose, suture opisthoparian, 8-1 O thoracic
segment, small pygidium. e.g. Acidaspis.
'
Order 8 Proetida : (Cam.-Perm.) Opisthoparian suture, large glabella, isopygus, ~1out gcnal
spine, 8-10 thoracic segments. e.g. Proettts, Phyllipsia.
16.4.3 Ecdysis and Ontogeny (Fig. 16-8)

Like all other arthropods, trilobites began its life from an egg from which a larva was
eventually hatched. From this larval stage the trilobite grew through periodic moulting. The
exoskeleton became disarticulated and broken,'llong the cephalic sutures during each moutling
process and the old skeleton was discarded as cast shell often called exuvia. The soft body of
the newly grown form soon secreted a new skeleton. Successive moulted exuviae are often
preserved as fossils and may give a record of successive ontogenic development from the earliest
larval stage. In this way, a single individual had produced a large number of skeletons in its
entire life. The different stages of ontogenic development of trilobites are as follows :
The earliest stage of this growth, protaspis was a small disc-like body carrying a segmented
central lobe (later transformed to glabella) with tiny eyes located anteriorly without any distinct
facial suture. Ma~y protaspic forms were spiny which later disappeared totally (indicating their
different mode of life, possibly planktic at this stage). The second stage of moulting is called
meraspis when developed a transverse furrow sep~rating cep~alon- from pres~1?1ptive pygidi.u":1.
When pygidium became free, thoracic segmentation started in front of pyg1dmm. Merac;p1s 1s
usually numbered as o, 1, 2, 3, etc. depending upo.n the numbers of thoracic ~g~nts ~ed
·t· ry pyg·i d'i um · When
· this segmentat10n became
. complete the tnlob1te attained
f rom t rans1 1ona , .
the adult stage called Jiolaspis which me.an~ ~ 'complete shield . Growth however, continued
by further ecdysis until the death of the ind1v1dual.
· t " ·ts of different ontogenic stages, often preserved along with the adult
F or paIaeontoIog1s , 1oss1 . .
. bl ·n recognizing a population whether belonging to one or several
forms cause serious pro em 1 . • • •
·' . f h three different ontogemc stages frequently exh1b1t different
species. As exuviae o t ese . .
PALAEONTOLOGY
morphology. and hence sometimes give u wrong impression of the presence of more than one
spe~ie~ wi thin a fossil po pulation.
16.4.4 Ecology
In ga neral. a ll tril obi tes were marine, basically crawlers on the shallow sea botton which is
evident from thei r nature of limbs (endopodites). Pilamcntal gill branches (exopodites) might
ha,-· hel ped them not only in respiration but also in partial swimming at least near the bottom
ne kt bcnth . . Palaeontologists have also tried al their best to interpret the function of the
ther morphologic features which could be matched with their life habit.
Eyes in trilobites may exhibit various degree of development or suppression. Blind trilobites
. metime·· interpreted as nocturnal and some consider them living in dark caves (within
,i.: .. re
reef. r at a greater depth of sea or even they might be burrowers. Such trilobites exhibiting
mooth arapace and wider thorax might also be burrowers. A form with very large eyes might
indi ate a degree of all-round vision and more probably it was a swimmer, swimming upside
d wn keeping a sharp look for preys or enemy predators. Some trilobites like trinucleids
exhibited a wide cephalic fringe surrounding the brim. This types of brim possibly helped the
anima l to align themselves towards some prevailing current which may imply that they were
tilter feeders.
Spines in trilobites are common features and it is asserted that besides protective function
they also acted as baffles to increase water resistance and helped to prevent their sinking within
sedi ment. Spines on ventral surface probably acted as support of body. Large genal spine might
have had supported the entire skeleton, to rise the skeleton above the sediment level.
Some trilobites of Ordovician often developed a highly convex exoskeleton with thorax
harply bending from occipital ring. Such a morphology is unsuitable for epifaunal crawlers or
wimmers and poss ibly they were largely sedentary suspension feeders lying partially buried
thorax and pygidium) within sediment.
J6.4.5 Trace fossils related to Trilobites

Trace fossils believed to have been produced by different trilobites are found within many
Palaeozoic sediments, though, there is still some uncertainity about their origin. Even, it is not
possible to say that a particular structure is related to a particular species or genus. Some of
the most common trace fossils are :
(a) Cruziana These are bilobed chevron-marked trails indicating traces of rapid
c~capc or crawling, or even burrowing movements .
.
(b) Ru ophycus Resting nest, burrows or surface exuviarion-murks created by trilobites.
(c) Protichnires Spot-like walking-i mpressions m.1rking movement of individual limb.
All the movement impressions arc generally classified as repid,inia and the resting
imprcs~ion as cubicl1i11/a .
J6.4.6 Stratigraphic uses

Trilobites are of considerable importance us regards the zonntion of Cambrian-Ordovician
rocks and are succe. sfully used as good zon~ fossils in muny countries. Almost everywhere in
:::,..
;:,;
~
~
,:
0
-
..C,
~ s
CAMBRIAN ORDOVJCJON SILURIAN DEVONIAN CARBO~IFEROUS PERMJAJ,;
REDUCHIIDA
-
,.
CORYNEXOCHIDA ,.
AGNOSTIDA -
,.
PTYCHOPARJIDA ,.-
-
~
ODONTOPLEUIDA
......
PROETIDA ,--
.....
PHACOPIDA
-
LYCHJDA -
,.
N
FJG. 16-9 : GEOLOGICAL RANGE OF TRILOBITES (orders)
~
210 PALAEONTOLOGY
marine sequence their first appearance usually murks the beginning of Lower Cambrian. Many
trilobites also attained a widespread geographic dispersal (possibly happened for its mobile
larvae). Some imponant index fossils are Redlichia rwet/ingi, (Lower Cambrian of Asia and
Australia), Paradoxide.,· pim,s (Middle Cambrian of Europ~ and North America), Olenoide.r
(Middle Cambrian of North America) etc.
16.4.7 Ancestry and Geological history (Fig. 16-9)

Trilobites appeared at the very beginning of Cambrian and this causes difficulty for
palaeontologists to make any suggestion about their origin and ancestry. However, many
considered that arthopods evolved from some Precambrian annelids and this root-form is named
arcl,etypa/. Larval protaspis stage of trilobites is similar to some larvae of annelids but it has
no similarity with larvae of other arthropods except arachnids. From this observation, many
have suggested the origin of trilobites from some group of primitive arachnids or both might
have evolved from a common annelid ancestor. Arising from some annelid ancestry, trilobites
appeared for the first time in Lower Cambrian marine beds and very soon became the most
dominant animals of that period. This appears to some palaeontologists somewhat anamolous
and they considered that the group might have existed before Cambrian in Proterozoic times.
Absence of their fossils in Proterozoic rocks is possibly due to the lack of hardparts. However,
Cambrian-Ordovician may be considered as the climax of this animals where they are found
mainly associated with inarticulate and primitive articulate brachiopods. They became fewer in
number after Ordovician. Only 10-12 genera were recorded in the Carboniferous-Permain times.
They became totally extinct at the end of Permian, possibly failed to compete with the incoming
fishes and large cephalopods both of which made their first appearance in Ordovician.
I
1
J
Chapter 17
ECHINODERMATA
17.1 GENERAL CHARACTERS
Echinodermata constitutes a group of well-defined, exclusively marine invertebrates which are
characterized externally by spiny skin from which it derives its name (Gk.- echinos: hedgehog;
derma : skin). Internally, they possess a 'water vascular system' which is also unique for this
phylum. Echinoderm may be considered as the closest relative of the phylum Chordata in having
an endoskeleton which however, unlike chordata, a mesodermal skeleton, covered by a thin outer
skin epidermis that encloses the inner soft part (instead of supporting them like chordates). As
the outer skin is lost during fossilization, the skeleton appears apparently as external. Echinoderms
also have a distinctive body plan based generally upon a five-rayed or pentameral pattern.
Depending upon the angle between these five rays, the skeleton may exhibit a radial or bilateral
symmetry. The skeleton, often called test is made up of numerous calcareous plates. They are
animals of triploblastic-coelomate grade with oligomerous body plan.
Echinoderms include such living animals like sea urchins (echinoids), starfishes (asteroids),
brittle stars (ophiuroids), sea cucumbers (holothuroids), sea lilies (crinoids) and the extinct
blastoids and cystoids (Fig. 17-1 ).
Internally, the animal possesses a true coelom (body cavity) where lies a well-defined
digestive system starting from mouth and ending at anal aperture (Fig. 17-2). Depending on
life habit and symmetry, the position of mouth and anus on the skeleton may change. Food
grooves along the mid-line of ambulacral areas are provided with cilia whose rhythmic
movements produce water current towards mouth where the food-particles are picked up. Inside
the skeleton also occur the water vascular system (performing various functions including
locomotion and respiration), reproductive system, and an ill-defined nervous system.
Most of the living echinoderms like sea urchins and star fishes are vagile benthos living at
shallow depth of sea. Most of the crinoids are sessile, attached to sea floor by skeleton-built
stems. However, many extinct fossil groups like cystoids, blastoids were also sessile.
Echinoderms have a long geological history from Cambrian to Recent.
In search of ancestry of this group one should go back into Precambrian as their earliest
members appeared in Lower Cambrian and most of the existing and advanced groups appeared
by Orodovician. The study of embryology of the modern echinoderms suggests that some
Precambrian annelid worms possibly chaetognaths might be the ultimate ancestors of this
phylum but between them there should be an intermediate simple bilaterally symmetrical form
as represented by the early larval stage of echinoderms, often termed as dipleurula (meaning
little two-sided). This also indicates the existence of a primary bilateral symmetry of the phylum.
In some respect echinoderms are related to hemichordates and chordates in the manner of
forming their body cavities and mesoderm. According to some authors, echinoderms belong to
a special branch of the animal kingdom in the chordate-line which arose in the early history of
life (Lower Cambrian) and developed independently.
211
PALAEONTOLOGY
2\2
• \D (Pe11racri11us)
CR\1'10
HOLOTHUROID (Holotl111ri11a)
ASTERO\D (Asterius)
ECHIN01D (£chin11s)
OPH\URO\D (Op/1/oderma)
flG. 17 - l : SOME MEMBERS OF LIVING ECHINODERMS
·-~
--
t:CHJNODERMATA
213
Md M
~..___..--~.---Re
~~~~---Sc
Ili~- Rdc
Fl
u..--CP
Ophiuroid
Holothuroid
Am
Asteroid
Md
n,...-1-1--- Ft
~'/~~~Sc
Crinoid
Echinoid
FIG. 17 - 2 : BROAD SIMILARITY IN INTERNAL MORPHOLOGY OF DIFFERENT

GROUPS OF LIVING ECHINODERMS (cross section passing through
mouth and anus)
A: Anus , Arn: Ampulla I Cp: Calcareous Plate I Ft: Food Tube, M: Mouth, Md: Madrcporite ~ Re: Ring Cann!•
Rdc: Radial Canal, Sc: Stone Canal (Md Sc Re Rdc Am Together Form Water Vascular System).
214 Amb-Phllc
Madrcporite
Porifcrous Zone
Ambulacrai A
Oculo-Genitul rca
Ring
Apical System
lntcr-Ambulacra,
Arca
lnterambulacra
- - - - Ambitus
Ventral (Oral) View
Dorsal (Aboral) View
(a) BASIC MORPHOLOGY OF A REGULAR ECHJNOID
Anterior Lah:ral +-----ti-.'i...-o:,:..M~

Ambs (2)
Pos!crior-------~~'rjr;
Later~I Ambs (2) .,..""""'-.,,,.
Oculo-Genita14"--------~MD
Ring
Granule View
(b) BASIC MORPHOLOGY OF AN IRRHilJI .AR ECHINOID
Anterior
A Anlcrior-poslcrinr Linc llf Hilarcral Sym111c1ry
A
Length _____F_ _ _____ l B
Ventral
Side View
(c) ORIENTATION
FIG. J7 - 3
~g~rN~~~O~~~~L FEATURES OF REGULAR AND IRREGULAR
ECHINODERMATA
215
Circular Ambilal Outline Dorsal
4'
Circular
5'
Compressed Circular
4 '',,f4
Dorsal View ',~1o~' ("'>..
( 1-5 : Five Planes of Radial Symmetry) ·--~~(__ _
--~ Semicircular
Side View
(a) SHAPE OF REGULAR ECHINOIDS
u
.!:
:i
0
'3
i<
Pentagonal Hean Shaped
Circular
~ .~
L-.Y.J ~~~!
·· ~ Planoc~nvex cii
Semicircular Concavo-Convex . Subelhpucal
(b) SHAPE OF IRREGULAR ECHINOID
A
M~uth (Central Pentagonal)
Mouth & Anus A
Both Circular
Diametrically -'----'--Mouth Crescntic
Opposite , Anteriorly- ·
Shi°fted ·Along
Ventral ~mmctty Linc
Dorsal Regularia
A1 Ventral
A
Anus Shifted
Posterriorly Along
Symmetry Line'
P • Dorsal P'----Anus
Supramarginal
Marginal
Venu1ll
(c) SHAPE & POSITION OF MOUTH ANUS lnframarginal
FIG. 17-4 : MORPHOLOGICAL FEATURES OF ECHINOID TEST
( D : DORSAL V : VENTRAL A : ANTERIOR P : POSTERIOR )
Palae(Gco)WP-28
216 PALA £0/l'f OUJO,
All echinoderm skeletons are composed of calcareous plates, spicules and ~p,n . Tii
skeleton has a characteristic microstructure in which regularly arranged minute pa.. ' ~ -
permeate the calcite, and thus in section, skeleton may be recognized by its typical honeycoo
pattern.
17.2 CLASSIFICATION
Although, classification presented in the 'Treatise on Invertebrate Palaeon10/ug Moore
et al., 1978) represents the most recent and details of subdivi sions of the phylum a r~e
simplified classification is presented here mainly following that of Moore and other t952J.
This classification is based on mode of life, presence or absence of fixed or flexib le arm, , ~
and symmetry of skeleton etc.
Phylum : Echinodermata :
Subphylum : Pelmatozoa (Gk.-pelmetos : stalk; zoan : animal). sessile echinoderrm.
Class 1 Cystoidea (Cam.-Dev.) : (Gk.-kystis : bladder; eidos : resemblance).
globular or sac-like calyx with perforated test and five arms e.g. Sinocynis.
Class 2 Blastoidea (Sil.-Perm.) : (Gk.-blastos : flower-bud). common ly caJ cd
'sea-bud', no arms e.g. Pentremites.
Class 3 Crinoidea (Cam.-Rec.) : (Gk.-krinon : lily). commonly caJled '.sea-lily'.
with five mobile arms e.g. Encrinus, P~ntacrinus.
Subphylum : Eleutherozoa (Gk.-eleutheros : free). Free-living echinoderms .
Class 4 Stelleroidea (Ord.-Rec.) : (Latin-stelle : star) skeleton star-shaped with
or without arms e.g. Devonaster, Asterias and ophiceroid like Ophioderma.
Class 5 Holothuroidea (Ord.-Rec.) : (Gk.-holothurion : water-polyp) commonly
called sea-cucumber; have a sac-like body e.g. Holothuria.
Class 6 Echinoidea (Ord.-Rec.) : animals resemble spiny hedgehog, commonly
called sea urchins, variably shaped, bilateral or pentagonal symmetry e.g.
Cidaris.
In this chapter a details of skeletal morphology of echinods together with a brief rnorphologjc
features of blastoids and crinoids have been described.
17.3 ECHINOIDS
17.3.1 Morphology of fossil Echinoids (Fig. 17-3, 17-4)
A. Symmetry (Fig. 17-4)
The calcareous skeleton of echinoid is called test which is made up of numerous calcareous
· plates serially arranged. Normally, the skeleton exhibits a five-rayed pantameral symmetry. Some
modern and fossil echinoids are found having a bilateral symmetry superimposed upon the radial
plan. Accordingly, most of the palaeontologists have subdivided echinoids into two dit.•is ions :
regularia or endocyclic echinoids (with radial symmetry) and irregularia or exocyclic echinoids
(with bilateral symmerty). All other morphological features on the skeleton in each group are
usually arranged in accordance with the symmetry of the test. These two morphogroups possibly
represent two separate modes of life. Most of the regularias are vagile benthic and irregularias
are infauna) burrowers.
ECHINODERMATA 217
B. Shape of tests (Fig. 17-4 a-b)

Tht' shape of cchinoid tests is variable and sometime reflects its symmetry. For regularia,
tests .1re limited within globular, subglobular, he mispherical or spheroidal shape and in all the
cases. they exhibit a circular outline of the test. Irregular echinoids, on the other hand, exhibit
more variable shapes, such us he mispherical, ellipsoidal, flattened pentagonal and a special type
called heart-shaped. The side of the test bearing aperture for mouth is the ventral or oral side
and the opposite is the dorsal or aboral side which may or may not bear anal aperture.
Peripheral outline of the test is called ambitus.
C. Other morphologic features (Fig. 17-3-17-7)

Peristome A circular/subcircular area enclosing mouth or oral opening surrounded by
imbricately arranged soft plates. In fossil specimens these plates are mostly
lost. In regularia, the peristome is situated at the centre of oral side; in
irregularia, it may be central or may be shifted towards anterior direction
along symmetry line. The oral aperture may show a circular, subcircular or
cresentic outline. In regularia, five pairs of gill notches are present at
junction of interambs and peristome margin (Fig. l 7-4c).
Periproct : A circular/subcircular areas enclosing anal aperture and is covered with a
leathery skin studded with small overlapping soft plates not usually
preserved in fossil. Like peristome, periproct may be circular, subcircular,
elliptical or wedge-shaped in outline. In regularia, periproct is found at the
centre of aboral side just diametrically opposite to mouth. In irregular
echinoid periproct is always eccentric and shifted posteriorly along
symmetry line and it could have three positions (Fig. l 7-4c).
(a) Supramarginal : Periproct on posterior aboral side e.g. Stygmatopygus.
(b) Marginal : Periproct on posterior margin e.g. Breynia.
(c) Inframarginal : Periproct on posterior oral side e.g. Echinolampas.
Oculo- : A cirlet of IO plates, 5 smaller triangular ocular plates alternating with 5 larger
genital ring and polygonal genital plates occurs around the periproct. Each of these plates
(Fig. l 7-3a, is perforated (ocular pores and genital pores) and function in vision and
I7-5a) reproduction respectively. In irregular echinoids, anus is situated outside the
oculogenital ring and the latter may be variously modified changing its
pattern and number of plates.
Apical system : Oculogenital ring when encircles the central anus in a regular echinoid; it
(Fig. I 7-5a) may be of two types :
(i) Insert : When ocular plates extend upto periproct margin separating
the two adjacent·genitals completely.
(ii) Exsert : When ocular plates do not extend upto periproct margin and
the two adjacent genital plates are in contact near the margin.
218
PALAEONTOLOGY
Ocular Plate
Exsert Dorsal view

Insert f Compound Plate '1
(a)
TYPES OF APICAL SYSTEM IN REGULAR ECHINOID
- - - ~ Ambulacral A
CDiademoi<l] (Echinoit!J
____.._ Areas (Amb)
Jntcrambulacral fp
I Areas (lntcramb) ·
~ S Ambs Bilaterally
5-Ambs all similar, Radially Arranged (ii) Ambulncral Plates
Aligned in a Pcntameral Symmetry lrregularin
Rcgularia (i) Ambs' Arrangement
All Ambs
Anterior One Largest Anterior One Smallest Equal
Sirnpk Petaloid Subpetaloid
Posterior .l'nir Smallest Posterior Pair-Largest
(iii) Shape Of Ambs
(iv) Relative Size Of Ambs In lrregulari11
(b) MORPHOLOGY OF AMBULA.CRAL AREAS
.Nonconjugate Pore
Pair with--1..
Circular Pores
Pore Pairs with Slit-like Pores In

Uniserial / Non Biscri_aJ Conjugate Circuh1.-
Pore Pairs In p P . slit-like and Circular Pores Peripodium
Conjugate ore nirs
Compound Plates pores
(c) PORJFEROUS ZONE IN AMBULACRAL PLATES
FIG. 17- S: MORPHOLOGICAL FEATURES OF ECHINOID TEST
ECHINODERMATA 219
Madreporite: Right anterior genital plate which becomes somewhat larger than other and
(Fig. 17-5a) contains either a large perforation or numerous sieve-like small perforations;
connect~d with the internal water vascular system; with respect to this genital
plates an echinoid test cannot be called symmetrical.
Corona The part of the test excluding periproct, peristome and oculogenital ring.
Ambitus Peripheral outline of corona, that gives the outline-shape of the test.
Ambulacral areas
(Ambs) Five narrow bands on corona _originating from each ocular plate and
(Fig. l 7-3a, continuing upto peristome in regularia; composed of at least two alternate rows
17-5b) of perforated calcareous plates called ambulacral plates.
Interambulacral areas
(lnterambs) : Five broad bands (alternating with five ambs), each extending from one
(Fig. l 7-3a, genital plate and continuing up to peristome in regularia; each composed of
17-3b) at least two rows of imperforated plates called interambulacral plates.
Coronal Number of calcareous plates constituting ambs and interambs; may be of two
plates types (Fig. 17-3a, 17-5b-c)
(i) Simple plate : Plates composed of a single piece; characteristic
of many primitive and advanced echinoids.
(ii) Compound plates : Composed of fusion of more than one piece,
generalJy one of which becomes slightly larger;
segments are called demiplates. These are of two
types;
Diademoid : lowermost segment is the smallest one,
Echinoid : lowermost piece is the largest one.
Alignment of ambulacral areas [Fig. l 7-5b(i)]

In regular echinoids, the five ambs are all of similar-sized and shape and are equidistant
from one another conforming with the radial symmetry of the test. Mostly they are flushed
with the surfa,;e.
In irregular echinoids, the five ambulacral areas are bilaterally arranged conforming the
symmetry of test; an anterior lateral pair on either side of an anterior unpaired amb and a pair
of posterior laterals; of these five, anterior unpaired one may be the largest or smallest depending
on the shift of centre of test towards anterior or posterior direction. In the former case, posterior
laterals will be the largest and in other case they will be smallest. The angles between anterior
unpaired and anterior laterals and the latter with the posterior laterals may not be the same;
ambs may be flushed with surface, may be elevated or even depressed. In _irregularia. the main
ambulaeral areas are restricted on the aboral surface only.
220
PALAEONTOLOG y
Primary [
Spine I Shaft
Stalk
Periproct
Secondary Tubercle _ ___;:~:..;

Primary Tubercle
A Magnified lnteramb Plate
(a) TUBERCLES AND SPINES OF REGULAR ECHINOm
Endopcla lou
Peripctalous
Fasciole
Fnsciole
Anal
Fasciole
Lateral--~·'-~
Fasciole
Anal Fasciol ---.,;i~.ll Arrnl Fasciole
Dorsal Vi«:w
Posti:rior View
(b) DIFFERENT TYPES OF FASCIOLES IN IRREGULARIA

Margin of
Phyllode Peristomc (Mouth)
Labrum
Bourrelct
Trident ate
Globifcrous
(d) PEDICELLARIAE
FIG. 17-6 : ORNAMENTAL FEATURES ON ECHINOID TEST
ECHINODERMATA
221
Shape of ambs : Ambulacral areas may be variable in shape as follows.
[Fig. I 7-5b(iii)]
(i) Simple or
Ray-like : Ambs of most of the regular echinoids and a few irregular echinoids
are simple ray-like, increasing in width from periproct to peristome and
it is maximum at periphery of the test.
(ii) Petaloid or
Subpetaloid : Most of the irregular echinoids exhibit ambs having the shape of the
petal of a flower or nearly so; many irregular echnoids have a
combination of both simple and petaloid ambs. ·
Poriferous zone : Series of pores are present on the outer edge of each simple ambulacral
(Fig. I 7-5c) plate or on each piece of a compound plate; each plate bears at least
one pair of pores called pore-pair; pores in a pair may be circular, slit- ·
like or a combination of the two.
Interporiferous : The mid-zone of each ambulacral area lying in between the two pore
zone bearing margins without any perforation.
Uniserial : When each ambulacral plate bears one series of pore-pairs; generally
poriferous zone characteristic of ambs with simple plates.
Biserial/friserial : When each ambulacral plate bears poriferous zone consisting of two or
poriferous zone three (even four) rows of pore pairs. This is found in case of compound
ambuJacraJ plate where each segment of a plate has a pair of pores.
Conjugate pores : Pore-pair in a plate when joined by a tranverse groove or slit.
Peripodium : Pore-pair situated within a depressed zone surrounded by an elevated
rim.
D. Ornamental features (Fig. 17-6)

Surface of echinoid test is ornamented mainly by spines which are articulated with some
elevated spherical bodies on plates called tubercles. Spines and tubercles are variable in shape
and size. The chief types are as follows : -
(i) Primary spines Large movable projections which occur singly on inter-
ambulacral plates. Each spine has a basal concave depression
(attachment zone) called aeetabulum, surrounded by a flange
(serving for attachment of muscle) called milled ring. followed
by a smooth tapering part, stalk and finally the main body of
spine called s/,a/t (may be smooth or corrugated); at junction
of shaft and stalk occurs a ring-like structure called collar or
neck.
(ii) Primary tubercles Prominent rounded elevations generally on ambulacral and
interambulacral plates of regular echinoids which bear primary
spines. Each has two parts : an elevated and rounded boss at
the · c.e nter on which occurs a small spherical mass called
222 PALAEONTOLOGY
mame/011, tip of which is perforated. Acetabulum zone of a spine 1

is articulated with convex mamelon; spines are activated by
several muscles attached with milled ring. Areole is a depression
on which boss is placed.
(iii) Scrobicular Small rows of granules surrounding the areole that bear fine
granules spines.
(iv) Secondaryffertiary/: Tubercles/granules of variable size from small to very small are
l\1iliary tubercles· found scattered over the amb and interamb plates bearing similar
sized spines.
(v) Pedicellariae Group of two, three or more minute snapping jaws mounted at
Fig. 17-6d) the tips of smaller spines scattered over the surface of the test
and used mainly for defence; may be triphyllous, ophicepha-
lous, tridentate and globiferous types. Such spines are detached
from tubercles after the animal's death and are fossilized
separately, often as microfossils.
(,;) Fasciole Apparently smooth and narrow bands on the surface of some
(Fig. I 7-6b) heart-shaped echinoids found at localized areas on the test
bearing minute milliary granules which in living condition bear
hair-like cilia. According to its position it may be endopetalous
(within the petaloid ambs), peripetalous (a ring surrounding
petaloid ambs), subanal (beneath the anus), anal (occurring
round the periproct) and lateral th~t leads from peripetalous
fasciole to the rear side along the flanks of test.
E. Other features
Floscellae (Fig. l 7-6c) A flower-like structure possessed by the.members of cassiduliocl
group, centered around the peristome; It has two parts :
depressed leaf-like areas corresponding to dorsal ambulacral
segments called phyllodes, that are separated by bulging areas,
bourrelets corresponding to interambs of dorsal side.
. . .
Labrum (Fig. I7-7b) A single elongated and elevated plate next to ventral peristome
found in some spatangoids representing the anterior most part
of posterior unpaired interamb occurring as a projected lip below
the mouth (peristome).
Actinal funrrows ~adial gro~ves, leading to peristome from periphery of oral

(Food grooves) side found m clypeasteroids, serving to transport food particles
(Fig. 17-7a) to the m?uth. In some forms they are branching and reaching
the margm of oral side.
Lunules/Marglnal Five enclosed perforations, two pairs in front of posterior paired

notches (Fig. I 7-7a) ambs and one. on posterior interamb behind the periproct (anal
lunule) found m some flattened clypeasteroids which are mostly
ECHINODERMATA
223
sand dollars. Such forms also sometimes exhibit five notches or
embayments on the margin in front of each petaloid amb.
Ambulacral lunules possibly act as short-cut ways for transferring
food along with water from aboral side to the mouth; also act as
hydrodynamic stabilizers; anal lunule acts as an outlet for excess
water drawn into the mouth by ciliary feeding currents.
Plastron (Fig. 17-7b) A flattened area behind the mouth on posterior interambulacra,
densely covered with tubercles bearing flat, paddle-shaped
spines, found in some heart-shaped echinoids.
Aristotle's lantern Masticatory or jaw-apparaturs of echinoids found inside the
(Fig. 17-7c) mouth aperture, consisting· of about 40 calcareous plates
(ossicles) operated by 60 muscles arranged in seven sets;
described first by Aristotle and bearing a fancied resemblance
to a lantern; Ossicles are arranged in five identical groups which
include five rod like teeth, five pairs or pyramids (partly fused)
five pairs of epiphyses, five rotulae and five pairs of compasses
(each pair fused). The entire structure may be raised of lowered
by muscle-activity.
Perignathic girdle Inside the peristome occur projected plates two from each
(Fig. 17-7c) ambulacral area (auricles) and two from ·each interambulacral
area (apophyses); together they form perignathic gird/~ which
supports the lantern.
F. Orientation and Dimensions (Fig. 17-3c)

Anterior Recognition of anterior direction for a regular echinoid can be
done only by the location of madreporite which has an anterior-
right position. The direction of ambulacral area immediately left
to modreporite is the direction of anterior of the test. Finding
out the anterior direction in an irregular echinoid is rather easy
where direction of unpaired amb indicates the anterior direction
of the test.
Posterior Opposite direction of anterior; in irregular echinoids, it is the
direction of shifting of periproct.
Ventral (Oral) Mouth-bearing side of the test.
Dorsal (Aboral) Opposite to ventral side; in irregularia, it · bears the ambs and
interambs and in regularia it bears central periproct.
Length Line joining anterior-posterior direction; parallel to bilateral-
symmetry line of an irregular echinoid.
Width Maximum lateral distance on ventral or dorsal side at right
angles to length.
Thickness Maximum distance between ventral and dorsal surface.
Palae(Geo)WP-29
224 Pr\LAEONT LOCY
,.,..____:::i...----- food Gro<,~ C Or

"Actinnl Furrow
Margi11nl NNdl
Aboral View Oral View

(a) SOME UNIQUE FEATURES OF CLYPEASTEROIDS
Labrum
• ,\nu,
Ornl View
(b) SOME FEATURES OF SPATANGOIDS
Epiphysis
Outwardly -~~~11111~
Projected Auricle
Form Each Amb
:'/'i~ml'l;,l=~.........11,.,£..----- lnwardl,Y Projected
-~~:.U Apophyscs From
Euch lntcramh
(c) JAW APPARATUS OF A TYPICAL REGULAR ECHINOID (Aristotle's lantern nnd Pcrignuthic 1irdle)
FIG . 17-7 : SOME SPECIAL MORPHOLOGIC FEATURES OF SOME ECHINOID TESTS
ECHINODERMATA 225
TABLE-13
Distinction between Regular and Irregular Ecbinoids
Regularia (Endocyclic) lrregularia (Exocyclic)

I. Symmetry Radial, pentamerous Bilateral
2. Shape Ambitus . circular in outline; Ambital outline variable; circular,
shape may be spheroidal, elliptical, pentagonal; shape may be
hemispherical. hemispherical, oval, discoidal,
heart-shaped.
3. Peristome Central on oral side, circular May be central, or may be shifted
in outline. towards anterior on oral side along
symmetry line; circular, subcircular,
cresentic, pentagonal in outline.
4. Periproct Central on aboral side dia- Always posteriorly shifted along
metrically opposite to mouth, symmetry line; may be supramar-
circular in outline. ginal (posterior-aboral), marginal
(posterior margin) or inframarginal
(posterior-oral); may be circular,
sub-circular, elliptical, wedge-
shaped in outline.
5. Oculogenital ring Central, surrounding peri- Always outside the periproct; may
proct, hence forming an be modified variously, no apical
apical system. system.
6. Ambulacral areas
(a) Arrangement : Ambs equidistant from each Ambs bilaterally arranged; one
other, radially arranged. anterior unpaired, two anterior
laterals (pair) and two posterior
laterals (pair): angular distance
between two adjacent ambs at one
s.ide may be different.
(b) Size : All ambs equal/similar. All may be equal in size; but
generally" anterior one become
largest and posterior· pair smallest
or it may be reverse.
(c) Shape : Simple ambs. Ambs may be simple, petaloid or

subpetaloid; a combination of these
three may occur, such as anterior
one ray-like others are petaloid.
226 PALAEONTOLOGY
Regularia (Endocyclic) Irregularia (Exocyclic)

(d) Plates/poriferous Plates may be simple or com- Plates usually simple, poriferous
zone: pound, poriferous zone may zone usually uniserial, pores may be
be uniserial/biserial/triseral; circular/slit-like or a combination of
pores usually circular. these two.
(e) Relative position: Amb and interambs are m Ambs and interambs may be flushed
same level, flushed. in same level or ambs may be
depressed or elevated.
7. Ornamentation (a) Surface ornamented with (a) Surface usually ornamented with
variable sized-tubercles/ small spines and fine granules.
spines, primary, secondary or
miliary.
(b) Such structures like flos- (b) These structures are found in
cell a, fasciole, labrum, some groups of irregular echinoids.
ptastron absent.
17.3.2 Functional morphology .

A. Water vascular system (Fig. 17-2, 17-8)
Water vascular system of echinoiderm is a unique internal organ consisting of some canals.
and tubular elements.
In a regular echinoid; the system starts from madreporite of the dorsal side. This sieved-
plate leads internally to a calcified tube (stone canal) which comes vertically downward from
anus to mouth (dorsal to ventral). Below the mouth aperture it forms a ring surrounding the
oesophagus (circum oesopliagal ring canal). From this, five canals radiate out and each passes
below the mid-way of each ambularal area (radial canal). The end part of each radial canal
finally comes out through each ocular-pore of apical system. From radial canals arise at intervals,
paired lateral tubes on either side each leading to a tube foot with inflated sac-like base called
ampula. Each tube foot is finally divided and passes through each pore-pair of ambulacral plates.
Tube feet can be extended out of the test when they expand with water and withdrawn inside
when retract. The entire system is filled up with water when it operates. The system performs
mechanical as well as some physiological functions like locomotion, respiration, excretion etc.
Most of the regular echinoids have tube feet with sucker at their tips used for grasping the
hard substrate, locomotion and respiration. These become internally divided by some longitudinal
membranes dividing oxygenated and deoxygenated fluid.
Irregul~r echinoids on the other hand, have different types of tube feet. Such echinoids like
clypeastends and spatangids which are mainly sand burrowers have developed broader and
flattene_d tube feet, mainly used for respiration. Although, tube feet are not preserved in fossils
but theu number and nature of function can be determined by the nature of the pores within
I 'lll ,\ '1)11/ 'Ni\t\l .\
~loolh
R l:HNbltJ AL NO OORS VENTRAL PL".NE
I 11t>1.' I \·,·t - ---l
a.---Musl.'lc
l\111lt-l'h1tl)
- ~ - - l\!1 • t'nir
/\mpullil
(b) A C..:IR 'llLAR TUBli t,'()()1' WITU SllCKFR AT TOP AND CIRCULAR PORES AT BASL
11. S. l'IIIH llKill A TlJt\E t-=l:t-=Tl
- Alli/l. f>lu1ti •·-.
11. S Tlll<OUOII A TUBE F~hTI

tu) A f'LATl'FNED Rl~SPIRATORYTIIBE FOOT ANDSLIT-U~ PORE
I
.l
.\ FIG. 17 • 8: WATER VASCULAR SYSTEM AND ASSOCIATE FEATURES
'
:fi
228 PALAEONTOLOGY
poriforous wn, of the :unbulacral urea. For example, a respiratory tube foot leaves on the test
a pair of por ·s oftt!ll joined by u furrow (conjugate pores). Occasionally, these pores become
elongated (slit-like) and reasonably. they ure found on the test of burrowing echinoids. Tube
feet with su k ~rs an.! characteristic of vagilc bcnthic forms used for locomotion and anchoring
and such tub, foct leave circular pore-pairs for their passage through the amb-plates.
B. Fnsciole
Fascioks 111·" found only in some burrowing echinoids such as spatangoids. Fascioles are
smooth :rnd narrow hands on the surface of the test of these echinoids apparently appearing
free from any tubercle. However, on close observation very small microscopic granules are found
here. These microgranules in living condition bear numerous hair-like cilia, the movement of
which creates water current. For an infaunal form such current is necessary as it increases the
amount of oxygenated water crossing the petaloid ambs and also aids in feeding and sweeping
away excreta. It also prevents mud/clay particles to settle on the poriferous zone so that water
vascular system can function properly.
17 .3.3 Classification
Subdivisions of the class Ecl1i11oidea into Regularia and Irregularia is simply a convenient
and provisional method rather than u phyletic grouping. The classification erected at present
by Moore er al. ( 1966, 67) in Treatise on Invertebrate Palaeontology was based upon a wide
variety of characters and skeletal features, such as rigidity of test, morphology of plates, nature
and composition of perignathic girdle and aristotle's lantern and so on. The classification upto
the level of order is given below :
Class : Echinoidea
Subclass 1 Perischoechinoidea (Ord.-Rec.) : regular, endocyclic, plates simple/
compound, perignathic girdle simple or absent, latern with grooved
teeth. e.g. Echinocystis.
Subclass 2 Cidaroidea (Dev.-Rec.) : regular echinoids, with two columns of
plates in ambs and interambs; perignathic girdle with only apophyses
e.g. Cidaris.
Subclass 3 Euechinoidea (Trias.-Rec.) : a large group with bicolumnar ambs and
interambs, perignathic girdle with both apophyses and auricles;
lantern may be absent.
Infraclass 1 Echinoithuroidea : echinoids with flexible test and pseudocompound
amb-plates.
lnfraclass 2 Acroechinoidea : upright lantern, compound amb-plates.
Cohort 1 Diadematacea : teeth grooved, lantern cidaroid type, e.g. Diadema.
Cohort 2 Echinacea : keeled teeth, solid spines, compound plates and complex
perignathic girdle.
Order 1 Stirodonta : auricles and apophyses in perignathic girdle, epiphyses
short. e.g. Plesiocidaris, Hemicidaris.
I-' '1//Vl I fN.\1.\1:.\ 229
Or,kr ~ Aulod01ll:1 : rigi i t --st. pt·rignathi girdle , ith auricles and apophyses,
l~l'th without er'" · \t' N keel. e.u . Di{l{/ mopsis.
Or<kr 3 CamaroJt nta : Pt'rignathic gi k wi th onl y auricles. epiphyses larger
e.g. cd1i11us.
Ct)hort J ltw~ulnria.
Supt•rorder t E0gnathostomata : di:tinctive t pe of lantern structure ; nearly
hemispherical tct.
Order t Hok ·typoida : the test retains r.idial symmetry except the shifting
of periproct t posterior. e.g. Holectypus .
Order 2 Pygusteroida : periproct key-whole shaped. extending beyond the
apil·al disc. e.g. Pygaster.
Superorder 2 Neognuthostomata : pentagonal or elliptical in outline, discoidal or
hemispherical. latern present at least in juvenile stage.
Order I Cassiduloida : subglobular; floscella present, lantern in adult. e.g.
Ec/1i110/ampas, Stygmatopygus.
Order 2 Clypeasterioida : discoidal, food grooves on oral side, e.g. Clypeaster.
Superorder 3 Atelostomata : hean shaped, no lantern.
Order 1 Disasteroida : apical system split. e.g. Disaster.
Order 2 Holasteroida : elongated apical disc. e.g. Ho/aster.
Order 3 Spatangoida : compact apical system without posterior genital,
marginal periproct e.g. Miscraster, Hemiaster, Breynia.
17.3.4 Ecology and mode of life (Fig. 17-9)

Echinoids are t!x.clusively marine benthic animals mainly ,·agile or infaunal burrowers or
cavt! dwellers. In seu, they are r,mging from intertidal down to abyssal depth.
Males and females ure separated; fertilization is external; eggs develop into free floating
larvae which settle down to become adult echinoids. In some forms, larvae are not set free;
instead they are protected by the mother. The young echinoids grow within the spines of the
mother until they become adult.
Regular echinoids mostl live on the sea tloor, moving on their stout tubular tube feet
provided with terminal sucker· They are generall y restricted to hard rocky bottom (reef zone)
and they sho;--various modifications which pem1it them to withstand the surge of waves and
I currem ·. Large spines and sucker-bearing tube feet give them secured anchorage on the bottom.
Some cave dwdling forms may occupy natural holes and while growing they enlarge the holes
by their spines and strong teeth making them as thdr permanent houses (Fig. 17-9d)]
I
lrregularia represents a divergent lines of adupt,ltion to lifo on soft bottom ,md mostly they
are burrowers living in colonies within sand ,md mud. They have succe.-.sfully U\c~loo all ~he
problem · of burrowing organisms such as how to breath, cat and excrete. All such features hke
l
,}
,.
PAI.AEONTOLOGY
230
Mouth
Wnrer
Currenr -----.•..l
Fnccol Product
Food uorhcrins
Tube Feel Prom
U11p11irc,d Amh
Mo111h
Plustron Spines Preventing Wutcr Flow

I.11 brum
(u) HlJRROWER SPATANOOID AND ITS .MECHANISM OF POOD COLLECTION
(c) A SHOL.LOW SAND DOLLAR CLYPEASTEROID

(b) SPl/tfD ECIIINOIO
(J) AHP.GULAR ECHINOID WITHIN ROCK-CAVE F11Tl20 BY ITS LARGE SPINES
FIG. 17 ·.9: DIFFERENT MODES OF LIFE OF ECHINOIDS
ECHINODERMATA -- I
change of the shape (highest posterior and sloping a;1teriur) and symmetry (bilateral) of the
test shift of the mouth (near anterior) and anus (marginal) in opposite dir 'l:lion. el vat ·d labrum
bcl~w the mouth, depressed ambulacral areas arc some features r·lutcd to their successful
adaptation to burrowing life. Several fascioles on the aboral surfaCl' are clearly helping them
to create several water currents directed al different direction for apturing food part id "'S.
performing respiration and removing the waste away from the body (Fig. I 7-9a).
Infaunal and burrowing habit was found to establish in echinoids since Crctat·eous. The
spatangoids or heart-urchins like Edti11ocardiu111 me rather deep burrowers living at 10 to 20
cm deep burrows (Fig. l 7-9a). They keep connection with wuicr surface hy digging a hole with
the help of a long slender tube foot with bristly termination like a chimnt:y sweep's brush.
This functions as incurrent funnel drawing water inside the hole. A similar but blunt tube foot
extends from anal area parallel to the surface which serves as a sanitary 'soukuway (Nicol,
1959). The inhalant tube foot ends above the apical part of anterior amb which is usually
surrounded by a fasciole (peripetalous) with numerous cilia that create a downward current
forcing water to pass along the sloping and depressed anterior unpaired amb where sticky food
gathering tube feet capture food particles and pass them to the mouth lying more anteriorly
on the ventral side. Another current sweeps over the paired petaloid ambs where flattened tube
feet perform respiratory function . A third current is created by cilia of sub-anal/anal fasciole
to carry the waste product away from mouth down the drain. When faecal materials fill up the
sanitary drain the animal moves anteriorly retracting the tube feet by rowing action with the
help of flattened spines on plastron resulting co1\apse of the burrow-hole·. In the new position
it creates another hole by the tube feet. Besides this normal burrowing habit, as shown by most
spatangids, some unusual forms appeared in Cretaceous which developed a snorkel-like proboscis
called rostrum projecting to the surface of water of burrow-hole, thus allowing the animal to
lie at a much deeper level minimizing the chance of predation. These are often called spired
echinoids e.g. Hegenowia. They also reduced their size to improve gaseous exchange within
the burrows (Fig. l 7-9b).
Sand-dollars like clypeasteroids possess several structural adaptations for shallow burrowing
life habit ·often in the rough tidal or intertidal zone (Fig. 17-9c). Most of them lie flat on the
bottom with a thin film of sand covering the aboral side. Their flattened and discoidal test and
short fur-like spines help in their slow movement through the sand. They increase the area of
petaloid arnbs to enhance respiration by flattened tube feet. Ventral food-grooves and marginal
notches of the test allow a smooth passage of water currents with food grain towards mouth.
Additional water currents with faecal materials goes away from the body through lunules. As
the animals are inhabiting in somewhat rough zone of sea, they have to strengthen their test
and stabilize their position by taking suitable adaptation. They develop unique internal
partitions/pillars in the form of radiating ridges within inner peripheral zone of test. This divides
the interior into narrow tortuous passages giving the tests sufficient strength.
For improving feeding capacity sand-dollars diversified their ambulacral canals with
development of extra-accessary tube feet served by microcanals located on internal walls. Some
of these canals are used by juveniles to store grains of heavy minerals selecting them from
sand that increases the stability of their tests and consequently they arc mostly confined to the
beach of the sea where such mineral grains are available. These modifications have made the
P..ilae(Gco)WP-30
,. •·
--· Scanned by CamScanner
Rcgul:tr echinoids lrrcguh1r cchinoids N
'J)
Clypcastcroids Spatangoids t.J
u
ii[
5 t:
~
- la
c
g"'
u
L_
u
~r
i JI "'
"O
~~
--- -
,. ,
"'
-- _..... ---
>.
u
"'
"O
·o
0
=
:c _____ .,..- _,
11 .S
.c
u
u
ca
u
UJ I
-~ I =
=-
e
:,
c..
·au
•
1-~
-::::..M =
0
>
8
i·;:
-=c;;
-i
·o
~
~
s:
! ('?-
0
<:
~
FIG. 17 - 10 C)
STRATIGRAPHIC RANGE AND RELATIVE ABUNDANCE OF THE COMMON GROUPS OF ECHINOIDS t--
c
C)
~
233
E'CH/NODERMATA
tests of most of sand dollars sufficiently strong increasing their fossilization potentiality and
for obvious reason burrowers' fossils are more common than their epifaunal counterparts.
As poriferous zone, through which tube feet are ejected, is the most vital area of the test,
all echinoids, regularia or irregularia, always prefer a zones of clear and agitated water. Muddy
und silty water may be fatal for their life, as fine silt or clay particles will soon clot the pores
causing their death. After the eruption from the Visuvius in 1906 a huge quantity of ash was
accumulated in the water of the Bay of Nappies and echinoids were virtually wiped out from
that locality within a few years.
17.3.S Origin, evolution and geological history (Fig. 17-10)

Echinoid fossils are first observed within the Ordovician rocks which resemble the Lower
Ordovician star fishes of the order somasteroida having similar looking tube feets, not completely
enclosed within ambulaeral plates. This suggests their derivation form an ancestor in which
like the star fish, the tube feet occurred in between ambulacrals. Moreover, these Ordovician
forms lacked the lantern and possessed jaw-like structures like star-fishes. Again, radial structure
and fixed initial larval stage of many star-fishes indicate their derivation from some sessile
pelmatozoan ancestry.
From Ordovician, echinoids made a steady progress with the appearance of other groups
but their number was rather limited upto Carboniferous after which regular echinoids became
diverse but never abundant. With the end of Permian, all but only one genus Miocidaris became
extinct and latter became the ancestor of all post-Permian echinoids. From this, first appeared
Triassic cidaroids. After this, they showed a great diversity with the appearance of many taxa
of regular and irregular echinoids, adapted for different modes of life. True irregular echinoids
appeared in Jurassic which might have a polyphyletic origin from different groups of regular
echinoids. Holectypoidi. show almost a radial symmetry except a slight deviation of anal aperture
from centre. Cassidulo1ds, spatangoids and clypeasteriods all appeared by Jurassic and Lower
Cretaceous. Echinoids became the most diverse during Early Teritiary times. But after Pliocene,
cassiduloids declined greatly as did the clypeasteroids. The older group holectypoids are at
present, at the verge of extinction.
One interesting feature of phylogeny of echinoids is that the precise relationship between
structural and functional differentiation is perhaps more or less clear. This is because the
evolutionary changes in the Mesozoic and later echinoids was rather slow, so that the derivation
of one group from other can be chalked out. This was not seen in many other invertebrate groups
where vital intermediate stages among the contrasted groups became very much short-ranging
and are often lost in the record, causing difficulty in linking one taxonomic group with the
other. For example, the remarkable functional difference between Early Mesozoic cidaroids and
Late Mesozoic spatangoids, or clypeasterids has been achieved by slow and gradual modificat·
f h . . f . ion
o t .e ex1.stmg organs o t.he1r ancestors. For this reason, functional significance and structural
mod1ficat1on
. . of many extmct groups
. may be interpreted from the presence o f s1m1· ·1ar organs
with modified forms and functions found within their modern counterparts.
PALAEONTOLOGY
2. 4
--,,
~s:ca,~~~~,---.. .
I
I
Anal Pyramid
calyx
I
I
fCrown
I
I
Rad ials--==::iU..JI I
B;i.sals -=~aV ______ l
t:J----,Columner Plate
Stalk _ __.. _ __., Hold Fast
(a) THE CALYX
Anal Pyramid Anal Pyramid
Aricu.farion To J (b) ANIMAL ATIACHED J . .

~ Anculat,o~ To
'===:;;;s
V L160r~)
Brachia BY STEM Brachia Supra
Jr, MM M ~ Radials(l)--£2 Uniserial
l)AV, .V V. U,. Basals(l) 00 0 0 CfJ

LJ [j Cl L) ~asals(l) .=(} 0 0 CJ lJ Biserial
/cl MONOCYCUC CUP (side layCllfl) (c.l) DECYCLIC Cl rp (Side Layout)
(e) BRACHIAL
FIG. 17-11 : STRUCTURES OF CRINOIDS
- - - Ambulacr-------.
--Ddtoid---~ --Spiracle
Radial--~
(b) Top View
(a) Side View

STRUCTURE OF A TYCICAL BLASTOID (P£NTREIT£S) TEST
(j) ~~---.Spiraclcs(S)
- , . . - - • Dcltoids(S)
----.> Basals(J)
(c) SIDE LAY OUT OF THE CUP
FIG. 17 - J2 : MORPHOLOGY OF BLASTOIDS

ECHINODERMATA 235
17.3.6 St.ratigraphic importance
Use of echinoids in stratigraphy is, however, limited. This is partly due to their limited
environmental occurrence and partly to the conditions affecting their preservation. Most regular
echinoids have delicate test hence are rarely preserved in good condition. Irregular echinoids,
because of their harder tests and burrowing habits are more readily preserved and hence are of
greater stratigraphic importance. Many irregular forms are used as zone fossils especially in
the classification of Cretaceous rocks such as the succession of species of the genus Micraster
in M. leski-M coranguinum ljneage have been used in the zonation of Upper Cretaceous rocks
of Europe.
17.4 OUTLINE OF MORPHOLOGY OF CRINOIDS (Fig. 17-11)

A. General morphology
Crinoids are marine sessile echinoiderms whose fossils are abundantly found within
Palaeozoic rocks and unlike other sessile forms of this phylum they are still existing in the
present day. They are popularly called by the name sea-lilies. Most of the Palaeozoic forms
possessed stalks for attachment but recent forms may be with or without such structure.
Skeletons are calcareous made up of numerous plates. In general, the skeleton has three parts:
calyx, arms and a stem or stalk. They range from Middle Cambrian to Recent.
The calyx (theca) is composed of two parts, a lower conical, globular or bowl-shaped-cup,
below the zone of arm-attachment (also called dorsal or aboral) and a flat to highly domed
tqmen above (bearing mouth and anus). The cup shows several circlest of small to large plates
arranged in a pentameral symmetry. Two types of cups are found monocyclic (with only one
circlet of plates) and dicyclic (with two circlets of plates). In the monocyclic type, the topmost
circlet of plates that bear the arms are called radials and the next circlet of plates below radials
and alternating with them are basals. In dicyclic type there are two rows of plates below radials
caUed infrabasals and suprabasals. Generally five plates constitute each row and that gives
the pentameral symmetry of calyx. Tegmen is made up of few to many plates above the radials.
The five plates around the mouth are calted orals or interambs located in interadial position.
Plates alternating with them are ambulacrum bearing food grooves. Anus is usually situated
on one of the interambs. A domed plate above the anus is called anal pyramid. Sometimes
anus may be present as a raised plated tube called anal tube. Mouth is located at the central
part of the tegmen. Arms are feeding appendages, made up of numerous plates called brachials.
They are cylindrical or wedgeshaped and bear ex.tension of cilated food groove on their upper
side. Brachia! plates may be uniserial or biserial. Arms may be unbranched or branched one
or few times. Again, each brachia! laterally bears small projection on either side called pinnules
also bearing food grooves. Calyx and arms are together called crown.
Crinoid stem (Fig. t 7-1 Ob) is usually composed of short disc-shaped plates called columnals
or ossicles which may be rounded, pentagonal or star-shaped in cross section. Each columnal
contains a central cavity, lumen, through which canals and nerves extend . Two types of
columnals are found : large or nodals bearing small projection called cirri (with very small
columnal plates) and smaller without cirri called internodals. A crinoid stem usually ends into
a structure like roots (made up of several cirri) that attach the stem with substrate. This is called
holdfast. There are some free living crinoids without stem bearing a series of cirri developed
from their basal cup that hold them with sea-floor.
PALAEONTOLOGY
236
8. Geological history . f Middle Cambrian and .they became fairly

Crinoids are re~~rded for ~he first ti:~ti:~: to diversify and reached their peak in early
common by Ordov1cian-Devoman. They c . t the Permo-Triassic boundary except a few
·c c ·
Carbom,erous. rmo1 s
"d almost became extinct a
.
. . .
. 1 t group that apeared first m Triassic and
. 1· h"
d1cyc 1c types w 1c h possibly gave nse to art1cu a e , .
. . . .d re more common in deep water environment
became gradually abundant. The hvmg crmo1 s a .
where stem bearing forms are less diverse than free-lymg .forms.
Crinoids were important sediment builders in the sh:lf ~ea .of Palaeozoic times formin~ great
crinoidal limestone of huge thickness. Two important crmo1d-n~h roc~s are Upper C~rbom~er~us
Mississippi Group, Indiana, USA and Permian System of Tm~or island, Indonesia. Cnno1ds
are mostly found in some restricted zones as disarticulated fossils , calyx often detatched from
stem.
17.5 OUTLINE OF MORPHOLOGY OF BLASTOIDS (Fig. 17-12)

Blastoids are the largest and one of the most advanced group of sessile echinoderms,
exclusively of Palaeozoic age. The skeleton is calcareous and composed mainly of two parts :
a conical to globular calyx or theca which is often looked like a flower-bud (hence the name)
and very often a stem also composed of numerous plates, fixing the animal with sea floor. A
few forms lacked stem. Lower or dorsal part of calyx is composed of three circlets of plates :
three basals (often unequal), five large radials and five small deltoids. The upper surface of
calyx is marked by five petaloid ambulacral areas, each depressed at midway where occurs a
ciliated food groove. In fact, each radial plate is intercepted at its middle by each ambulacrum
on the oral side and each deltoids is located in between two ambs at its peristomal side. Mouth
is situated at the centre of calyx encircled by 5 holes called spiracles, one of which acting as
anus. Each ambulacrum is composed of two rows of fine plates (lancet-plates) centrally located
and two rows of lateral plates along its two margins. They usually bear fine brachials. One
advanced morphol.ogic character of blastoid is its greater tendency to develop a pentameral
s~m~etry and to ~ncrease the num?er of ~late~ with_in calyx. Blastoids are ranging between
Silurian and Permian, a~d they attamed their chmax m Carboniferous. Their fossils are found
from shallow sea deposits, sometimes along with Palaeozoic reef Ii t A t'. ·1·
c · of Carboniferous age.
· p entrem,tes
,oss1-1 1s mes ones. very ,am11ar
Chapter 18
GRAPTOLITES
18.~ GENE~'~ MORPHOLOGIC FEATURES (Fig. 18-1)
Graptol,.~es is ~ v~rnacular term used for some fossils found in marine Lower Palaeozoic
rocks, espec.1~lly w1.thm black shales, that resemble some ancient writings on rocks. (Gk.-
gra~tos: wntmgs; htes : stone). Faunal affinity of these extinct group of animals remains a
subJect of controversy for many years.
The skeleton of the colony is chitinous consisting of a series of inter-linked hollow tubes.
The first ~om:ied part of the colony is a conical body called sicula, having an aperture downward
and termmatmg at the apex and extending to a long thread like structure called nema. The
sicula is divided into two parts showing different ornamentation. An upper prosicula with
longitudinal and spiral striae and a lower metasicula with concentric ring-makings indicating
~riodic growth. These ri~gs are called fusella . In fact, these are half rings joined along a median
zig-zag line at one side and on other side, they are joined to a stout, longitudinally directed
rod-like structure called virgella which projects beyond the aperture. In front, the aperture is
marked by two projection called apertural spines.
The first tubular theca develops from a notch at the edge of metasicula on its virgellar side.
From this develop successive thecae away from sicula. A thecal aperture points upwards and is
provided with a pair of apertural spines.
Cavity of first formed theca and that of sicula is connected through a foramen and all other
successive· thecae are likewise linked up internally with one another through a common callal.
Each theca has two parts : Protheca, proximal to nema through which common canal runs and
metatheca, which is the outer part of theca divided up by .a medium partition-wall (median
septum) producing two common canals at distal end of the theca. Within each theca possibly
inhabited small individuals of the colony called zooids, Chitinous skeleton of the colony is called
rhabdosome, which may be branched or unbranched. Each branch is called stipe.
Some graptolites possess a stipe with two series of thecae united back to back wi,h the nema
in between when it is called biserial. Some are imiserial with one series of thecae.
The stipe-alignment in a rhabdosome shows all variation from stipes hanging downward
(pendent), horizontal, to vertically upwards (scandent). The intermediate forms are declilled
(between horizontal and pendent) and reclined (between horizontal and scadent). Biserial thecae,
according to their growth pattern, are conventionally expressed 11 - t 2, 2 1 - 22 and so on. Th 11
develops first; from which grows Th. 12 ; Th. 2 1 arises from the t 2 in the opposite side. Thus
thecae on either side having same number pass in front of the sicula, each time forming a crossi11g
canal. There may be development of a double crossing canal system when Th. 22 arises from
Th. l I instead of Th. l 2.
Thecae may be simple tubular in shape. But in some cases there are lot of variations and
complications of thecal pattern like hooked, ':::te, triangular, geniculate~ = ~ a t e : ! ~
PALAEON1'0LOGY
2)8
.-----Ncm•
Ncma .------1
H/\LF•l<INO (._ll<UWTII
IN '!<EMF.NT OJI
Apcrture-- I USP.LLAH 011
Ml.fASI 'ULA
Virgella ~ - - -
STRUCTURE
OFSICULA
Two
Common
canal nn, "Al: AND SI 'ULA
(Apcrtural view)
Tubulat
UIStRIAI.
RI IAU[)(JSOM
SECflONOF
RH~DOSOME
Hooltc:d
A Single --t-'ffl
Common
canal
(In Section)
---Pendent
J
VARIOUS POSITION s ·npE
TYPES OF THECAE
I; p
Th
l 2'
Bi-St.ipcd
Th
Multi5tiped
4-Stiped Monortipcd Double c..c. Single c.c.
NUMBER OP STIPES (Branch) IN GRAPTOLITF.S
NUMBER Of CROSSING CANALS cC C.)
FIG. : 18 - I MORPHOLOGICAL FEATURES OF GRAPTOLJTES
GRAPTOLITES
239
18.2 ECOLOGY
Grnptolites were exclusively mari . d · ·
epiplanktons. Thi s is evident from ne_ an maJonty. of them were f~ee floating planktons or
. d h f '" . their occurrence m all types manne rocks irrespective of
their ept o ,ormat1on and also fro th · . . · · ·
. . . H . . · m e1r association with other marine fossils of Ordo-
Si 1un an age. owever, there is more abundance of graptolites within Ordo-Silurian black shales
(rocks supposed to have been forn d d d · ·
. . 1e un er re ucmg environment). Graptolites were definitely
not mhabttants
. unde r such environm en t . B ut t he dead skeletons of graptolttes,
· accumulated in
such
. basms, probably
. escaped elim1·nat·1on ca · d b h 1· · · ·
use y ot er 1vmg scavangmg animals (as ltvmg · ·
amm~ls cannot thnve under reducing environment) which possibly took these skeletons as their
food m normal condition .
More recently, some supplimentary floatation mechanisms of graptolites have been described.
~ome of them had flat expanding webs surrounding the proximal of colony which could have
mcreased the surface area. Many monograptids exhibit vertically radiating vanes, arranged around
the nema which helped in stability during floating. Recently, a hypothesis of automobility of
graptolites has been advanced (Kirk, 1969, 72). It has been suggested that some graptolites were
not merely passive floaters, but can swim actively. Although, the mechanism of swimming
remains unclear, it is inferred that ciliary ornamentation of zooid might have helped in this
function. Decrease of number of stipes, their alignment and complex thecal forms, are all seen
as parts of the evolutionary response to their mode of life.
18.3 BIOLOGICAL AFFINITY

For a long time graptolites were considered as forms having a close affinity to cnidarians
and most probably to hydrozoans. Most of the authors of later period rejected this view and
pointed out a close resemblance of graptolites with small modem deep sea rhabdopleura, which
are colonial organisms belonging to subhylum Hemichordata of the phylum Chordata.
Rhabdopleuras have their fossil relatives as far back to Ordovician. They exhibit tubular
exoskeleton like graptolites, each bearing zooids. Most of the recent authors have accepted
graptolites as a separate subphylum within chordates.
18.4 GEOLOGICAL HISTORY AND EVOLUTION

Graptolites has a very short geological history. They appeared for the first time in Ordovician
from dendroid group and continued upto Silurian. Owing to their planktic habit they attained
wide dispersal within their short life-time. This makes most the species of graptolites ideal index
fossil of Lower Palaeozoic times. Within Ordo-Silurian succession of Europe three distinct
graptolite faunas have been recognised which are divisible into 43 faunal zones. (11 in
Ordovician and 32 in Silurian). Some of the important fossils of these faunas are :
Monograptid fauna (Middle to Upper Silurian)

Monograptus, Cyrtograptus, Dimorphograptus.
Diplograptid fauna (Middle Ordovician to Lower Silurian)
Dip/ograptus, G/yptograptus.
Dichograptid fauna (Lower to Middle Ordovician)
Tetragraptus, Didymograptus, Dichograptus.
240 PALAEONtOU>Gy
Strikingly, graptolites exhibited some morphologic changes ,with time. For example, primitive
Ordovician graptolites usually showed uniserial thecae, but multiple stipes. Staurograptu.f had
numerous stipes, Dichograptus possessed eight stipes, Tetragraptu.f four stipes and Didym0 .
graptus had only two stipes. But most of the Silurian forms are unistiped (Monograptu,,),
Similarly, earlier Ordovician graptolites mostly exhibited all types of stipe-alignment starting
from pendent to reclined. But Silurian forms have mostly scandent types stipes.
Chapter 19
RECORD OF INVERTEBRATES FOSSILS

FROM PHANEROZOIC ROCKS OF INDIA
19.1 OCCURRENCE OF INVERTEBRATE FOSSILS IN INDIA
In India, invertebrate fossils are known to occur in marine and fresh water Phanerozoic rocks
deposited at different parts of Peninsula and Extrapeninsula. Since Cambrian period, marine
Palaeozoic rocks in India, however, are mainly localized in various extrapeninsular basins of
Spiti, Kashmir, Simla, and Garwal. Naturally most of the fossils of marine invertebrates are
found within these basins. In Mesozoic periods, along with the marine deposits in the Himalayan
basins, several marine transgressions took place along the different parts of western, eastern
and southern peninsular coast. In Kutch, the transgression came in Middle Jurassic and the
marine condition continued upto Mio-Pliocence with a few breaks in the sequence. So rocks
of Kutch basin yield a rich marine invertebrate fauna ranging in age from Jurassic onwards.
The south-east coromandel coast (Kaveri basin) and the east coast (Assam and Bengal basin)
of India witnessed the marine transgression at Upper Cretaceous (Cenomanian) times and the
marine condition continued there upto Eocene-Oligocene and the corresponding rocks yield a
rich marine fauna. Besides these major areas (Himalayan basins, Kutch basin, Kaveri basin and
Assam-Bengal basins), isolated marine beds are found at different parts of Peninsula such as
marine Umaria/Manendragarh beds (Permian) of Rewa, marine Bagh beds (Cretaceous) of
Narmada Valley and patches of Miocene beds at Baripada in Orissa and Quilon in Kerala.
Reports of fresh water invertebrates fossils, although have been made from different continental
beds but they are of less importance, except a few. These continental beds are Gondwana Group
(Lower Permian-Lower Cretaceous) and Deccan Intertrappeans beds (Palaeogene) of Peninsula
and Siwalik Group (Neogene) of Extrapeninsula. Neogene beds of Assam and Bengal basin
also yield a few invertebrate fossils of freshwater to estuarine types. Most of these continental
invertebrates fossils are geologically less important and fails to attract the palaeontologists to
study them intensively.
The name (mostly generic) of only those fossils, generally considered stratigraphically more
important are listed in the undergoing portion. Specific occurrences of some particular forms
are given within parenthesis. Some species used as index fossils are also asterisked.
19.2 PALAEOZOIC FOSSILS

Inarticulate branchipods, trilobites and corals constitute the bulk of the Palaeozoic fauna,
mostly collected from the Himalayan basins (Spiti and Kashmir). ·
Himalayan Basins
Cambrian
Brachiopods Oho/us, Obollela, Neobolus, Acrothele, Acrotreta, Lingu/e/la,
Botsfordia, Nisusia.
''
'I
241
. ""1,
242 PALAEONTOLOc;y
Trilobites Redlichia noetlingitft, 0/enus, Agnostus, Anomocare, Ptychopczria,

Microdiscus, Tonkinella*(K}, ChuanKia*(K), lrunia (K), Conncoryphe
(K), Solenpleura, Bo/aspidel/a•, listmia, DikeJocepha/ite.'i, Blountia.
Echinoderm Eocystites.
Gastropod Hyolithes wynnei*(S).
Ordovician (mostly from Spiti basin)

Brachiopods Orthis, leptaena, Sowerbyel/a, Strophome,w, Refinesquina,
Plectambonites, leptol/oides, Triplescia (K).
Trilobites Asaphas, Calymene, 1/laenus, lichas, Dalmcmites, Acidaspi.\'
Pelecypods Pterinia, Ctenodonta.
Gastropods Be//erophon ganesa*, Hyo/ithes, Cyclonema, Tentaculites.
Cephalopods Orthoceras, Cyrtoceras, Gonioceras.
Ostracods Beyrichia, Dinulites.
Brayozoans Monotrypa, Prosopora, Ptilopora, Phy/lopora.
Graptolites C/imacograptus (K}, Didymograptus (K).
Corals Favosites forbesi*, Syringopora, Streptolasma, He/iolites.
Siluria11 (mostly from Spiti basin)
Brachiopods lingula, Orthis, Da/manel/a, leptaena, Rafinesquina, Plectambonites,
Strophomena, Atrypa, Rhynchospira, Conchidium, Euspirifer, Chonetes,
Pentamerus ob/ongus*, Triplescia.
Corals Favosites, Ha/ysites wallichi*, Syringopora, Zaphrentis, Conophy//um.
Trilobites Calymene, Phacops, Acidaspis, 1/laenus.
Cephalopods Orthoceras.
Gastropods Hyolithes, Belleropl,on.
Echinoderms Caryocrinus, Cheircrinus.
Sponge Dictyospongia.
Devonian (from Kashmir only)

Conodonts : Po/ygnathinus, Ancryodella.
Carboniferous (mostly from Spiti)

Brachiopods Products, Spirifer, Neospirifer, Syringothyris cu.'ipidata*, Reticularia,
Derbyia, Dia/asma, Athyris, Chonetes, Lingu/a, Orthis, Aulosteges.
Pelecypods Modiola, Avicu/opecten, Oricrassate//a.
Gastropods Pleurotomaria, Bel/crophon.
Trilobites Pltillipsia aff. chiftordi*.
Bryozoans Feneste//a, Protoretipora.
.,_----- --- -- -
243
RECORD OF INVERTEBRATES FOSSILS FROM PHANEROZOIC ROCKS OF IND/A
Permian
(a) Himalayan basins . . L tt ia Richthofenia*,
Brachiopods Spirifer rajah*, Products, Mergmifera*, Chonetes, . ;y .o~ •
Spirigera, Athyris, Streptorhynchus Derbya, Fus,spirifer.
Corals 7Apherentis, Waagenophyllum indicus*, Clisiophyllum.
Gastropods Pleurotomaria, Bellerophon, Conularia.
Pelecypods Eurydesma*, Deltopecten, Pterina, Nucula, Schizodus, Aviculopecten,
Solemya.
. *
Cephalopods Xenaspis, Xenodiscus, Cyclolobus, Sageceras, Gastrwceras ·
Foraminifera Parafusulina *.
Bryozoans Protoretipora, Fenestella, Dystintella, Acanthocladia, Rhabdopora.
(b) Manendragarh-Uamria basin

Brachipods Products, Spirifer, Streptorhyncus, Reticularia, Athyris, Trigonotreta.
Gastropods Pleurotomaria, Conularia, Peruvispira, Keeneia.
Pelecypods Eurydesma*, Detopecten/Nuculopsis, Phestia.
19.3 MESOZOIC FOSSILS

The major constituents of Masozoic marine invertebrate fossils are cephalopods of the
ammonoid group (mostly index fossils). The other dominant groups are pelecypods and
gastropods.
Triassic
Himalayan basins (Spiti and Kashmir)
Cephalopods Otoceras, Ophiceras, Meckoceras, Aspidites, Xenodiscus,
Hedenstroemia, Ptychites, Ceratites, Hollandites, Sturia, Gymnites,
Joannites, Beyrichites, Keyserlingites, Monophyllites Sibi-rites,
Hungarites, Flemingites, Pinacoceras, Juvavites, 1i.betites, Tropites
subbulatus*, Atractites . .
Pelecypods Pseudomonotis, Daonella, Lima, Monotis, Pecten, Halobia, Megalodon,
Lilangia, Fontolium.
Brachiopods Rhynchonella, Spiriferina, Dialasma, Spirigera.
Corals Thecosmillia, Montlivaltia, Thamnasteria.
Jurassic
(a) Himalayan basins
Cephalopods Phylloceras, Lytoceras, Analytoceras, Stephanoceras, Macrocephalites
traingularis, Virgatosphinctes, Pertisphinetes, Oppelia (Str~bli.tes),
Aspidoceras, Spiticeras, Acanthodiscus, Hoplites, Belemnites.
244
PALAEONTOLOGY
(b) Kutch basin
Cephalopods
Macrocephalites macrocepha/us*, Indocephalites Prio ·
Reineckia, Perisphinctes anceps*. Peltoceras athelata* •Rubertcentes,
· .-'
0 nono1ues, . . • oceras,
A ·a Terame//1ceras, Maymtes*, Torriuatisphinctes*
"7 ,
A•ax,·
,, oceras
sp, ocera_s, Streb/ires, Katro/iceras, Waagenia, Hildoglochiceras •
Virgatosphmctes, Ptychophylloceras, Be/emnites. '
PeJecypods Avicu/a, Astarte, Goniomya, Ostrea, Corbula, Trigonia.
Corals Montlivaltia, Stylina, Thamnasteria.
Cretaceous
(a) ffimalayan basins
CephaJopods Acanthoceras. Acanthodiscus, Dip/oceras, Holostephanus,
Perisphinctes, Hoplites neocomens*, Belemnites.
Pelecypods Cardium, Pseudomonotis, Ostrea, Gryphaea, Hippurites.
Foraminiferas Textularia, Nodosaria, G/obotruncana*, Orbitolina.
(b) Kaveri basin

Ceph.alopods · Turri/ires, Ham1tes, Scholenbachia, Belemnites, Acanthoceras.
Mammites, Pachydiscus, Baculites, Lytoceras, Phy/loceras, Ptychoceras,
Tetragonites, Placenticeras, Anisoceras, Desmoceras, Kossmaticeras,
Sphenodiscus, Parapachydiscoceras, Nautilus, Stoliczkaia.
Pelecypods Lucina, Gortriana, Trigonarca, bzoceramus, Protocardium. Trigonia,
Gryphaea, Exogyra, Alectryonia.
Gastropods ; Rostel/aria, Cyprea, Fulguraria, Hemifusus, Cerithium, Turritella.
Dentalium, Neptunea, Cancel/aria, Solarium, Lyria, Neithea.
Corals : Astrocoenia, Caryophyllia, Stylina, Thecosmi/lia, Isastrea,
Thamnasteria, Heliopora, Trochosmil/ia, Cyc/olites.
Echinoderms Stygmatopygus elatus*, Hemiaster, Cidaris.
Foraminiteras Globotruncana*, Rotalipora, Marginotruncana, Rotalia.
(c) Narmada basin (Bagh beds)

Cephalopods : Knemiceras, Narmadaceras.
Pelecypods : Plicatula, Protocardium, Trachycardium, Grotriana, Ostrea,
/noceramus, Neithea.
Gastropods Fulguraria, Fasciolaria, Turritella, Lyria.
Echinoderms Cidaris, Salenia, Cyphosoma, Hemiaster, Echinobrissus.
Brachlopods Malwirhynchia, Rhynchonella.
RECORD OF INVERTEBRATES FOSSILS FROM PHANEROZOIC ROCKS OF INDIA 245
(d) Kutch basin (Ukra bed and Trigonia bed)
Ccphalopods Australiceras, Cheloniceras, Colombiceras, Crioceras.
Pelecypods Trigo11ia vemrecosa*.
(e) Assam basin (Mahadek Formation)

Cephalopods Nautilus, Baculites, Stoliczkaia, Anisoceras, Puzusia, Belemnites.
Pelecypods Neithea , Vola, Chlamys, Atectryonia, Gryphaea, ltioceramus, Ostrea,
Trigonia, Nucula.
Gastropods Nerita, Natica, Turritella, Rostellaria, Cyprea, lyria Fusus, Mitra .
Echinoderms Echinoconus, Hemiaster, Stygmatopygus elastus* .
Brachiopods Rhynchonella, Terebratula, Concinnithyris.
Foraminiferas Orbitoides, Globotruncana*, Gumbellina, Obitoides, Siderolites.
Echinoids Stygmatopygus, Hemiaster, Euspatangus, Cassidulus.
19.4 CENOZOIC FOSSILS

Cenozoic marine invertebrate fossils are dominated by foraminiferas (mostly index fossils),
echinoids, molluscas, and corals. Palaeocene fossils are found locally within, Niniyur beds of
Kaveri basin, Nerinea beds of Pondicherry, Duddukura bed of Andhra Pradesh and also from
Assam. Some distinctive Palaeocene index fossils are :
Cephalopods : Nautilus danicus (KV), Pelecypod : Cardita beaumonti (D).
Foraminiferas : Globorotalia uncinata (P), G. trinidadensis (P), Globigerina pseudobulloides
(A), Miscellanea miscella (A), lockhartia (A), Nummu/ities thalicus (A), Discocyclina ranikoti
(A), Assilina ranikoti.
Eocene
Assam-Bengal-Kutch basin
Foraminiferas Nummulites atacicus, N. obtusus, N. stamineus, N. fabianii (A),
(all index fossils) N. beaumoti, N. acutus, N. irregularis, Discocylina sowerbyi,
D. dispansa, D. omphalus, D. javana, D. sella (A), D. assamica (A),
Dictyoconoids cooki, Faviania indica, Assilina exponens, A. granulosa,
A. spira, Pellatispira indica (A), Hantkenina alabammensis (A).
Echinoids Echinolampas.
Oligocene
Kutch basin
Foraminiferas N. jichteli*, Lepidocyclina dilatata*, Spiroclypeus ranjanae,
Miogypsina complanata*.
)
r
246 PALAEONTOLOGY
Miocene
Kutch basin
Foraminiferas Miogypsina dehaarti*, M. globulina*, Spiroclypeus ranjanae,
Lepidocyc/ina sumatrensis, L. tournoueri, Taberina ma/abarica*,
Austrotri/lina howchini, Globigerina, Orbulina, Bo/ivina.
Megafossils other than foraminifera of Cenozoic age may be divided into two divisions as
Palaeogene fossils and Neogene fossils. There is an overall similarity of fauna! types of different
localities and most of them are long ranging forms persisting upto the present day. Most of
these fossils are found from Tertiary beds of Kutch and Eocene-Oligocene beds of Assam.
Palaeogene
Echinoids Cyphosoma, Salenia, Eurhordia, Hemiaster, Schizaster, Cidaris,
Conoc/ypeus, Echinolampas, Euspatangus, Micraster, Breynia,
Clypeaster.
Gastropods Lyria, Cerithium, Conus, Turritella, Architectonia, Fusus, Murex,
Vo/uta, Mitra, Nerita, Natica.
Pelecypods Ostrea, Spondylus, Cardium, Corbula, Lucina, Pecten, Venus.
Corals Montlivaltia, /sastrea, Thamnastrea, Cyclolite, Trochosmi//ia,
Astrocoenia, Euphyllia, Meandriana.
Neogene
Pelecypods Pecten, Doscinia, Venus, Cyprea, Cyrena, Lucina, Pitar, Nuculana,
Arca, Glycimeris, Ch/amys, Nucula.
Gastropods Turrite/la; Mitra, Oliva, Voluta, Natica, Conus, Nerita, Murex.
Echinoids Breynia carinata*, Euspatangus, Echinolampas, C/ypeaster,

Echinodiscus, Schizaster.
Lower Miocene marine fossils having close affinity to Kutch fauna, are reported from
Andaman, Quilon bed of Kerala and also from Baripada bed, Orissil. Some fresh water-estuarine
invertebrates from Lameta bed (Cretaceous) and Deccan Intertrappean beds (Eocene) .are as
follows :
Ostracods Cypris submarginata*.
Gastropods Physa (Bullinus) prinsepii*, Melania, Turritella dispansa, Lymnaea,

Paludina, Valvata, Cytheria.
Pelecypods Unio, Modiola, Tellina, Ostrea, Lymnaea subulata.
* Index fossil.
Abbreviations used : K : Kas~mir basin, S : Spiti basin, KV : Kaveri basin, D : Duddukuru bed,
P : Pond1cherry Formation, A : Assani hasin.
PART - III
VERTEBRATE FOSSILS
Chapter 20
MAJOR SUBDIVISIONS OF VERTEBRATES
20.1 VERTEBRATE FOSSILS
The animals bearing dorsal nerve cords and axial notochords are included within the phylum
'Chordata'. There is a primitive group of chordates which possesses some sort of impersistant
or persistent but a poorly developed notochord (subphylum Protochordata). The more advanced
chordate has a skull, a bony or cartilaginous internal skeleton, and a dorsal bony/cartilaginous
rod enveloping and protecting the dorsal nerve cord, evolving into a jointed vertebral column
(subphylum Vertebrata). The skeleton also includes two pairs of appendages in the forms of
fins in fishes and limbs in tetrapods. The forelimbs of birds are modified to a pair of wings.
Fossil chordates are mainly composed of vertebrates, the history of which is rather long
and complex since their appearance in Ordovician. One characteristic feature related to the fossils
of vertebrates is their disarticulated nature. Many vertebrates, especially the larger terrestrial
groups, after their death and subsequent loss of the soft parts and ligaments tend to be
disarticulated along the weak bone-joints due to the effect of various erosional and depositional
processes. These fragmented parts of the skeleton are preserved separately either in different
places or within an area in a scattered condition. The toughest challenge to a vertebrate
palaeontologist is to collect these fossils as perfectly as possible and to reconstruct these pieces
in the laboratory to get a complete picture of the fossilized animal. For this, a palaeontologist
should have a comprehensive knowledge about the different groups of chordates and the
characteristic skeletal morphology of each.
20.2 GENERAL PLAN OF VERTEBRATE SKELETON (Fig. 20-la-b)

The generalized characteristic features of a vertebrate skeleton may be summerized as
follows:
(a) Vertebrate skeletons are internal, bilaterally symmetrical, may be cartilaginous or bony
or more commonly a combination of both.
(b) Vertebrates normally have their long axis of the body horizontally placed and any
deviation from this position represents a specialization (e.g. man).
(c) The sense organs are mostly concentrated at the anterior part, situated within a bony
case called skull which in turn is articulated with the dorsal vertebral column.
(d) The main axial skeleton (vertebral column) emerging from the base of the skull moves
dorsally towards posterior where it forms a tail. The entire vertebral column is flexible
and composed of several bony pieces called vertebrae.
(e) The middle part of each vertebra is called centrum and on either side of it are a pair
of processes (neural processes) which curve backward forming a dorsal arch called
neural arc/, through which passes the dorsal nerve cord . .A pair of similar processes
249
\
r
250 PALAEONTOLOGY
Vertebral Column
Pelvic Girdle
Pectoral Fins
(a) SKELETON OF AN AQUATIC VERTEBRATE
Tail
Vertebral
Column
(b) SKELTON OF A TERRESTRIAL TETRAPOD VERTEBRATE
..,___ Neural
Spme - - " ' 7 1 1
Hmflerus _ __....,··' ~---Femur
Fibula
Dorsal View Lateral View
(c) A SINGLE VERTEBRA
Carpals,---..,~~-- T11r~'!ls
Metacarpals Metatarsals
Fingers i)--- Fingers
i-- For Forelimb For Hindlimb-+

(d) LIMB BONES OF A TETRAPODA
FIG. 20- I: MAJOR SKEL~TAL ELEMENTS OF AQUATIC ANO TERRESTRIAL

VERTEBRA TES
, · Bt>WISI( . OF \ ERTEBRATES
251
al~ f r~n n. ventral nr_c h (liaemal arcl,) enclosing a passage for main dorsal artery. Two
p~tin-. 0f ~pm~s Un! dn-ected from each vertebra, ·o ne extending dorsally and the other
, ~nt .1\1~ L·alkd neural spines and laaemal spines respectively (Fig. 20-1 c).
\t ln m0~t f th ,ertebrates, there are two pairs of appendicular skeletons, each consisting
0f se,·eml pisces of bones. They form two pairs of fins in aquatic group and two pairs
~ i limbs in terrestrial group. In bird. the anterior pair consists of two wings while the
p stt.'rior pair remains as legs. In addition, the aquatic vertebrate usually bears median
fins ~md a prominent tail fin (combination of a pair). All these skeletons function for
stt~~ring and 1'alnnce for locomotion, for propulsion and in bird for flying. The paired
nppe ndi ular skeletons are articulated with the main axis by two bony girdles, the
anterior one is called pectoral girdle and the posterior one pelvic girdle (Fig 20- I a-d).
g) Ext~n ing latem.lly from vertebrae there are pairs of ribs, encircling and protecting the
inner rgans and soft parts. In all higher vertebrates (except one group) the mouth is
pnwi ed with an upper and lower jaw, often bearing teeth .
..\ frw ther characteristic features related to soft parts of the vertebrate are :
3) The ma.in nerve cord and all other internal organs are placed dorsally (ventrally placed
in case of invertebrate).
Respiration is done by means of gills (for aquatic animals) or lungs (for terrestrial group).
20.3 BROAD SUBDIVISIONS OF CHORDATES

A. Generalized classification of the phylum Chordata is given below mainly following the
s hemes of Colbert ( 1969) and Romer ( 1950).
Subphylum : Protochordata (Ord-Rec.) : Primitive chrodates with partial/temporary/
cartilaginous notochord.
Class : Hemichordata : Notochord at anterior part only. (e.g. Balanoglossus).
Class : Urochordata : Notochord at larval stage only. (e.g. Sea squirts).
Class : Cephalochordata : Persistant but cartilaginous notochord. (e.g.
Amphioxus).
Subphylum : Vertebrata (Ord.-Rec.) : Possesses a true notochord or vertebral column.
Section : Agnatha (Ord.-Rec.) : Jawless vertebrates, aquatic.
Class : Ostracodermi (Ord.-Dev.) : Primitive, armoured, jawless fishes. (e.g.
Cephalaspi.'i ).
Class : Cyclostomata (Sil.-Rec.) : Jawless, recent, fish-like animals with
circular mouth. (e.g. Lamprey).
Section : Gnatbostomata (Sil.-Rec.) : Vertebrates with bony jaws surrounding mouth.
Superclass : Pisces (Sil.-Rec.) : Aquatic, bearing scales and fins.
PALAEONTOLOGY
Class : Placodermi (Sil.-Perm.) : Extinct fishes with primitive jaws. (e.g.

Coccosteus).
Class : Acanthodii (Sil.-Perm.) : Spiny fishes. (e.g. Climatius).
Class : Chondrichthye (Dev.-Rec.) : Cartilaginous fishes. (e.g. Shark).
Class : Osteichthyes : Bony fishes. (e.g. Rohu).
Superclass : Tetrapoda (Dev.-Rec.) : Generally terrestrial with two pairs of
appendicular limbs.
Class r Amphibia (Dev.-Rec.) : Glandular, s~aleless skin.
Subclass : Labyrinthodontia (Dev.-Trias.): Enamel of teeth lobed
showing transverse folds giving a pattern of a 'labyrinth'.
(e.g. Euryops).
Subclass : Lissamphibia (Carb.-Rec.) : Modern amphibias. (e.g. Frogs).
Subclass : Leposondyli (Carb.-Cret.) : Sp_?on-shaped vertebrae. (e.g.
Microsaurs).
Class : Reptilia (Carb.-Rec.) : Scaled or armoured tetrapods.
Subclass : Anapsida (Carb.-Rec.) : With solid skull-roof. (e.g. Turtle).
Subclass : Synapsia (Crab.-Jura). : Mammal-like reptiles. (e.g.
Cynodonts).
Subclass : Diapsida (Perm.-Rec.) : Two skull-openings separated by
postorbital squamosal bones. (e.g. Crocodiles).
Subclass : Euryapsida (Perm.-Cret.) : A single skull-opening surrounded
by postorbital and squamo·sal bone. (e.g. Plesiosaurus).
Subclass : Parapsida (Trias.-Cret.) : One temporal opening surrounded
by post-frontal and supratemporal bones. (e.g. Icthyosau-rus).
Class : Aves : Frontal limbs transformed to a pair of feathery wings; body
feathery, a beak in front of mouth.
Subclass : Arachaeornithes (Jura.) : Jurassic toothed birds. (e.g.

Archaeopteryx).
Subclass : Neomithes (Cret.-Rec.) : All modern birds.

Class : Mammalia (Trias.-Rec.) : Tetrapods with hairs in body; young
suckled milk by mother from her milkgland.
Subclass : Prot~theria (Pleis.-Rec.) : Primitive, egg-laying mammals but

fossil record unknown before Pleistocene. (e'.g. Platipus).
MAJOR SUBDIVISIONS OF VERTEBRATES 253
Subclass : Eotheria (Trias-Jura.) : Primitive reptile-like mammals of
Triassic (?)/Jurassic. (e.g. Triconodonts).
Subclass : Metatheria (Cret.-Rec.) : Pouched mammals. (e.g. Kangaroo).
Subclass : Eutheria (Cret.-Rec.) : Advanced placental mammal. (e.g.
Man).
20.4 SOME DIAGNOSTIC CHARACTERS OF MAJOR GROUPS OF VERTEBRATES

A. PISCES
I. A general torpedo-shaped stream-lined body tapering on either end.
2. Skeleton bony except the shark-group (cartilaginous).
3. Gill-slits usually covered by opercular flaps except the sharks where gill-slits are open.
4. Homocercal or diphycercal tail (except shark where it is heterocercal).
5. Jaw attachment usually holostylic (amphistylic in shark); jaw usually with homodont teeth .
. 6. Bony skull composed of multiple skeletal elements; a single element in shark-group called
palatoquadrate.
7. Body usually covered with scales of variable types.
8. Paired and unpaired fins supported by soft spiny fin-rays.
9. A pair of external nostrils but ear internal.
10. Swimming bladder or air-bladder usually present (except shark).
11. Cold-blooded animals.
12. Respiration by gills.
I 3. Youngs from eggs without egg-shells.
B. AMPHIBIA
1. Skin usually glandular keeping the surface moist.
2. Skull articulated with vertebral column by two occipital condyles.
3. Limbs tetrapodous pentadactyle type (may be less).
4. Teeth in jaws (homodont) or may be absent.
5. Vertebrae procoelous or amphicoelous type.
6. Jaw suspension autostylic.
7. Larvae passing through a fish-like 'tadpole-stage' in water.
8. Cold-blooded animals.
9. Respiration by lungs in adult.
10. Youngs from eggs without egg-shells.
254 PALAEONTOLOGY
C. REPTILIA
I . Body covered with horny scales, armours.
2. Skin dry without any gland.
3. Limbs tetrapodus-pentadactyle type, digit terminals with claws.
4. Skull with a single median occipital condyle for articulation with vertebral column.
5. Mandible with several skeletal elements, articulated with skull by quadrate bones
(autostylic).
6. Jaw usually with teeth, normally homodont type.
7. Vertebrae gastrocentrous, procoelous.
8. Ribs forming a true sternum.
9. Respiration by lungs.
l 0. Cold-blooded animals.
11. Youngs from eggs with egg-shells (amniotic).
D. MAMMALIA
l. Skin usually covered with hairs (except whales).
2. Glands present on skin.
3. Mammary glands functional in females for nourshing the youngs.
4. External ears present.
5. Teeth normally heterodont (except whales).
6. Skull with two occipital condyles for articulation with vertebral column.
7. Reduction of number of bone. elements in skull.
8. Lower jaw with a single bone (dentary) on each side; Jaw suspension craniostylic,
streptostylic.
9. Vertebrae gastrocentrous, acoelous types.
10. Limbs tetrapodus; pentadactyle type, feet plantigrade, digitigrade or un 1· ad . te · I
· · prov1·de d wit
of d1g1ts · h c 1aws, na1. s or hoofs. gu 1gr e, rmma s
1
11. Warm-blooded animals, respiration by lungs.
12. Youngs have direct birth.
E. AVES
1. Skin covered with an exoskeleton of feathers.
2. Forelimbs modified into a pair of wings 'd d .
clawless digits; hindlimbs used for w lk" provt e_. with fe~ther~ for flight with three
digits with claws. a mg, perching or sw1mmmg, each bearing four
r , •
'._ .
.,. ..
'
..'. ~
MAJOR SUBDIVISIONS OF VERTEBRATES 255

3. Skin-gland absent.
4. Bones spongy, light and hollow.
5. Skull monocondylic having a single occipital condyle.
6. A pair of horny beaks, no teeth.
7. Lower jaw with several bones~ jaw suspension streptostylic.
8. Vertebrae procoelous, amphicoelous or heterocoelous.
9. A number of vertebrae of anterior part found to form synsacrum white posterior caudal
vertebrae fused to form a pygostyle.
10. Sternum broad usually with a longitudinal ventral keel for attachment of flight muscles.
11. Digital bones often found.
12. Warm-blooded animals, respiration by lungs.
13. Youngs from eggs with egg-shells.
Chapter 21
OUTLINE OF MORPHOLOGY OF
S01VIE SKELETAL ELEMENTS OF VERTEBRATES
21.1 SKULL
A. Three cmb)onic components
Th' skdetal fran1ework of the head region of a vertebrate is called skull. The structure of
skull becom~s progressively complex with the appearance of more and more advanced gro~ps
f v~rt "brat s. A primitive jaw less vertebrate had an incomplete cartilaginous box enclosing
th' brain. In lowe r groups of fishes the skull is a cartilaginous cover of brain with isolated
lower and uppe r jaw. But in higher vertebrates the skull is bony with welded upper jaw and
suspend~d lower jaw. Skull develops from three embronic components viz. (i) clzondrocranium
is c nsisting of a braincase and cartilaginous capusles of otic, olfactroy and optic region,
tii) spla11c/mocra11ium is derived from the cartilaginous visceral skeleton which is replaced and/
r ne\, ly invested by bones in higher vertebrates (iii) dermatocranium is consisting of dermal
bone that later become attached with the chondrocranium. This is absent in lower groups of
fishes including sharks.
B. De, elopment of embyonic components (Fig 21-1)

After the formation of the initial chondrocranium from embyo, the mesenchyme cells form
a membrane covering the brain where appear two pairs of cartilaginous plates, the parachordals,
,~nd the !'re~lwrdals. lying b~low the po~terior and anterior to the brain respectively. Ring-
It ke carttlagmous elements anse surrounding the three sense organs and there are three pairs
of elements, one each for optic, olfactory_ a~d otic c~psules. Parachordals and prechordals
gradually grow larger_and fuse along the mi.dime of brain forming a single basal plate leaving
a small gap at the middle calle~ hypophys,al Jenestra for the position of pituitary gland. In
the ne xt stage olfactory and ot1c capsule grow larger to join and fuse w·th th b
formmg · ·1 · h d · 1 e asa 1 p1ate
a cart1 agmous c on rocramum as a continuous floor of the b · ·
. ram. 0 pt1c capsu 1es
however remam free and usually become fibrous to form sclerotic f th . . .
b ·1·
mo 1 1ty. Af h t· · t· I ·1
ter t e ormat1011 o t 1e f oor, the basal plate beoins t ° e eye permitting its
. . . . o o grow upward along the
side of the bram and 111 shark 1t completely covers the brain a .. b II .
· h b. - J h~ · .
8 ut 111 ot er verte rates, on y t e postenor or occipital region of brain s a ox ca ed neurocramum.
cartilaginous bones, and the rest has only a dermal membr· Ch d g~ts roofed over by
· ,
- ·ct · • · · f h ·k ti · ti"
g<tp at m1 -postcno1 o t e s u c.i ~d forame11 magnum th ane. hon rocranmm
. . leaves· a large
roug which spinal cord emerges.
The splanclmocranium or a visceral skeleton grow, f. h .
wall of pharynx between the gill-slits for their suppo t Th
s 10m t e splanch
·
od
. me mes erm m t e
· h
be several pairs of gill-arches as bars of cartilage r · : hese ~re called g1ll-arclies. There may
. . . on e1t er side Ea h · · · .
tormmg some horse-shoe shaped visceral arclies Th fi , . ·. · c pair 1s Joined ventrally
. . . e irst pair is then . d.b l
1s hyoid arch and the rest are called brancli,·al a I h · ia11 ' " ar arc/a, the second
. .
all Jaw-bearing vertebrates the madibular arcl,
re ,
,est at supp0 rt ·11 f
~ g1 s o lower vertebrates. In
goes to ,orm uppe d .
r an 1ower Jaw for supporting
.. 256
~
c::::
-,
Opt ic capsule
-
r-,
t"l,
a
""I'\
~
Lens I 19 0
Midbrain- / a, ~ \ _Hypophysis ~
~
Hindbrain - .- ll! .... - ::i:
Auditory vesicle Hypophysial 0
t""-
Notochord I llti'RD ~ 1::,#f I Otic capsule It 0
fenestra C)
Parachordal • - C7 ~
cartilage (a) APPEARANCE OF CARTILAGES (b) FORMATION OF ETHMOID "'Basilar plate 0
(c) FULLY GROWN ~
IN HEAD OF EMBRYO AND BASILAR PLATE
CHONDROCRANIUM c,,
FlG. 21 - 1 : THREE STAGES OF DEVELOPMENT OF CHONDROCRANJUM a
~
t:,
Orbit
Chondrocranium vi
;:>;:
C'?']
Ligament r-
rr;
Basal process ....;
::t,:
C'-
Basal process rr;
it'CJ H)omandibuiar C'-
t:,
Palatoquadrale ~
rr;
<'.
....;
AMPHISTYLIC vi
Orbit Spiracular ligament Basal _ _ _ __, Orh1t a
jll"OCl!~S .........,...._ Palawquadratc ~
Ethmopal~11ine
Ligamcnt ~ - - - - - - - Cranium Otic process ,.,
~
yomandibular ~
- - - - Columdla or stapcs ""
r:;
P:ilatoquadmtc :::::
~
:::::,..
--;
M
V:
..&,Z I lyoid arch
HYOSTYLIC
AUTOSLYLIC
FIG. 21 - 2: DIFFERENT TYPES OF JAW SUS PEN ION
N
V,
'-l
,.
•, __
258 PALAEONTOLOGy
the mouth. Hyoid arch in higher vertebrates functions as a support of jaw. Branchial arches i
tetrapods are highly reduced forming hyoid apparatus and cartilages of larynx. n
Formation of a skull stops at cartilaginous stage in agnathas and in lower groups of fishes·
but in bony fishes upward, dermal and cartilage bones become incorporated with the skull:
Dermal bones appear in head region of bony fishes as scales. In higher vertebrates several
cartilaginous bones are added in parietal region, olfactory and otic capsuls and also in upper
and lower jaw for performing various other functions. The different segments of the skull are
shown below and the chief skeletal elements of each segment in different groups of vertebrates
are shown in the Table- I 4.
Chondrocr~nium cartilages
I. Occipital segment most posterior part of skull
2. Parietal segment anterior to occipital
3. Frontal segment anterior to parietal
4. Olfactory capsule smell organ (nose)
5. Otic capsule auditory organ (ears)
6. Optic capsule vision organ (eyes)
Splanchnocranium
7. Lower jaw
8. Upper jaw
9. Hyoid arch
Dermatocranium : dermal bones
21.2 JAW SUSPENSION (Fig 21-2)

The method by which the upper and lower jaw of vertebrates are suspended from the
chondrocranium · is an important feature of the skull and often helps in the recognition of
different groups of vertebrates. The major types of jaw suspension are as follows :
(a) Autodiastylic : Jaws are merely attached to the skull by ligaments; hyoid arch occurs
as branchial arch. e.g. early fishes.
(b) Amphistylic : The basal and otic processes of upper jaw are attached to chondrocranium
by ligaments; hyomandibular of hyoid arch is attached to the skull at one end and from
its other end both the jaws are suspended. Thus it is a double suspension of jaws by
both bones and ligaments. e.g. primitive sharks.
(c) Hyostylic : Both the jaws are suspended from hyomandibular which in tum is attached
to chondrocranium. e.g. modern sharks and bony fishes.
(d) Autostylic : Upper jaw is completely fused with skull while quadrate of upper jaw
articulates with articular of lower jaw. e.g. some bony fishes and tetrapods (except
mammals).
(e) Craniostylic : The articulation between lower and upper jaw is done by two dermal
bones, squamosal of upper jaw and dentary of lower jaw. e.g. mammals.
-
OUTLINE OF MORPHOLOG Y OF SOM E SKl:.LE'/'A/, J:,'U ~'Ml:NT\' OF VEUTEIIRATES 259
(f) Streptostylic : In most f the birds and in a few reptiles articulali on is like au1os1ylic
suspension but quadrate instead f being fixed firmly is loosely attached to articular so
that it is movable at both ends. e.g. Lizard, snake and hird.
21.3 DIAGNOSTIC FEATURES OF SKULi S OF TETRAPODS (Fig. 21 -3)

A. Amphibia
I. Several bones found in fish- skull arc absent ~uch as, pleurosphcnoids, post-parietal,
prefrontal, post frontal, lacrimal, ectethmoid, pterot ic, ectoplerygoid, hyomandibular,
symplectic and gill covers.
2. Skull more flattened, especially the ventral side.
3. Embryonic chondrocramium partly replaced by cartilage bones.
4. Basioccipital, supraoccipital, exoccipital, basisphenoid and presphenoid not ossified.
5. Otic capsule with a ventral aperture, the fenestra ova/is onto which the stapedial plate
of columella fits.
6. Dorsal surface solidly covered by dermal bones leaving opening for nares and eyes.
7. Two occipital condyles.
8. Jaw-suspension autostylic mostly without tooth.
B. Reptile
l. Entire chondrocranium ossified except the naso-ethmoidal region.
2. More dermal bones than amphibia.
3. Parietals fused together (except in chelonids) with interparietal foramen.
4. Between the orbits present an inter-orbital region and behind it one/two temporal fossae
(except in chelonids).
5. Parasphenoid and basisphenoid fused.
6. Four occipital bones surrounding the foramen magnum with a single occipital condyle.
7. A transverse bone (ectopterygoid) between maxilla and pterygoid, and a vertical
epiterygoid running from preotic to pterygoid.
8. Each half of lower jaw with one cartilage and five dermal bones.
9. Autostylic or streptostylic jaw-suspension; jaws with homodont, sharp and conical teeth.
l 0. Quadratojugal absent; hyoid arch forming columella of middle ear from its hyomandibular
part and the rest part forming hyoid.
C.Aves
I. Comparatively larger and lighter for presence of pneumatic bones; bones of skull fused
together with practically no suture.
2. Large arched cranium; orbit large, continuous with a temporal fossa supported by a thin
interorbital septum; each orbit composed of a ring of dermal bones, the sclerotics.
3. A large pointed beak formed by premaxillae and dentaries, covered by horny sheath,
without any teeth.
260 PALAEONTOLOGY
Nnsal---.. Frl)t1\al - - - -,S upruoccipital
(l r~Ill l\lc iII n l'n:opcri:ubr
Mnxillu
-;=-i'C":::1-- Opcri:ulur
, ..:........- Sul:opcrculur
- - - - - - - Quallratc
---Nasal capsule
Angular
Jug. i i - - - - Prc1m1xilla--::,,1::t~
Articular External
FISH (lalcral \·icw) naris -~i;__-flllffl,:lill
Nasal--,-..,...-~
Ptcrygoill _ _,_,_""111111 ParasphcnoiJ
Qumlrnto-j11gal Ptcrygoid
Orhi tal foss;1
Qu.idrato-jugal
Post-orbital Supraurhital • l\uu1 h>f) \. .tp~uk
fnr.11111: 11 l' Columclla

111ag1111m : cc,nllylc xoccipital
I' Ventral
Dorsal View
.
AMPHIBIA
View
~"!S'~r--Maxilla
S"-:::;;;~~~,pL.;L._ l.acrymal
Occirital Dcntary Optic Iossa
---~fas1sphcnoiJ
-----Pam:tal
Supra.:ccipital
Articualr _ _ _ __,
o, hiw~pli\.'11oiu l'lcl) guiu Squamos:11
:-.fax iII a --,"-,-,,r..,~~ Zygomatic
REPTILE (lateral view) Pal.,tinc ..-"-,!~t----arch
--7"---:::~~---...-~~ Basioccipital
Occipital
---eondylc
Orhilal foraman
Incisors----..:~,----
Zy~1m1atic pmccss Frunlal Tympanic bulla
Squamosal lntcr-orhital ~cptmn
Pa rieta1---,
Mol;n tcc1h
O.:dpital Nasal chamber
MAMMAL(latcral vicwJ
condyle l'rc111axilla
Ma'\illa
JuHal
Dcntar\'
Quadratc--- .__ _ _ _ _ _ _ _ _ _Vomi:r
'ata1111c
Quudratojugitl _ _.J,.?...,"'--_..r
Jugnl Articular Angular
A YES (lutcrul view)
FIG. 21 - 3 : MAJOR SKELETAL ELEMENTS OF SKULL OF VERTEBRATES
OUTLINE OF MORPHOLOGY OF SOME SKELETAL ELEMENTS OF VERTEBRATES 261
4. A single occipital condyle formed by basioccipital and exoccipital; skull at right angle
to vertebral column.
5. Parietal and frontal very large forming the roof and much of the lateral wall of cranium.
6. Auditory capsules sink inwards with fused bones.
7. Jaw suspension streptostylic.
8. Hyomandi_bular b?ne forming the 'columella' and 'stapes' of middle ear, rest of hyoid
arch forming hyo1d apparatus along with first branchial arch.
D. Mammals
I. Los_s of many bones such as parasphenoid, quadratojugal, prefrontals, post-frontals, post-
orb1tals, quadrate and all bones of lower jaw expect dentary.
2. Fusion of basisphenoid and alisphenoid; fusion of bones of temporal region; in some, all
occipital bones fused to form one bone.
3. Dermal bones dorsally joined with cartilage bones and the joining lines form sutures.
4. Some cartilage bones and membrane bones fused together.
5. The cranium expanding dorsally and laterally to accommodate large-sized brain; its cavity
closed in front by a cribriform plate.
6. Bones of facial region closely united and in higher forms facial region lying below the
cranial region instead of in front of it.
7. Skull with two occipital condyles for articulation with vertebral column that lying at right
angle to it.
8. Bones of otic capsules fused to form a petrosal enclosing the inner ear.
I
'I 9. Petrosal, in some forms, fused with squamosal and tympanic bone to form a petrous region
[ of the temporal bone.
1
' 10. The tympanic part of the temporal bone forming an external auditory meatus.
11 . Jaw suspension craniostylic.
12. Orbit formed by lacrimal, maxilla, orbitosphenoid, palatine and also ethmoids.
13. A bony palate formed by premaxillae, maxillae, palatine and pterygoid bones.
14. Palate pushing the internal nares backward and forming a floor below the respiratory
channels separating them from buccal cavity, enabling the mammal to chew and breathe
at the same time.
15. Two respiratory channels separated by an internasal septum formed by mesethmoid and
inside the channels turbina/s formed by ectethmoid.
16. Lower jaw with dentary only; teeth on both jaws heterodont type.
17. Hyoid arch partly forming hyoid apparatus and partly thyroid cartilage of pharynx, and
parts of internal ear; the remaining visceral arches forming thyroid, epiglottis etc.
21.4 VERTEBRAE AND VERTEBRAL COLUMN

Vertebrae of various vertebrates are different. Even, within a group, vertebrae located at
different parts of vartebral column are dissimilar. But all vertebrae conform to a basic plan
(Fig. 21 -4a).
- -· - L ' - .. J.: . •»· ., •' l ~" •
PALAEONTOLOGY
R--------Neural Spine
Tuberculum---.
~-----Neural Arch
~----·Neural Canal
Diapophysis
ffS:~----Hypapophysis
Ccnturm
i,...---- Hacmal Arch
-"4J.-----Haemal Canal
l~----- Hacmal Spine
(a) A TYPICAL VERTEBRA
Anterior
Posterior
~
f A(~
Neural Arch
Neural Canal - -......!It.Ii
Convex ,': .:. ~Cc~
Posterior Face._.,. ·;:~: :; C--vc
At Both Ends
lntcrcenlr\lm PROCOELOUS AMPHICOELOUS
Ccntrum Saddle-Like
Ccntrum With Convex Ccntrum Flat On Eithc:r
Anterior And Concave HETEROCOELOUS
Posterior Side
OPISTHOCOELOUS ACOELOUS
(b) VERTEBRA-PAliERN BASED ON TYPES OF CENTRUM
FIG. 21 - 4 : MORPHOLOGY OF VERTEBRA
oUTLINE OF MORPHOLOGY OF SOME SKELETAL ELEMENTS OF VERTEBRATES 263
A typical vertebra has a c~lindrical body ca11ed centrum. On either of it is a pair of processes
which curve backward formmg a neural arch that encloses the neural canal through which
dorsal nerve-cord passes. Neural arch i~ projected dorsally into a neural spine. At lower part
of centrum is a haemal arch enclosing haemal canal which is the passage of dorsal artery and
vein. The haemal arch is also produced back into a haemal spine. Besides, other type of
processes and articulation zones may be found on a vertebra. Zygapophyses mark the
articulation zones for successive vertebrae. Some processes develop transversely (transverse
processes) such as parapophyses and diapophyses from neural arch or ·centrum serving
articulation of ribs in thoracic zone. Hypapophysis is a mid-ventral projection. Centra of
successive vertebrae within the vertebral column are placed end to end in a row and the shape
of the two end surfaces of each centrum is _of importance for articulation. There are five different
types. (Fig. 21-4b).
(a) Amphicoelous : Centrum has concave surface on both sides and thi~ is supposed to be
the most primitive as found in all fishes.
(b) Procoelous : Centrum surface is concave anteriorly and convex posterior_Jy as found in
amphibias and reptiles.
(c) Opisthocoelous : Centrum surface convex anteriorly and concave posteriorly; found
loca11y in a11 groups of vertebrates except fishes.
(d) Heterocoelous : Vertebra has centrum transversely saddle shaped, the anterior end is
convex dorso-ventrally but concave sideways and posterior end is just opposite to this
as found within cervical vertebrae of birds.
(e) Acoelous : Centrum surface on either end is flat as found in mammals.
21.S SOME CHARACTERISTIC FEATURES OF VERTEBRAE OF DIFFERENT

GROUPS OF VERTEBRATES (Fig. 21-5a-d)
(a) Most fishes exhibit vertebrae all similar looking, amphicoelous type.
(b) In most amphibias, all vertebrae, except the first (at!as) and last (caudal), are similar,
procoelous type. The last vertebra representing caudal region is projected into a long
triangular urostyle.
(c) In reptile, the vertebrae within the vertebral column can be divisible into four regions
depending on their different appearance and locations. From anterior to posterior these
are : cervical, thoraco-lumber, sacral and caudal; all vertebrae are procoelous.
. ..
(d) In aves, vertebral columns is divisible into four region such as cervical, thoracic, sacral
and caudal, each with different forms; mostly heterocoelous. The last thoracic, few
lumber, sacral and caudal vertebrae are fused forming a synsacrum. The posterior caudal
vertebrae are projected into a smalJ plough-shaped pygostyle.
(e) The vertebral column of a mammal is usually divisible into following zones : cervical,
thoracic, lumber, sacral and caudal. Most of the vertebrae are acoelous type. Sacral
vertebrae are fused to form a sacrum.
Palac(Geo)wP-3 4
.j
264 PALAEONTOLOGY
Neural Arch Postzygapophysis

-----Neural Spine
Prezygapophysis
L-•---- Transverse
Process
----- Postzygapophysis
Neural Canal
~ - - - - - Ccntrum
Pl,sl zyg.apophysis - - - - - Centrum
Posterior View Dorsal View
(a). AMPHIBIA
Neural Arch
Neural spine - - -
Prczygapophysis
Postzygapophysis
Transverse
process
Neural CanaJ
Thoraco-Lumbcr (Dorsal View)
Caudal (P.osterior View)
(b). IH:PTII.E
11111/l.----Neural Spine
Neural Arch--....
- - - Postzygapophysis - . - - - Neural Spine
Transerse Process Neural Arch
Transverse Process
Centrum
(c) BIRD (Anterior View)

· - - - - Centrum
Thoracic Verteb~ (Anterior View)
(d.) MAMMAL
FIG. 21 - 5: PAITERN OF VERTEBRAE OF TETRAPODS
ouruNE OF MORPHOLOGY OF SOME SKELETAL ELEMENTS OF VERTEBRATES 265
prccoracoid Epistcmum
~ - - - - - - - - - Omostcrnum
t--------Claviclc
~;a.----- Supruscupulo
c;;...a..-- Scapuln
-~-~il\U~~~-- Coracoid
T---------Mcsostemuna
Epicoracoid - - - - - - - Wr----------Xiphistcmum
Epistcmum
--w~-- Epicoracoid
r-1'---.:£.lt..a.-- Clavicle
~~-.... Suprascapula
Scapula
r------ Glcnoid Cavity
..-,..o.r,
" ' - - - - - Corncoid
"'.':!a!Plilr--------stemal Plate
Stemul Ribs
(b)
Metastcma\ Process------------,. - - - - - Glenoid Cavity
Oblique Xiphoid Process----------,dl.a ---Scapu\a
Lower Xiphoid Process-----.
Mctastcmum
s,ema\
Kcc\/Crest Pectoral Girdle (Hal() Outer View
Suprascapula (c) Bllm
Vertebral Part Of Rib
. A ~ - - Manubrium
(Prestemum)
~. ~csostcmum~"-•Capitulum
,.I (With 5 Picccss
';, . A Mammalinn Rib
Xphistemum Of Stcmabrac)
: \
'',, ~ - -Xiphoid Canilagc

,\
Glcnoid Cavity Sternum (Ventral View)
Stcmum And Pctoral Girdle
(d) MAMMAL
_-,
I.
I
l:
·'
'
FIG. 21-6 : STERNUMS AND PECTORAL GIRDLES OF TETRAPODS
:,
PALAEONTOLOGY
266
Cavity For
Trochlea
Radio-
Ulna--....,
.--Carpals
Ulna
Radio-Ulna
Humenis Hand Bones
(a) AMPHIBIA
- - Phalanges
..."---Claws
(b) REPTILE
Fuse<l Metacarpals
Hnnd Bones
Humerus Ra<lius-Ulna
(c) BIRD
Humerus Rndius-lJlnn Hand Bones

(d) MAMMAL
FIG. 21 -7 : BONES OF FORELIMB OF VERTEBRATES
ouTL!NE OF MORPHOLOGY OF SOME SKELETAL ELEMENTS OF VERTEBRATES 267
zt.6 VERTEBRAL COLUMN AND FISH-TAIL (Fig. 21 _9 a)
In the fish the shape of the tail is g
.
II . . ·
enera y correlated with the nature of terminal portion
f the postenor part of notochord/vert b 1 I . ·
o . e ra co umn. Four types of fish-tall may be recognised
accordingly.
(a) Protocercal : ~his is possibly the most primitive type where notochord extends straight
to the ~n~ ~f tail and the caudal fin becomes symmetrical and rounded e.g. cyclostomes,
and pnm1ttve fishes.
(b) D~phycercal : ~ertebral column is ending before the posterior terminal point making
tail-fin symmetncally rounded. e.g. Dipnoi, Latimeria, Polypterus.
(c') Heterocercal : Vertebral column bends upward dorsally in the posterior-most region of
tail so that the caudal fin exhibits two unequal halves, a narrow and elongated larger
dorsal lobe and a smaller broad ventral lobe. e.g. sharks and some bony fishes. In some
fishes it may be reverse e.g. Cypselurus.
(d) Homocercal : Terminal point of vertebral column is short of posterior end but it bends
upward so that dorsal lobe of fin disappears and the ventral lobe is marked by two
equal halves, the end of which may be truncated, rounded, or medially notched. e.g.
most of the modern bony fishes.
21.7 RIBS (Fig 21-6d)

Thoracic region of many vertebrates except the lower groups is provided with ribs growing
in pairs from some anterior thoracic vertebrae. Ribs of cartilaginous or bony fishes are either
fused or articulated with axial column. In fishes, there are two kinds of ribs, dorsal and ventral.
Some fishes have one pair of ventral ribs. Some fishes have either dorsal or ventral ribs hut
some possess both the types. In tetrapods there is only one pair of ventral ribs from each
vertebra, joining ventrally with the mid-ventral sternum bone and dorsally with thoracic
vertebrae by two heads. The upper tuberculum articulates with diapopliysis of the neural arch
and the lower head capitulum articulates with parapophysis of centrum. Besides, at the bifid
nature of vertebraside of a rib, in case of birds, there is an additional backwardly projected
process called uncinate process.
21.8 STERNUM (Fig. 2 l-6a-d)

Sternum is an elongated rod-like bone along mid-ventral region of th~rax, absent in fishes
but present in most of the tetrapods. It is actually a series of cartilages or bones or both. In
tetrapods sternum articulates with pectoral girdle, dorsally and ventrally with the ribs forming
a thoracic busket. Some characteristic features of sternum of each group of vertebrates are as
follows :
(a) In an amphibia, sternum exhibits two parts, an inverted Y-shaped bone (omosternum)
with an expanded cartilage head (epistemum) anterior to clavicle and a backward portion
posterior to ·coracoid with a rod like initial part (mesosternum) and an expanded
cartilaginous head called xipliisternum.
(b) In a repitle, sternum is a rhomboidal and flattened cartilage (sternal plates) behind and
between the coracoids with three pairs of short and free thoracic ribs.
PALAEONTOLOGY
268
Pelvic Bone
(lschopubic Rar)---Nf'i:Y"
Femur-----. lschium
Radials.---c:::::::::11m~-
Basipterygium ----ill:J.
Pelvic ------fiffH-ll+i~
Cartilage
Somactids
Pelvic Girdle
Pelvic Fin,--......,1 {one half in lateral view)
Astragalus-----:IIIY
>.a~-- Calcancum
Tarsals:---tll
(a) FISH 0 Distal Epiphyis
0
~?t' 0~--~ Metatarsal
~) I t Phalanges
Epipubis _ _ _...., Hind Limb Attachment
With Pelvic Girdle
(b) AMPHIBIA
Ilium - - -
, "' ~---Tibia
l,&,l..""""--Fibula
Tarsnl:; Acetabulum---~
Metatarsal
Phalanges lschiun-----'~
Pubis
Pelvic Girdle (Left Half Ventral View)
Acetahulum lschium (c) REPTILE
Preacetahular Ilium Pubis

Pelvic Girdle (~.ft half outer vii:" 1
(d) Bird
Acetabulum Tarsals
(e) MAMMAL
flG. 21 - 8: PELVIC GIRDLES AND HIND LIMBS OF VERTEBRATES
_.-/. '. ·
< l ·l f I: )F M RPHOLOGY OP S ME ,
. , . . . ' Sl<ELF.TAL ELEMENTS OF VERTf:BRATES 269
c) Sternum m a bird 1s well-dcvclo d
callt~ br~ast Mne. Here ·tJ .Pe tu, a broad bone lying in the breast region, also
s emum as a boat-shaped b . , d f . ,- . .
r late (ma11ubrium) an t . u one compose o . an anterior vertical
, an enor lateral proicction ( la l I
tnin,•ular stout bony l· t ( . J me !t erna process) and a broad
'
0
- • P a e can,aa or sttrnal t) · · ·
ventrnl side bearing tw . .. . . cre.f proJectmg vertically downward from
o pnu s of x1pho1d processn T-h ·t f b' d .
functions for attach t f . · e s ernum o 1r main ly
men o Vanous muscles used for flight .
d The mammalian sternum is m d f .
. . 'd · a c up O a senes of bones fixed together and arranged
,n rows ymg mt -ventrally with
1 .
_ . _ . an antenor prtsternum , middle mtsolternum (with
fi ve pieces tuscd to form st b ) d . . .
trna rate an a posterior x1plusternum which is long with
an expanded head of cartilage (xiphoid cartilage).
2 l.9 BONY GIRDLES AND LIMB BONES

Pai~ed limbs (~ne antcri~r and .one posterior) and the associated bony girdles by which they
are 3rt1cula~ed with the axial skeleton in different tetrapods are very similar in appearance as
regards their bone-elements.
Ec.ctoral girdle (Fig. 21-6a-d), placed anteriorly on dorsal side and articulated with anterior
sid of sternum is composed of several bony elements, such as a scapula, a suprascapula, a
precoracoid and a coracoid in each half. A glenoid cavity is present in between scapula and
racoid. The forelimb is composed of three parts : humerus, radius-ulna and hand bonts.
Fig 20-1 d, 2 l-7a-d). The anterior most limb-bone humerus becomes articulated with the
pectoral girdle at glenoid cavity. The distal limb-bone is composed of two parts called radius
and ulna in reptiles, birds and mammals but in amphibia they are fused to form a radio-ulna.
The digital part of the bone is composed of carpals, metatcarpals and a number of phalangts
forming the digits. Pelvic girdle (Fig. 21-Sa-e) placed posteriorly on dorsal side. has three bones
on each side which are ilium, ischium and pubis. At their junction is the foramen acttabulum
for the articulation of the proximal posterior limb bone/em11r. The two distal limbs bones tibia
and fibula (tibia-fibula in amphibia) are articulated with femur. At anterior to tibia-fibula are
a number of digital bones, tarsals and mttatarsals and pl,alanges. Limbs of tetrapods are mostly
pentadactyl type (five digital). In amphibia there is a third compoment of limb in between the
digits and tibio-fibula composed of two bones astragalus and calcaneum.
Bird has a different pattern of pelvic girdle where pubis bone is rotated backward to become
parallel to ischium and ilium and the ilium becomes greatly extended both anteriorly and
posteriorly to acetabulum. This gives pelvis a tetra-radiate structure in contrast to the common
triradiate structure of most of the other tetrapods. However, some extinct group of reptiles.
t.he omithischian dinosaurs, exhibited bird-like hip-joint.
Terrestrial mammals show three types of digital posture of hand and feet for locomotion
(Fig. 2 l-9b) :
(a) PJantlgrade : It is supposed to be the most primitive type in which the entire hand and
foot area is in contact with ground as found in bear, elephant and man.
(b) Digitigrade : It is commonly used by animnls for getting incrensed s~ed iri locomotion
where only digits arc placed on the ground. The wrist and the ankles are lifted above
at the time of locomotion as found in dog, cat and many other cumivor-us.
N
...:,
0
Diphycercal
Protoccrcal
Hypoccrcal
Heterocercal
(n) TYPES OF CAUDAL FlNS IN FISH
I
,, ' )
) \
\
'"
-- _...... ---- --· \) \
, .,. -- - ___ ,
Plantigrade
Digiligrndc
(b) FOOT POSTURES OF MAMMALS Unguligradc
~
s:
~
FIG. 21-9: CAUDAL FINS OF FISH AND FOOT POSTURES OF MAMMALS <:
~
8
C)
"'<
<)UTUNE OF MORPHOLOGY OF SOME SKELETAL ELEMENTS OF VERTEBRATES
271
( ) Unguligrade : It is an extremely , · r d · ·
· ti · f d' . specia ize type of d1g1t-alignment for locomotion in
1 1
w 11c 1 on y 1e tips o 1g1ts are ·
I ·
. . . m contact with the ground surface. Normally such digital
ups are provided with hoofs such as found m · h orse, d eer, catt Ie etc.
21.10 STAGES OF EVOLUTION OF LIMB BONES

Stu<ly of evolution of limbs exhibits three stages (Fig. 21-10).
(a) ~rchiptcrygium typ~ : It is the most ancient type seen in ray-finned fishes where a
hn bears a central axml skeleton from which diverge small radial elements called tins-
rays.
(b) ~ch~hyoptery~ium type : This exhibits the beginning of tetrapod-limb evolOtion and
md1cates an intermediate stage between a true fish and a tetrapod (as exhibited by
crossopterygians or lobed-fin fishes). Usually these fins are attached to some muscular
lobes. The skeletal elements inside the lobe are aligned in three rows : a proximal row
comprising a single basal bone radius (comparable to humerus/femur of a tetrapod), a
middle row with two radial elements (comparable to radius-ulna/tibia-fibula of a tetrapod)
and the distal-most row with radial fins (comparable to tarsals and metatarsals of a
tetrapod). The whole structure is articulated with axial skeleton foreshadowing the girdle
of tetrapods.
(c) Cheiropterygium type : This is exhibited by the first group of amphibia represented
by labyrinthodonts showing a primitive girdle and all such bones like humerus/femur,
radius-ulna/tibia-fibula and the digital bones.
21.11 TEETH (Fig. 21-11 a-c, 2 I- l 2a-d)

Vertebrates possess two types of teeth. Some primitive vertebrates develop epidermal teeth
which are small conical structures lying above the dermal papilla but are derived from epidermis.
True teeth or dermal teeth are supposed to be derived by transformation of placoid scales of
fi sh. Recent view is that both teeth and placoid scales are modified remnants of bony dermal
plates of ostracoderms and placoderms. In lower vertebrates, teeth may be replaced several times
in life which are called polyphyodonts. But most of the mammals are diphyodonts having
only two sets of teeth, one milk set replaced by a permanent set each with a fixed growth
period. A s regards the nature of fixation of teeth, there are few types (Fig. 21-11 a). The
primitive pleurodont teeth are fixed to a shelf at inner margin of jaw-like indentation. In many
advanced vertebrates teeth are fixed directly to the crest of jaw-bone, called acrodont. Neither
pleurodont, nor acrodont type of teeth have any root. The most advanced type of tooth is
tliecodont where each tooth has root (one/two) that is embeded within a socket of the jaw
bone. In this case, a tooth exhibits two parts : a crown above the jaw and a root below the
jaw. Thccodont teeth are shown by most of the reptiles and mammals.
Structurally, when all teeth in an animal are similar looking, the pattern is called /wmodo11t
but when an animal bears different types of teeth in its jaw, each type performing different
function, the pattern is called heterodo11t. (Fig. 21-11 b). Most fishes, amphibias and re ptiles
exhibit lwmodont teeth whereas most of the mammals have heterodont teeth (except whales).
Snake has normal small conical homodont teeth but poisonous snakes have a pair of long and
sharp teeth, their poison fangs, for injecting venom into the body of their prey.
Palac(Geo)WP-35
.. . •y--- ...
PALAEONTOLOGY
272
Axial bone
Crossopterygian
fish Humerus
Actinooterygian fish
(b) ICHTHYOPTERYGIUM TYPE
(a) ARCHIPTERYGIUM TYPE
Advanced Tetrapod
(c) CHEIROPTERYGIUM TYPE
FIG. 21 - l O : EVOLUT10N OF LIMB BONES FROM FISH-FIN
Some primitiv~ extinct amphibias and extinct crossopterygian fishes had developed a special
type called laby~mtliodont teeth which were conical teeth but made up of several longitudinal
ridges, each radially shows complicated fold patterns ·resembling a labyrinth (Fig. 21-1 lc).
Heterodont teeth of mammals (Fig. 21-J lb) functionally are of four types. Each type is
located at some definite part of the jaw. For ideal case, on each side of a jaw, there are incisor,
canin.e, premolar and molar teeth from front to backward direction. Usually, the incisor is
chisel-shaped having a single root, used for cutting food materials. The canine has sharp points
and edges used for tearing food and also with one root. Premolar and molar, also called the
cheek teeth, fundamentally are similar and are made up of some cusps placed on their flat
upper surface of the crown. They usually have two/three roots. Cheek teeth are used for crushing
and grinding of food materials. All mammals not necessarily possess all these four types teeth.
For example most of the herbivoras lack canine teeth. Sometimes some teeth may be modified
for performing other functions. For example the tusks of an elephant is actually its second
pair of incisors which is evergrowing. Molars of carnivors also p0· ,~ss cusps with sharp edges
unlike the other mammals.
The number of teeth is variable in heterodont mammals and is usually fixed and dignostic
for each group. Mammalian heterodonty is thus expressed by a dental formula which is an
expression of number of each type · of teeth in each half of the jaw, the lower jaw having
underlying the upper and the number of each type tooth is shown numerically. Thus the dental
formula of man is 2123 = 32 meaning that it has on each half of jaw, two incisors, one canine~
2123
two premolars and three molars. On each side of lower and upper jaw number of teeth is 16
and as mammals are bilaterally symmetrical, the other half of the jaw also contains I 6 teeth.
It is assumed from fossil evidences that the ancestral mammal had a total of 44 teeth and its
3143
dental formula was = 44.
3143
Dental formula of s~me other living mammals are as follows :
Rodent Rabbit f~jj = 28
Cat 0033 = 32
Carnivora 3133
Horse 3134 = 44
Perissodacty le 3134
The origin and evolution of compl~x ~remolar and molar tooth (Fig. 21-12) in ~am~als is
a matter of speculation. The theory which 1s generall~ accepted and supporte~ ~y fossil evidences
states that in the primitive conical tooth of a reptile were added two additional cone-shaped
buds giving rise to a triconodont shape of tooth. Fossils of some extinct reptiles of synapsid
group (mammal-like reptiles) and some primitive Mesozoic mammals exhibited this type tooth
even in Upper Triassic period. Gradually, these projecting cones shifted apart to give rise to
three tubercular cones (one original and the two added) or cusps in a tooth arranged in a
triangular fashion, called trituberculine type of tooth. This type of tooth is found among the
fossils of early mammals in Late Mesozoic. The original cone of the tooth is called protocone
PALAEONTOLocy
-~ f
4 - , M - - - t - - - Root
Al(,41,l.---::i~- Alveolus (Socket)
n."<d, 11 (Fish & \111 phibill) ThccoJont (Reptile & Mammal)

Plcurodont (Amphibia & Rcplilcl
(a) ATI"ACHMENT OF TEETH TO JAW
Canine
Homod Ill teeth of a n:p1ih.: (l izard)
Heterodont teeth of a Synapsid

1 1 erenci:ucd check teeth (mammal•like reptile)
Canine
Premolars ( 4 ) - - - - - -
Incisors (3)
Hc1croJ0111 1cc:th of a primitive

Mc~z.oic mammal (Triconodont) Hctcrodont teeth of a mammal with full
set of 44 teeth (HyracothC'r111m)
b) MORPHOLOGICAL VARIATION OF TEETH IN TETRAPODS
Enamel Folding Central Pulp
Dentine
T. S Of Tooth
Entire Tooth A Portion of T. S (Enlaged)
c) LAOYRINTHODONT TOOTH
FIG . 21 • II MORPHOLOGY OF TEETH OF VERTEBRATES
OUTLINE OF MORPHOLOGY OF SOM/:: SKELETAL ELEMENTS OF VERTEBRATES 275
Cama.~sial or secodont
(Cumivora) Denticulate
(Crab-eater seal)
(a) PATTERN OF CROWN STRUCTURE OF
Triconodont
CHEEK TOOTH Tritubcrculalc {fossil mammal)
(ta.~sil mammal)
(b) PRIMITIVE CHEEK TOOTH
Crown
,-------,-~r1r--- Dcntary
Gum
11•.---- Jaw bone

NP----cement
---Pulp cavity
(with nerves & blood vessels)
(c) L.S OF A HUMAN MOLAR TOOTH
Side view Side view

Side view Top view
Side view
p.c~~r---Enamel
ridges
:zt::~J.---Transvcrsc
,,.,Ni3.~ ridges or
lophs
Brachydont & 1-lypsodont &

Brachydont & Lophodont (elephant)
Sclenodont (tapir) Sclenodont (horse)
Bunodont (man)
(d) TYPES OF CHECK TEETH OF SOME RECENT MAMMALS
FIG. 21 - 12: MORPHOLOGY OF CHEEK TOOTH
!j
°'
Anterior covered pan
w ~- II~ Neucleus - \\\\({([~

(i) Posterior exposed part
Grooves ~
Cycloid Teeth (spine)
Ctenoid
--------11!!---Epidennis
r--Cosminc layer
':ffJ..t ·l':1 ,,),~~-
._\ ' 1. I~ , • -
Pu IP~·.·, .:.:. ·: .•.:•'~Middle layer
. - .. ~·1\ ~-
~
cavny -~ ,·:. , .., _ .:
-- .... ::!& .... - .
- · .ar-Dcntanc
(ii) Cosmoid
(i. Knie ii. section)
Ganoid
?
s:
Placold ~
~
t"-'
C
C)
FIG. '2\-\'3 : DIFFERENT TYPES OF DERMAL SCALES IN FISH ""<
and the other two are metacone and paracone. From these cusps later developed extra cusps,
ridges, folds to give rise to varied types of cheek teeth found in advanced mammals.
Considering the relative length of root and crown, molar tooth may be of two types (Fig.
21- I2d). Tooth with a larger root and a shorter crown is called brachydont as found in primates.
On the other hand, tooth with a larger crown and a smaller root is called liypsodont as found
in horse and many other grazing mammals.
On the basis of nature of the upper surface of crown, a molar tooth also shows a lot of
variations depending upon the type of food and feeding habits of mammals (Fig. 21-12d).
Bunodont tooth has small separate rounded cusps for grinding food as found in man. Secodont
tooth has pointed and sharp edged crown for tearing food as found in carnivoras (often called
carnassial tooth). In selenodont, the tooth becomes squarish with cresentic cusps of hard
enamel enclosing softer areas of dentine often filed up by cement. This tooth is used for
grinding and grazing of grass-like harsh food, and is characteristic of horses and catties. Many
mammals develop various intermediate forms of tooth like bunoselenodont as found in giraffids.
A lophodont tooth has cusps on crown surface joined with each other forming some transverse
ridges often intricately folded and covered by hard enamel. In between ridges occurs soft dentine
elements. These teeth are used for grinding coarse plant materials. In elephant only one
lophodont molar becomes functional at a time in each half of the jaw. A combination form
bunolophodont is found in some extinct group of elephants.
Structurally, an ideal tooth (Fig. 2 l-12c) is · made of dentine, chemically similar to bone.
At the upper surface of the crown, the dentine is covered by a very hard and shining materials
called enamel. The neck and the root of the tooth are covered by rather dull-lustred substance
called cement. Chemically cement, dentine and enamel are similar but show only a variation
in their content of some inorganic salts.
21.12 ACCESSORY SKELETAL ELEMENTS

A. Fish scales (Fig. 21-13)
Fish scales are usually mesodermal in origin. Primitive fishes developed two forms of scales.
(a) Cosmoid : The earliest known vertebrates had an armour of large bony plates which
became smaller and was transformed into cosmoid scales of placoderms. 'nlese scales
are found today only in the living fossil Latimeria. A cosmoid scale has four distinct
layers : the lowest layer of isopec{ine or dentine (compact and bony), the mid~le two
layers composed of spongy bones with vesicular spaces for blood vessels, the t~trd and
the uppermost layer called cosmine may be again covered by an outermost thm layer
of shining enamel.
(b) Ganoid : Ganoid scales are found in primitive bony fishes. This scale h~ also a basal
layer of isopedine above which there may or may not be a reduced cosmine la~er. The
uppermost layer is made up of ganoin which has a thick cover of enamel. Ganotd scal~s
are found in fossils of many primitive bony fishes and at present, they are found m
fishes like Polypterus and Lepisosteus.
i ·,'
I I It has been implied that evolution of fish-scales proceeded along two dirnherent linegsra~:~
',;
~-:
~·,
"~
· l ·
the very early history of the fish. In the hne of evo utton
f th
? e ga h
noid scale t ere was
0
wever two.types
u loss of the upper ganoin layer giving rise to thinner leptoid scale. There are
~ '
.... -.- --
27K PALAEONTOLOGy
H tr) I i,,(k:nMI SI.in ..,.___ Horny

Epidennai
Covering
a.:TJ--- Epidermal
Layer
cntr.il
R11ny Core Prong Horn (Anti/ope) Bony Core
Kn<'b Horn (G,mffc) Fibre Horn (Rlrlnoceros)

Horny Epidermal Layer
Bony Core
Dony Horn SI.in
Mature Stage (Antler) Hollow Hom Bison (True Horn)

Immature Stage (Ante/er)
fa) DIFFERENT TYPES OF HORNS OF MAMMALS
IL.1 ·h i~ ,,r Shafr ------tiN

(Sc:,pu~I
Calamus-------~
Down Feather
Contour Fcntncr
(b) FEATHERS OF BIRDS
Nail Plate (Ungu,s)
Nall Bone
Claw Of Oird & Rcpcilcs
Nail Of Mon (Sagltllll Section)
Subun,ui5
Cuneuli
(c) DIGITAL TIPS OF VERTEBRATES Hoof of Uqulales
FJO. 21 -14 : DERMAL SKELETONS OF SOME VERTEBRATES
OUTLINE OF MORPHOLOGY OF SOME KEU.TAL EU£M FNT\' <W \ RlfffllNA'/'l~S .• i 'J
of teptoid scales namely cycloid and ctenoid both having II singk 111 l'r of isop •din,~. l 'y ·loid
scales are circular in outline, thicker at middl . having n low •r l11yor of lihrnus conm· ·1iw 1iss11rs
and an upper bony layer of isopedin". They also show cone •ntri · )otl'OWlh lin ·s ofl ·n 111urki n1,t
the age of a fish. These scales lie with posterior part of •a ·h ovurlupping th• nnt ·rior portion
of the scale behind and in this way the body is cov ·r ·d by u dnubl ·-scul • lay •r. Th • conccakd
part of scale may have a wavy margin. Ctenoid scal .. s are probnhly un udvunc ·d fnrni of 9duid
scales having the same shape and structure but they differ in huving small tc"lh or cte11i on
their free poste.rior part and their anterior concealed part muy hnv • notched or sculopcd mnrgin.
In another line the cosmoid scale has lost its three out · r luy rs, retaining the fourth ·num ·1-
like dentine layer and becomes somewhat enlarged to form placoid seal •s. Such sent "S ar · found
in the body of sharks where they are also provided with lutcrnl and mediun spin ·s.
B. Fins
Fins are appendicular skeletal features of fishes. Two types of fins ure found in u fish body.
paired fins (pectoral and pelvic) and unpaired median fins on dorsal and ventrnl sides. ln ideal
ray-finned fishes, the main body of all fins are structurally similar but the mode of auuchmcnt
of paired and unpaired fins is different.
A typical median fin is usually traingular in outline with u narrowing busnl part. The body
of a fin is composed of a series of fine rod-like cartilage bones often with spiny terminals
called pterygoniphores or somactids. Usually a wide strip of ligamentous tissue connects
somactids with notochord. Somactids diverge from base of a fin where they are occasionally
fused. Laterally a membranous layer connects somactids with each other. In shark, somactids
bear in front a double series of numerous fine horny rays called ceratotrichia.
The paired fins of both pectoral and pelvic region are attached with the axial skdeton by
means of cartilaginous girdle-bones. The pectoral girdle is formed by the fusion of two cartilage
bones on either side below the pericardium. Each bone is called scapulacoracoid whose
outerpart is scapula and the innerpart is coracoid. It bears a few foramens for the passage of
arteries and nerves. The main body on the fin with somactids becomes joined with corucoid
by three fused small cartilages namely protopterygium, mesopterygium and metapterygium.
Pelvic girdle on the other hand is a flat rod of cartilage, called ischopubic bar, also produced
by fusion of two cartilages (ischium and pubis). On each side of the bar is an acetabular region
to which is attached the basipterygium (fused basal part of somactids) of the fin (Fig. 2 I-Sa).
In all the cases, muscles attached to bones help the movement of fins.
C. Corneal structure-Horns (Fig. 2 I- l 4a)

Ho~s are generally found within ungulate group of mammals. Five types of horns may be
leCOgmsed.
(a) Keratin fibre horn : Horns of Rhinoceros are of this type. 1t has no skeletal element
,. but' is made up of keratinized cells of epidermis that consist of matted keratin fibres
bou~ tog~ther. ~t is a permanent epider~1al structure in head and if broken, it grows
' agam. lnd1an Rhinoceros has two but African has only one horn.
(b) Prong horn : lt c~nsists of a permanent projection of the frontal bones of head covered
by hard horny epidermal sheath which is occasionally forked forming one to three
prongs ~~c up o~ only horny sheath which is shed annuully. This type of horns are
found w1thm Russian antelopes A11tilocapra.
Palac<,Gco)WP-36
PAL.AEONTOLOG'I
280
(c) Antler . This type of horns are found within males of deer family ~t are 11ometi:C1
resent .in both sexes in a few groups. An antler cosists of a brandc mlg otuThtgrowt I of
P · · h b h ·ry skin often calle ve ve . e vc vet
frontal bone covered durmg its growt Y a ai · .• bo
1 The
is shed after the full growth of the antler when it is only a naked derma nc.
horn is shed after each breeding season.
(d) Hallow horn : This type of horns are found in catties, sheeps, goats and in ":1~ny other
· d I · ·n both sexes It has a prominent bony dermal core arising from
art10 acty s occurring 1 · · fi d ·d ·
frontal bones, covered by a permanent hollow horn derived from corm ,e epi enms.
They are unbranched and non-shedding.
(e) Giraffe horn (Knob horn) : They are like those of antlers, wit.h a bony dermal core
covered by an epidermal velvet or skin which is never shed. Giraffe horns are short,
unbranched, permanent and are found in both sexes.
D. Digital tips (Fig. 21-14c) .

In most of the tetrapods, the terminals of the digits are provided with such acces~ory s~eletal
structures like claws, nails and hoofs which are derived from horny layers of ep1derm1s and
are found growing parallel to the surface of the skin.
A common variety of digital tips found in reptiles, birds and also in some mammals are
claws. They are made up of an upper hard scale-like plate called unguis underlain by a
relatively soft layer called subunguis, both converge to a point and cover the terminal part of
the last digital bone (phalanx). Nails and hoofs are generally found within mammals. In some
mammalian nail, the subunguis layer becomes reduced and continuous with a pad at the end
of the digit. These types of claws are found within carnivoras. Nails are found in primates
where the upper unguis layer is large, flat and thin while the subunguis is much reduced in
frontal part of the nail. The back portion of unguis, then, lies on a nail-matrix and the whole
structure is within a nail groove. Hoofs are characteristic features of ungulate mammals. In
hoof the unguis becomes very thick and it covers all around the end of the digit completely
enclosing the subunguis as a core of unguis layer. Behind the hoof is a soft pad. Both nails
and hoofs are modified forms of claws.
E. Feathers (Fig. 21-14b)

Feathers, found only in birds, are considered some modified forms of reptilean scales. lbey
are light, strong, elastic and waterproof showing many colours. They develop from the epidennis.
They form a protective cover and they regulate body temperature and support in flight. 1nere
are three ~ain types of feathers; contour feature, down feather, and filoplume of which the
first type 1s the most common and found all over the body of bird. It has three parts :· a stalk
at th~ base. (calamus), a rachis or stem called scapus and the membranous hair like nions
on either side of the axis called vane. po
TABLE-14 ~
MAJOR SKELETAL ELEMENTS OF SKULL OF DIFFERENT VERTEBRATES c:::.
~
+ Unpaired bone; ./ Present; • Absent. ~
0
Region of skull "'ti
Cartilage bones Dermal bones Mixed bones Fish Amphibia Reptilia Bird Mammal
~
OCCIPITAL Supraoccipital+ ./ unossified ./ ./ ./

~
i
C
Exoccipital ./ unossified ./ ./ ./ t""'
C
Basioccipital+ ./ • ./ ./ ./ C')
~
PARIETAL Basisphenoid+ ./ ./ ./ ./ ./ .,,C

V)
Pleurosphenoid ./ • ./ ./ • ~
C
Parietal ./ ./ ./ ./ ./ V)
Interparietal+ • • • ./ • ~
l"?']
Postparietal ./ • • • ./ ~
t""'
FRONTAL Orbitosphenoid ./ ./ ./ ./ ./ f:2
Presphenoid+ ./ ./ ./ ./ ./ ~
~
Frontal ./ ./ ./ ./ ./ ~
Postfrontal ./ • • • • .,,0
Lacrimal ./ ./ • ./ ./ ~
~
• • • ~
Pre frontal ./ ./ 0::,
./ ./ ./ ./ ./ ~
Mesethmoid+
OLFACTORY ~
V)
CAPSULE ./ ./ ~ ~ ~
Turbinals
Cribriform • • • • ~
Ectethmoid
~
• • • • N
00
Region of skull Cartilage bones Dermal bones Mixed bones
Fish Amphibia Reptilia Bird Mammal
Nasal ./ ./ ./ ./ ./
Vomer
• • • • ./
OTIC CAPSULE Epiotic

Septomax iBary
• ./
• • •
./ ./ ./ ./ ./
Prootic ./ ./ ./ ./ ./
Opisthotic ./ ./ ./ ./ ./
Squamosal ./ ./ ./ ./ ./
Supratemporal ./ ./ ./ ./ ./
Sphenotic ./ • • • •
Pterotic ./ • • • •
OPTIC CAPSULE Sclerotic • • ./ ./ •
P~ATE Parasphenoid+ ./ ./ ./ ./ •
Pterygoid • .
./ ./ ./ ./
Ectopterygoid ./ • • • •
Endopterygoid ./ • • • •
Palatine ./ ./ ./ ./ •
UPPER JAW Quadrate ./ ./ ./ ./ •
./ ?
Epipterygoid • ./ ./ ./
s:
Metapterygoid
./ • • • • ~
• • • • ./
~
Alisphenoid
,. s
C)
-..:
f ,;
;
~
'
~
, . C
Region or skull Cartilage bones Dermal bones Mixed bones Fish Amphibia Reptilia Bird Mammal c:::
Premaxilla ./ ./ ./ ./ ./
-
~
~
C
Maxilla ./ ./ ./ ./ ./ "!i
~
Jugal ./ ./ ./ C
./ ./
.,,
;:i:,
Quadratojugal ./ ./ ./ ./ • ::i::
C
t""'
LOWER JAW Articular ./ ./ ./ ./ • C
C')
Malleus
• • • • ./
""<:
C
"!i
Mentomekelian
• ./ • • • V')
C
Mandible • • • • ./ ~
t't]
Coronoid • • ./ ./ • V')
Splenial • • ./ ./ • ~
~
Angular • • ./ ./ • t""'
~
Suprangular • • ./ ./ • t't]
~
HYOID ARCH Hyomadibular ./ • • • • ~
Symplectic and ~
associated bones
./
• • • • a
"!i
• ~
Columella ./ ./ ./
• ::ti
~
Stapes • • • ./ • ti:,
~
GILL COVER Preopercular ./ • • • • ~
Opercular ./ • • • •
V')
Subopercular ./ • • • •
Interopercular ./ • • • • N
Gular ./
• • • • 00
'Jol
Chapter 22
SOME ASPECTS OF EVOLUTION OF VERTEBRATES
22.1 INVERTEBRATE TO VERTEBRATES
So far no undoubted representative of chordates has been found among the fossils of
Cambrain age and one general fact can be derived from the study of Cambrian fossils is that
they were all invertebrates lived in ocean. The first report of vertebrate fossils came from
Ordovician rocks of Colorado which are fragmentary remains of bones and scales of some fish-
1ike animals. Then vertebrates certainly arose from some sort of invertebrate animals of
Cambrian or still older age. But as is so often the cases, the exact ancestral group yet remains
unknown. One common question about the vertebrate ancestry is wheather ancestral vertebrates
were free swimmers or sessile forms. Tunicates, one of the simplest members of living chordates
belonging to protochordata exhibit free swimming tad-pole like larvae at the initial stage, each
bearing a notochord at their tail, a dorsal nerve and pharyngeal gill slits. At adult stage the
animals become sessile and filter-feeders (with loss of notochord and dorsal nerve) resembling
calms or oysters. Amphioxus, another member of the same group is however, fish-like swimmer,
filter-feeder and it retains the notochord throughout the life time. From this observation, many
workers like to suggest that the ancestral chordates were derived from some filter-feeder
invertebrates and that initially they were swimmers like their ancestor. But which group of
invertebrates could be the ancestor, or at least most closely related to vertebrates? Most of the
palaeontologists and neontologists have inclined to accept echinoderms as the desired group.
Echinoderms are closely related as regards their nature of larvae and larval development to
chaetognath worms in one hand and some protochordates on the other. They resemble chordates
in the manner of forming their body cavities and mesoderm. The idea is that there was a
hypothetical worm-like animal (derived from chaetognath worm), often named 'dipleurula'. This
was a bilaterally symmetrical, flat cylindrical animal with ventral mouth and anus, symmetrical
coelomic pouches on either side. They had also a ciliated band forming a loop running down
on either side with coordinating nerve plexus.
It is believed that chordata-echinoderm group on one hand and lophophorate group
(branchiopod-bryozoa) on the other, were independently derived from such an ancestor. The idea
is that the ciliated band of the ancestor dipleurula was fused along its midline to provide the
rudiment of dorsal nerve cord of the chordate in one direction, while torsion of alimentary canal
with an upwardly directed mouth and a posterior lateral anus led it to the direction of echinoderm
group. Today one may find a similar simple la!val stage both in echinoderms and potochordates.
A more plausible explanation is given by Berri II ( 1955) (Fig. 22.1 ). It is assumed that the
common echinoderm-chordate ancestor was possibly a small sessile arm-feeoing (lophophorate)
creature fed by waving ciliary tentacles. From this, was derived the primitive sessile echinoderm
on one direction and a sessile filter feeder stemchordate (pterobranch) on the other direction.
In the latter the external tentacles were replaced by an internal filtering apparatus in which
284
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, ,.,
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-
•.
· ·· · - - · / ..
.. -- c ·" £ ASP£C1 OFF.\ OLlfrlON Of VERTEBRA'/'/::S 285
! '
~t he nu chordate
( Acorn worm)
Primitive pterobranch
(Scssik arm feeder)
Primitive sessile
echinoderm
Primitive sessile arm feeder

Ancestral slu..:k
FlG . 22 - I : DIAGRAMATIC REPRESENTATION OF ORIGIN OF CHORDATES BASED ON
IDEA OF BERRILL (1955)
l'ulutoqundratc Hyomndihula
Maxilla
FIG . 22 -2 : EVOLUTION OF JAW FROM BRANCHIAL ARCH IN PLACODGRM FISH
286 PALAEONTOLOGY
food was entrapped inside the pharynx that gradually developed as external gill silts. From this
fonn evolved free-living hemichordates on one hand and sessile ancestral urochordates (tunicates)
on the other direction. All these forms however, develop initial free-living ciliated larvae.
However, some ancestral tunicates instead of producing such normal ciliated larvae (common
to all those previous groups) formed tad-pol~ larvae with aJI typical somatic features of
chordates. This typical protochordate larva by paedogenesis, suppressed the sessile adult stage
and developed reproductive organs at a premature condition and ultimately became the ancestor
of both cephalochordates and jawless vertebrates, the agnathas.
22.2 EVOLUTION OF JAWS (Fig. 22-2)

The second great event in the history of the vertebrates is the appearance of jaws as it opened
the vertebrates to new lines of adaptation and diversification and new lines of evolutionary
advancement. Jaws of vertebrates are believed to have originated by a morphological
transformation of gill arches, the bones supporting gills within jawless vertebrates. The primitive
vertebrates agnathas such as ostracoderms bore several pairs of gills and gill-arches. Each giJl-
arch was formed by a series of bones arranged somewhat in the fashion of the english word
'V' with the points directed posteriorly. In course of evolution, a few frontal pairs of gill-arches
were eliminated but one pair of gill arches was transformed into jaws of the higher vertebrates.
A study of embryological development of some early placoderms and modem fishes indicates
this type of origin for the jaws. This type of transformation of one structure performing a
particular function to another structure showing a different function had taken place several
times in the course of animal evolution and much of the progress of advanced animals was
brought about by such transformation of an existing structure to another.
22.3 FROM WATER TO LAND

The most significant event in the evolution of vertebrates is the appearance of terrestrial
tetrapoda from fish. Advent of fishes with jaws took place in Upper Silurian but their success
came in Devonian when they became the only dominant group of vertebrates in water. They
include cartilaginous and bony fishes. The teleostei, the most successful group of bony fishes,
however, is off the main line of evolution that led to the advance groups of tetrapods. In order
to search for the origin of tetrapods, we have to turn to the line of evolution represented by
the crossopterygians, a group of air-breathing fishes. They appeared in Devonian like many
other fishes. The early members of these fishes were fusiform having heterocercal tails, paired
lobed-fins like other lung-fishes, but in addition, they had completely bony skulls and jaws
comparable to those seen in early land living vertebrates. They also possessed two larger bones
between the eyes (homologous to parietal bones of tetrapods) and the latter were surrounded
by some circumorbital bones like higher vertebrates. In other words, the early crossopterygian
fishes had many such features which one should expect within the ancestor of land vertebrates.
That is why they are regarded as the direct progenitors of the first amphibia labyrinthodonts.
The labyrinthodonts also retained many features of their crossopterygian ancestor. Their name
refers to a complicated pattern of infolding of enamel of the teeth. A sim;;dr feature also has
characterised the enamel pattern of crossopterygian teeth. The mosaic pattern of series of bones
fitted edge to edge in the skull of labyrinthodonts corresponds in details to the bony plates of
head of crossopterygians. All these similarities leave a little doubt that crossopterygians were
the ancestors of the first land vertebrates, the amphibias (Fig. 22-3). The similarities between
SOME ASPECTS OF EVOLUTION OF VERTEBRATES 287
the Devonian crossopterygian Osteolepis and Eusthenopteron with the earliest labyrinthodont
arnphibia lchthyostega are very much conspicuous. .
From fossil remains it can be inferred that crossopterygian-labyrinthodont transition took
place in Upper Devonian times. From the pattern of skeletons within lobed fins of Devonian
rossopterygians it can be inferred that they used their fins to crawl ou·t on land as and when
nc ·essary. Early amphibias, although possessed many preadaptations for the life in land, it is
probable that they spent greater part of their life in water like their ancestors and like many
amphibias of the present day. But they were also capable of walking on land by their bony
paired limbs which are modified forms of lobed-fins of their ancestors. Possibly they left water
when they were forced to do so. But when did they do so? Obviously, they did not leave water
10 escape the predators since in the aquatic environment of that time they themselves were one
of the largest predators. Since they were carnivorous and prospective preys in the form of
animals were abundant in water, paucity of food can hardly be an explanation. The most possible
explanation is based on the idea that the ancestral amphibias lived in pools that dried up
periodically as do the pools in which some lung-fishes live today. Gradual evaporation of
stagnant water during periodic dry season leading to overcrowding, forced these animals to leave
their old house, possibly in search for neighbouring pools with better living conditions. Such
an adverse and stringent conditions ultimately led them to become true land dwellers.
22.4 RISE OF REPTILES

A leap from water to land towards the close of Devonian leading one vertebrate class to
another (from fish to amphibia) may be considered as one of the most significant event in the
history of vertebrate evolution. From this they entered a completely new environmental domain,
facing new challenges leading to several new lines of evolutionary developments over a
tremendous range of adaptations. Let us consider the new challenges faced by these early land
vertebrates and how they were able to cope with them. This can justify the cause of evolution
of the next group of vertebrates, the reptiles.
One of the problem with which the early amphibias had to contend was the mechanism of
respiration. Co_mmonly, the fish obtains oxygen from water and performs respiration by gills.
Land vertebrates have to draw oxygen from air and perform respiration by means of their lungs.
Solving this problem by early amphibias was not very difficult as they had merely to go on
using the lungs that they had inherited from their crossopterygian ancestors (lung fishes). But
unlike the ancestors that used their lungs as accessory breathing organs, amphibias are primarily
air breathing animals and they use their gills only at their larval stage of life.
The second problem confronted by the first land vertebrates was related to their naked skin
which failed to prevent the excess evaporation of water from their body in open land. Probably,
some earlier amphibias (ichthyostegids) lived more in water and never went far away from the
water as do by many modern amphibias. Some other amphibias retained the scales or similar
structures like those of their ancestors. Some Permian amphibias are found to develop a tough
skin often underlain by bony plates. Some modem amphibias are found growing much tough
and glandular skins for keeping their outside wetty. So they have adapted several means which
enable them to be increasingly independent of water but a complete·-and appreciable success
was never achieved.
Palae(Geo)WP-37
288 PALAEONTOLOGY
Reconstructed Archaeopteryx
(Barlic~r Bird)
Pm
T Pn
h
Thcrapsid Reptile (Synnpsid)
<Ancestor To Mammal)
Thecodont (DiapsidJ Rcprik
(Ancestor 10 Bird)
Cot)'losaur (Anapsid)
(Stem Reptile)
M
f-""::l,,-r--.,...~--L_
r
~~::::::::::::aG~?- up
Labyrinthodonl
,\n Sa
(Stem Amphibia)
Pf" p
Crossopterygian fish <>s1c:olc:pis-Crossop1crygian

(Accstor to amphibia) ·A (Lobed Fin Fish)
FlG. 22-3 : CHANGE or: SKULL PATTERN :\NO EVOLUTION Or MAJOR GROlTPS OF
VERTEBRATES
[A : Articular; An : Angular; D : Dentary; F: Frontal; G : Gulnr; It : lntertemporal: J : Jugal; L : Lacrymal:
M : Maxilla; N : Nasal; Op : Opercular; P : Parietal; Pd : Predentary; Pf : Post Frontal; Pm : Premaxilla:
Po : Post Orbital; Pop : Preopercular; Pos : Post Splenial; Pp : Post Parietal: Prf: Prefrontal; Q : Quadratc:
Qj : Quadrato Jugal; Sa/San : Subangular; Sop : Subopercular; Sq : Squamjosal; SI : Supratemporal :
T : Temporal. Th : Teeth; Tf : Temporal fossn: Tb : Tympnnic Bulla]
~~a~N~----Amnotic Cavity
~---Embryo
~"l+----Chorionic Cavity
FIG. 22 - 4 : TRANSVERSE SECTION OF AN AMNIOTE EGG OF REPTILE
_,,,_-
, .
soME ASPECTS OF EVOLUTION OF VERTEBRATES 289

A land animal have to cope with the power of gravity which for aquatic animals is less
problematic for they are living within denser water that lessens the effect of gravity. Early
land vertebrates had not only to devise some structures in counter action of gravity but also
had to walk on land freely against this force. Thus they developed stronger backbone and limbs
at their very early stage of evolution. The simple disc-shaped vertebrae of the fish are
transformed into interlocking structure in which these are articulated with one another by
muscles and ligaments forming a flexible but strong vertebral column that gives necessary
support to the body. Limbs bones are strongly supported and articulated with vertebral column
by means of two pairs of bony girdles; pectoral girdle in front and pelvic girdle in the back.
Such a locomotive device is just reverse from that of a fish whi~h swims by its body while its
fins and tail keep the balance. A land vertebrates, on the other hand, moves by limbs
(homologous to fish fins) while its tail functions as a balancing organ. Such a ba~ic skeletal
plan and pattern of locomotion initiated by the first amphibias are retained by all other
subsequently evolved terrestrial vertebrates.
However, the most crucial problem was related to the mechanism of reproduction. Fishes
hatch unprotected eggs in water where they develop. A land animal may have two options,
either going back to water to reproduce or developing a method of protecting eggs in open
land. Amphibias, though made several advances in the adaptations of their life in land but they
are unable to solve this problem and consequently they have to return to water at the time of
their reproduction for laying eggs from which develop their aquatic larvae called tadpoles
exhibiting many characteristic features of fishes. The external gill-slits of an amphibian larvae
resemble those of modern lungfishes. Within this failure of solving the problem of reproduction
in land, lies the seed of evolution of yet another group of new land vertebrate, the reptiles which
were undoubtedly derived from the amphibian ancestory.
The transition from amphibia to the descendant reptile is far Jess convincing from the fossil
record unlike the crossopterygian-amphibian transition. This is because the principal difference
between an amphibia and a reptile is not found in their skeletal forms but lies in their mode
of reproduction. The first reptile finally was able to overcome the problem of reproduction in
land by developing shelled amniote eggs, a device for allowing the embryo to grow in its
own self-contained liquid environment within egg shell. Within a pliable egg-shell lies a bladder-
like membrane (amnion) that contains the amniotic fluid in which embryo floats (Fig. 22-4).
The fluid protects the embryo from mechanical injury and drying. Thus a reptilian egg is just
the egg of a fish which is bottled up not within wate~ but in amnion. The water and food for
such an embryo are stored within yolk and albumen and oxygen is permitted to go through the
porous egg-shell and the latter also gives an external protection of embryo. The animals with
such an egg can wander freely on the land without having to return to water for reproduction.
The first animal to have such eggs is the reptile which makes him a true land vertebrate.
The other advances achieved by reptiles over their amphibian ancestor were their greater
ability to regulate the body temperature and body water. Most of the reptiles have some body-
coverings in the form of horny scales, armours and such other features which not only protect
the animal from external injury but also reduce the loss of body water through evaporation.
Now it becomes quite clear that distinction between the first reptile and its immediate
.i amphibian ancestor from the fossil record is not so easy because there is hardly any structural
difference between them. However, it appears that reptile's immediate ancestor was one type of
290 PALAEONTOLOGY
labyrinthodont amphibius EUld the transition probably took place in later part of Carboniferous,
although th " first fossil of amniotic egg has been obtained from the Lower Permian sediment in
North Ame ri ·n, long after the presumed time of first appearance of reptiles. But of course, we
have some fossils of labyrinthodonts showing somewhat intermediate characters between an
amphibia and a reptile. The most significant feature common to a labyrinthodont and a reptile is
their possess.ion of one occipital condyle, (the bony knob by which the skull is articulated to the
first vertebra of the backbone), while the modern amphibias have two occipital condyles. From
fossil record it appears thut cotylosaur reptiles were the most primitive and they appeared from
labyrinthodonts in L 1te Carboniferous to which they are similar in many aspects (Fig. 22-3). Many
palaeontologists considered Lower Permian Seymouria as a member of earliest reptile cotylosaur.
It was a lizard-like anirnal about 60 cm long possessing both amphibian and reptilian affinities.
It resembles labyrinthodonts in having a less ossified skull without temporal opening,
labyrinthodont-t.ype of teeth, differentiated vertebrae and a pectoral girdle more closer to skull.
Its reptilian affinities include, possession of more perfect muscular limbs that arise mid-ventrally,
anapsid t) pe of skull with complete roofing and a pelvic girdle, strongly articulated with the
vertebral column.
22.S FROM LAND TO AIR

Biologists have long, fascinated by a close anatomical resemblance between a bird and a
reptile. The tenn 'glorified reptiles' frequently applied to birds, suggests the fact they are similar
to reptiles in many ways. There is almost no doubt about the suggestion that birds arose from
some reptile ancestry most probably in the Mesozoic. But before bird possibly another group
of tetrapods, the flying reptiles (pterosaurs) had already invente~ the air.
The first requirement for a flying animal is to overcome the downward pull of gravity. This
is especially a problem for large forms like birds and flying reptiles than the insects. The flight,
so far we know, requires a high rate of body metabolism and the animal has to control its body
temperature by developing some devices, and also quite obviously it must have a flying device
in the form of wings.
For adaptation to this new domain both birds, and flying reptiles have developed light hollow
bones with thin outer wall, and a pair of wings which are modified forms of forelimbs. Along
with this, for efficient flight, they have strong backbone and powerful arm muscles to move
the wings up and down. They also have additional areas for attachment of muscle. Their sternum
or breast-bone becomes highly enlarged to provide anchorage for such muscles. Birds also have
'eye brains' that is brains showing pronounced visual areas with reduction of portion connected
with sense of smell. Besides, birds have some air-sacs connected to the lungs serving principally
as a ventilating system, dissipating the heat generated by their vigorous metabolic activity
necessary to fly. Being warm-blooded birds can control their body temperature. Feathers are
considered as some modified forms of scales of reptiles. The pterosaur has sternum bone and
its brain was comparatively larger than other reptiles but wheather being a cold-blooded animal
it was able to control the heat of the body or were able to generate heat required for flight is
a matter of speculation.
Then the question arises, 'which type of reptiles could be the ancestor of bird and wheather
pterosaurs and birds were the descendants of the same ancestor?' Here, we can recall the
.r
J
;,,\ SOME ASPECTS OF EVOLUTION OF VERTEBRATES 291
l
primitive reptile cotylosaur, as it has been more or less accepted as a stem reptile and could
;t be the ancestor of all other reptiles and also could be the distant ancestor of birds and mammals.
In fact, several lines diverged out in Late Permian-Triassic from this stem-reptile of which the
two important groups were the thecodonts and synapsids. The former gave rise to pterosaurs,
birds and other reptiles including dinosaurs while from the latter ascended the mammals (Fig.
22-3). Some of the unique achievements of thecodonts were their way of locomotion. Instead
.)
of walking or running on four limbs they adopted a bipedal locomotion running on two
hindlimbs as done by many dinosaurs, pterosaurs and birds. For this, their hind limbs became
elongated, forming a support upon which the body was balanced as on a fulcrum. The body
was projected forward from this fulcrum and its weight was counter balanced by a Jong tail.
Forelimbs, being freed, were used for grasping preys or other food materials. This thecodont
body plan, as Colbert (1951) stated is "the blueprint to dinosaurian forms". Birds and pterosaurs
are creatures showing parallel evolution after evolving form a common thecodont ancestor. The
first bird like the first pterosaur appeared in Jurassic times and the skeleton of the earliest bird
is recovered from Solenhofen Limestone of Bavaria, Germany, which is named Archaeopteryx
lithographica, sometimes called a 'missing link' between reptile and bird, possessing some
intermediate or common structures of both. Their reptilian characters include (a) presence of
scales (b) beaks provided with pe.g-like teeth (c) a tapering lizard-like tail (d) sternum without
a keel (possibly this bird was flightless) (e) presence of carpals and metacarpals (unlike bird
where they form carpo-metacarpus). The Avian characters of Archaeopteryx are (a) body covered
by feathers (b) larger brain and larger orbit (c) presence of beaks (d) tibia-fibula separated
(e) eyes with sclerotic ossicles (t) presence of clavicle and wishbone (Fig. 22-3). ·
What could be the nature of immediate ancestor of birds? It is difficult to accept some
ornithischian dinosaurs or pterosaurs as th~ immediate ancestor of bird though the former have
a similar pelvic structure, a rapid bipedal movement and a beak-like structure without teeth.
The pterosaur, like bird was a light-boned creature. But none of them have clavicle or wishbone
and sclerotic ossicle, characteristic of all birds and even present in Archaeopteryx. This must
indicates that birds evolved independently from some thecodonts showing some generalized
characters like possession of clavicle and sclerotic ring. Even there might be a common
thecodont ancestor of bird and dinosaur in Late Triassic.
It was a common belief that running birds (ratitae) appeared first from which arose flying
birds (carinatae) at a later period. But recently a fossil, Eleutherornis, is found (thought to be
the probable ancestor of ostrich) from Eocene of Switzerland that shows a closer affinity to
flying birds than does the present day ostrich. In that case, as believed by some palaeontologists,
carinatae may be more primitive and may be the ancestor to running birds which were readapted
to terrestrial mode of life to avoid competition.
As regards the origin of flight there are also several hypotheses. According to some, flight
resulted from rapid running on the ground, occasionally by leaping on their larger hind limbs.
Gradually forelimbs became enlarged and modified into wings. Another theory postulates that
ancestral birds were adapted to arboreal life and they occasionally climbed down from tree by
I ·' gliding to the ground as done by some modem flying mammals (like flying squirrels). Possibly.
i ·1 they possessed feathers in both the limbs. Eventually the forelimbs became enlarged and
'· .
I ' converted to wings. Some consider that birds might have originated from both arboreal and
:~ cursorial ancestors.
292 PALAEONTOUJG't
22.6 RISE OF MAMMALS

There is no fundnm •ntal diffor~ncc in skclt:lnl or strncturul puttcrn between a reptile and it
mammal. The differenc ~ is lying in thoir system uf reproduction and in their pattern of
reproducti,e organs. However, lik, hird, mnmmuls urc warm-blooded an~ huve an insulating
cover on their body in the form of hairs by which they cun regulate their body temperature.
Moreover, they have devclo1 •d more cftici ~nt heart, lungs and heterodont teeth (suitable for
all types of food . Shdled reptilian egg was 1111 imporlunt advunce over the amphibia and fish.
Mammals, on the other hand, cxhihit a mnjor advance by total abandon of reproducing the
e.xtemal egg. Instead, the egg is retained within the body of female where it is fertilized. The
devdopment f the embryo takes place within the body of female before being borne active.
Along with this, the animal has developed another specialization that permits the young to be
nourished by the mother before its full development; that is, the development of milk producing
glands in the female's body from which the class takes its name.
Fossils provide ample evidences from which it is inferred that synapsid reptiles were the
ancestors of mammals. This group of reptiles possessed several features common to both reptiles
and mammals which are as follows :
Reptilian features : (a) cranium relatively smaller than avernge mammals (b) parietal foramen
(c) a single middle ear-bone (d) reduced quadrate, quadratojugal and other lower jaw bones
apan from dentary.
Mammalian features : (a) typical upright mammalian limbs capable of running with
considerable speed (b) skull with two occipital condyles (like all mammals), a secondary palate
and an enlarged lateral temporal fossa (c) lower jaw with the largest bone dentary (d) dentition
heterodont consisting of incisors, canine and undifferentiated cheek teeth.
It is difficult to precise the line of mammalian ancestor among the synapsids but most of
the workers have taken the therapsid as the immediate ancestor. They showed far advance
towards mammals in possession of some features along with some primitive character. But the
mixture of these two types of characters are so variable that it is hardly possible to point out
a particular group of these reptiles as the ancestor of mammals. It is quite possible that
mammals might have a polyphyletic origin, that is several groups of mammal-like reptiles led
to the ancestry of different groups of early mammals. Most probably this tmnsition took place
at Late Triassic or Early Jurassic times. The earliest mammalian fossils are represented by
morganucodonts reported from Upper Triassic of Europe and Asia. They were tiny animals
having slender single-boned lower jaw with teeth differentiated into incisors, canine and post-
canine molars and premolars. •
Chapter 23
GEOLOGICAL IDSTORY OF VERTEBRATES
23.l JAWLESS VERTEBRATES
The earliest vertebrates are included within the group agnatha (Gk.-a : without; gnathus :
jaws) which possibly appeared in Cambrian. Conodonts, small tooth-like microfossils of
Cambrian-Ordovician are now interpreted as denticles of some primitive jaw-less fish-like
animals often named as Cladagnathus. Complete conodont animals have been reported from
Edinburg (Briggs et al., 1983). They were eel-like animals of nearly 4cm. length exhibiting
head, eyes, notochord, v-shaped tissue-blocks (myotomes) and a tail-fin. However, there are
still contradictory opinions about these conodont animals. Besides conodonts, there is also report
of at least two forms of Ordovician agnathas. Arandraspis has been obtained from Middle
Ordovician of Australia having a head shield made up of a dorsal and a ventral plate joined
laterally by branchial plates covering the gill-areas (Ritchie and Gilbert-Thomlinson, 1977).
Asrraspis, known from Ordovician rocks of North America had an extensive head-shield with
8-13 gill-slits (Elliott, 1989).
Ostracoderms (Fig. 23-1 a) were the most primitive extinct agnathas characterised by heavy
bony dermal plates, the frontal part of which often became fused to form a cephalothoracic
shield. They had a circular mouth-slit in front of head, a single nostril just above mouth, one
or two unpaired fins and 5-10 pairs of gill slit. The tail-fin was diphycercal type. Animals of
cyclostomata group are represented by a few living agnathas such as Petromyzon (lamprey),
Myxine (hag fish) etc. Though surviving, they are primitive in many aspects and possibly they
represent the modified and degenerated off-shoots of the primitive vertebrate ancestor. They
can be readily differentiated from fish by the absence of jaw, tooth and paired fins.
Although some doubtful poorly preserved remains of ostracoderms are reported from
freshwater Ordovician rocks of Colorado, the first undoubted evidence of this form came from
marine Middle Silurian rock of England. Ty.to genera have been described of which Jamoytius
is worthmentioning as it~appears to be very primitive and might have occupied a position close
to the ancestry of lamprey and its allies (Fig. 23-1 b). It was more like a cyclostom with tubular
body, suctorial mouth, single nostril, lateral eyes, 8 to IO pair of gill-slits and a diphycercal
tail. Fossils of armoured fishes (ostracoderms) are well documented in Middle Silurian to Upper
Silurian rocks. One of the best known fossil ostracoderm is Cephalaspis, whose remains are
found within Silurian-Devonian rocks of England (Fig. 23-1 a, 23-3). It was a small, fish like
animal having an elongated but flat head, heavily armoured with a cephalothoracic shield, often
projected laterally into two horn-like structures. True ostracoderms appeared in Silurian,
diversified in Devonian but became extinct towards the end of this period. Possibly they evolved
from some primitive ancestral form like Jamoytius.
PALAEONTOWGY
294 Caudal fin
Dor. al Crest Of Scutc Dorsal Pin
fused Trunk
Median Nasal----,
Lateral Hom - - -
(a) CEPHALASN~ (ostracoderm)
Gill Slits---.
Buccal Sucker (b)PETROMYZON (cycloslomata)
FIG. 23 - I : EXTINCT AND LIVING JAWLESS VERTEBRATES
Chondrichthyes (Charcarodon)
Placodcnni (Dlnichtl,yes)
()sleichthycs-crossoptcrygian (lotim.irla)
Ostcichthycs-telcostct'' (La6-o)
Cheirolcpis (earliest actinopterygion) Ostcolcpis (earliest sarcoptcryaian)
FIG. 23 • 2: REPRESENTATIVES OF MAJOR GROUPS OF FISHES (sketeches not to scale)
GEOLOGICAL HISTORY OF VERTEBRATES 295
23.2 ADVENT OF FISHES . . .. .

Fishes with jaws have two broad subdivisions : (a) fishes with pnm1Uve type of Jaw structure
and (b) fishes with advanced type of jaws. Primitive group, extinct at present,. has two
morphologic types : acanthodii, the spine-bearing shark-like fishes and placoderm1, mostly
armoured fishes.
Aca11 thodians are popularly called "spiny sharks" for their superficial resemblance to modem
sharks. In these ancient fishes, there was an enlarged palatoquadrate (upper jaw) without teeth
and a teeth-bearing well-developed lower jaw. The jaw structure of acanthodians though primitive
but nevertheless, is basically the same as that of a bony fish. They had also a pair of gill-
coverings or opercular flaps over each gill-slit (somewhat like opercular flaps of higher fishes) .
They had a number of paired and an unpaired median fins, each consisting of a web of skin
supported by a strong spine. The tail-fin was typically heterocercal. Skin was protected by small
rhombic ganoid scales. Mostly they were fresh water fishes. One familiar acanthodian fossil is
Climatius of Late Silurian-Early Devonian age (Fig. 23-3). It was a small-sized fish (few inches),
with rounded anterior and tapering posterior, and they had two large eyes indicating their higher
sense of sight and predatory nature. Acanthodians, the earliest known ganoid fishes appeared
in Upper Silurian, reached their climax in Devonian and became extinct at the end of Palaeozoic.
Placoderms, another form of extinct Palaeozoic fishes had more advanced type of upper
jaw that was firmly fused to the skull. The head and shoulder region was heavily armoured
consisting of a series of large bony plates firmly joined with one another by sutures, while the
remaining part of the body was naked. The head was strongly arched across its top from side
to side so that the head could be moved up and down. The two most common Devonian
placoderms are Coccosteus and Dinichthyes belonging to arthrodire group. They were rather
large-sized animals and the latter often attained 10-20 feet length. Dinichthyes had a very strong
lower jaw with terminal plates projecting in the form of sharp teeth (Fig. 23-2). Sharp teeth
and movable head probably indicate that they were active predators.
From the heterogeneous morphology of the different types .of placoderms, it appears that
they were possibly a specialized group _of primitive fishes deviated away from the main line of
evolution. They have a short geological history as they appeared in early Devonian but mostly
became extinct with the end of this period.
23.3 ADVANCED FISHES

Advanced fishes may be placed in two groups, the chondrichthyes, represented by cartilaginous
fishes, like the sharks, rays and skates and the ostrichthyes that includes bony fishes.
Chondrichthyes : They include sharks and their allies which possess some characters
a~peared to be more primitive than true bony fishes. These are cartilaginous skeletons, open
gill slits, heterocercal tail, skull with a single skeletal element and primitive amphistylic jaw
attac~ment etc. But observing the abundance and success of modem sharks, many authors have
considered them, in spite of their possession of several primitive characters, as a specialized
group and their cartilaginous skeletons as the products of $econdary development.
On~ of the earliest sharks known from fossil record is Cladosellache (Fig. 23-3) of Upper
Devonian age reported from Cleveland Shales. It was a small animal compared to modem shark
PalaclGcuJWP-)~
PALAEONTOLOGY
296
Cheirolepis
Coccosteus
cp/1(1/aspis Os1racodcnns
FIG . 23 • 3 : EVOLUTION OF DIFFERENT FISH-GROUPS
GEOWGICAL HISTORY OP VERTEIJl<ATES 297
having a generalized morphology and it cooJd t,e ~ ~m ~ nus from whkh evolved all other
sharks and allied formb like pleurocanth .hark-", typical fil hatkti , skates, rays, bradydonts and
rat-fishe~. Thi s form probably survived upto the end of Palaeowic. Pleurocanth sharks evolved
during Carboniferous-Permian time bu in contr~1 to other sham, they were fresh water. They
possessed a long donial fin extending upto the diphxcercal tail, a peculiar Jong spine projecting
backward from the back of skull, bi-cusped teeth etc. All the~ features probably indicate their
evolution in a specialized direction.
Bradydonts were a small group of extinct pecialiud sharks (Permo-Carboniferous)
characterised by a flattened and wide tooth-bearing plate, often called pavement.teeth, indicating
their adaptation for crushing shell s of mollusc . One representative genus is Janess, reported
from Permian of Europe.
The earliest and the most primitive representative~ of modern sharks were hybodonts, which
had a long geological history from Late Devonian to Eocene. Characteristically, they had narrow-
based paired fins, unlike Cladosellache broad based fin ') and hence probably were more
efficient swimmers. Hybodus was a common Permian-Mesozoic form.
From Mesozoic onwards evolution of sharks proceeded along two different lines. One Jed
to modem predator sharks and other group to ray-fishes and skates. Typical sharks are elongated,
fast, aggressive, highly predaceous fi shes with a streamlined body, pointed head and a widely
gaping ventrally placed mouth equipped with many sharp teeth.
The skates and the ray-fishes are specialized sharks, mostly bottom dwellers with enlarged
pectoral fins (used like wings while moving through water), narrow pointed tails often projected·
in the form a whip and flattened teeth used for crushing shells. Rat-fish forms a separate group
among the modern shark living in deep oceanic water with an elongated pointed rostruin,
autostylic jaw and a tail projected into a long whiplash. They are ,ranging from Jurassic to
Recent. Fossil history of sharks is not very encourag·ing as their complete fossils are very
uncommon. More commonly, fossils of isolated teeth, spines, fragments of jaws, skulls are
found. Observing the surviving forms it may be concluded that they are contin1:1ing as dominant
marine animals since Carboniferous showing well adaptation to their environment and are quite
successful in holding their position in spite of invasion of other marine vertebrates (including
advanced fishes) with the same habitats.
Osteichthyes : Colbert has classified osteichthyes into two groups viz. actinopterygii including
ray-finned fishes and sarcopterygii including lobed-finned, air breathing fishes. The other .
subdivision of these two groups are as follows :
Actinopterygii : (i) Palaeoniscoida (Dev.-Perm.) : ancestral forms.
(ii) Chondrostei (Dev.-Rec.) : primitive forms.
(iii) Holostei (Trias.-Rec.) : intermediate forms.
(iv) Teleostei (Cret.-Rec.) : advanced fonns.
Sarcopterygii : (i) Crosspterygii (Dev.-Rec.) : lobed fins, supported by jointed bones.
(ii) Dipnoi (Dev.-Rec.) : ldbed fins, leaf like.
.,..
t.'ftl: . '"...
298 PALAEONTOLOGY
Geological history of bony fishes is quite a long one. Fossils of the first group of bony fish
are obt.ain~d from the Middl.e De~onian fresh_ wate~ rocks of E_n gland, belonging ;;
palaeo111sco1ds. The genus Che,rolep.Hs had a basic oste1chthyan cranial pattern from which
started the evolution of other bony fishes with complex skulls. Among the skull bones were .
a series of rostral bones, a pair of bones forming skull-roof on either side of temporal bones.
circumorbital bones surrounding eyes, a tooth-bearing upper jaw and an amphistylic lower jaw~
It had also paired and median fins and a heterocercal tail. Palaeoniscus was a Permian form
of this group. However, within their short duration (Devonian-Permian) palaeoniscoids followed
many lines of adaptation and specialization.
It has been assumed that palaeoniscoids were the ancestral forms from which evolution of
bony fishes proceeded passing through three general stages indicated by successive appearance
of primitive chondrosteans, intermediate forms holosteans and the most advanced and recent
forms teleosteans in Permian, Triassic and Cretaceous Period respectively (Fig. 23-3).
During the Triassic, appeared some advanced forms of bony fishes showing a homocercal
tail, thinning of scales and shortening of jaws. One of the representatives of these forms was
the common Triassic genus Redfieldia. This form is assumed as intermediate between
chondrosteans and the more advanced holosteans. The latter mostly replaced chondrosteans at
the end of Triassic, and exhibited further advancement with typical homocercal tail, thin scales,
more specialized skull and jaws, frequent ossification of vertebral centra, reduction of fin rays
and complete . loss of spiracles.
The first holostean, comparatively a generalised form of Mesozoic is Semionotus. Deep
bodied fishes like Depedius, and Microdon anived in Jurassic. Culmination of this group was
reached in Jurassic-Early Cretaceous times. Since the beginning of Cenozoic they were declining
in number with only two genera persisting today, Lepisosteus of Mississippi and Amina, the
bow-fin fish of northeast United States.
Since the Cretaceous, the declining holosteans were gradually replaced by the incoming and
ever-expanding teleosteans. In teleosteans, the skull becomes more specialized with shortened
jaws and concentration of teeth upon_ premaxilla. The~ sho~ perfec~ homocercal ~ail, co_mpl~te
ossification of internal skeletons, vaned types of modification of paired and unpaired fins with
migration of pelvic fins towards head, and development of ctenoid or cycloid type of scales.
In the geologic history, teleosts are represented by the primitive form Lepto/epsis appearing
in Jurassic showing intermediate characters between holosteans and teleosteans. After Cretaceous
teleosteans exhibited varied and complex lines of adaptive radiation that have established them
as the most successful among the creatures of the water-world. At .present, they exhibit all sorts
of variations in their shape, size and habit.
Considering the whole picture of vertebrate evolution we can see that ·teleosts, showing a
climax of their development among the water-living vertebrates of the present earth, a~ off
the main line of evolution that led to the higher vertebrates. In order to search for the history
ebrates other than fishes, we have to look back to another small group of bony fishes,
of ve rt .d h . I
th sarcopterygians that include the lung fishes an t e1r al ey crossopterygaans.
. Th ey ma de
th:ir first appearance as early us Devonian, but remain always suppressed under the vast group
of actinopterygians.
GE'OLOGICAl HISTORY OF VERTEBRATES 299
The most generalized dipnoids of Devonian, are Dipterus and Osteolepis of Old Red
Sandstone of England ( .. ig. 23-2). They are similar in some respect with the Cheirolepis, the
s1em actinoptcrygian, in having a heterocercal tail and primitive types of paired fins. Osteolepis
differs from Cheiro/epiJ in several other aspects which are as follows-
Clieirolepis Osteolepis
I. Heterocercal tail without epichordal lobe. Heterocercal tail with an epichordal lobe.
2. Fins supported by parallel fine-rays only. Fins supported by short muscular lobes with
internal axial bony elements.
3. A single dorsal fin. Two dorsal fins.
4. No pineal opening m between parietal A pineal opening present.
bones in skull.
5. No internal narial opening. Internal narial opening helping respiration in
• air.
6. Ganoid scales. Cosmoid scales.
All these differences may indicate that there was a basic divergence between these two lines
of bony fishes from the very beginning since their arrival.
A central line of dipnoan evolution led to Ceratodus, a widely distributed genus of Mesozoic
found in many freshwater deposits of the world whose direct descendant is the present day
Neoceratodus of Australia showing a little change of morphology from its Mesozoic ancestor.
The other two lung fishes, now surviving are Protopterus of Africa and lepidosiren of South
America.
From the Devonian Osteolepis evolved the other group of air-breathing fishes, the
crossopterygians. They, however took a separate line from dipnoids, showing increasing
ossification of skull and internal skeletons, development of sharp and pointed teeth adapted for
grasping preys (possibly carnivorous) with complex radial enamel-foldings (labyrinthodont
types). They also developed lobed-tins, (quite different from those of dipnoids) having a single
proximal bone articulated with girdle, below which there were two additional small bones and
beyond this were still smaller bones radiating outward.
Such feat.ures of early crossopterygians like more complex pattern of skull, possession of
labyrinthodont-type of teeth, more ossification of bones, more effective nostril and three-tier
bonc11 within lobed-fins may put them as the most probable ancestors of amphibia, the first
tetrapod on the earth and land ns well. From the basal stock Osteo/epis. evolution of
crossopterygians proceeded in two general lines, one leading to the fresh water rhipidistians
and the other to coelacanthids. The Devonian rhipidistian (rhizodont) genus Eusrhenopreron
exhibited a more developed prophetic skull like the amphibia and more advance type of venebraJ
column ending before the tail fin. There is little doubt that rhizodonts were very close to fish-
umphibiun transitional 1.onc.
However, unothcr group of crossopterygians, the coelacanthids was deviated far away from
the muin line of evolution. They were deep bodied, lobed-fin fishes showing much reduction
r -
300 PALAEON1'0LOG)'
of internal skeletal elements, symmetrical but diphycercal typ" of mil nnd rm extra lob d-tin
between upper and lower pair.
Coelacanth, first appeared in Devonian but shows incre·1s " in numb ·r in M~s z.oic. The
typical fossil is the Cretaceous genus Macropoma. This was the last fossil of c lacunrhids mer
in the geological sequence and they were once thought extinct after thut period until I93 when
a living coelacanth was caught in South Africa. The specimen has been found r markabl) similar
with the Cretaceous form Macropoma. More living coelacanths have been a nilabl after I 5_.
The living genus is named as latimeria. From its remarkable similarity with thos tnci nt
fossils, latimeria is called a 'living fossil' (Fig. 23-2).
One interesting feature found in the evolution and development of bony fo,hes is rhar one
group of fishes were replaced by another with intervals of geologic time. The group whi h had
replaced the earlier one however retained many of the morphological features acquired by rhe
pre-existing group in response to their variable adaptations. It is quite obvious that for a life in
water, aII animals require some common structures such as a streamlined body, a tuilfin,
respiration by gills, large mouth with efficient teeth (for carnivoras). paddle-like pectoral und
pelvic zones and so on. As all the fishes, primitive or advanced type, appeared in diff-rent times,
had to confront the similar environmental problems, they tried to solve them in the same manner.
The result was the repeated development of similar forms and structures in different groups of
fishes at different geologic times (parallel evolution). But as new forms, evolved by means f
genetic processes and natural selections, show increasing efficient mechanisms in coping with
environments, they were able to replace the older pre-existing forms (ecologic replacement).
Thus deep-bodied chondrosteans were replaced in Triassic by holosteans of simi Jar forms ,, hich
in turn were replaced by teleosteans in Cretaceous (many of which also exhibit similar de~p
bodied forms). Although, chondrosteans were well-adapted to their environment ut th ir times.
holostean could be able to replace them because they were more efficient and the same ,ms
the fate of hoJosteans at the end of Cretaceous when they were replaced by teleosteans. Fig.
23-3 shows a diagramatic sketch of phylogeny of fishes.
23.4 AMPHIBIAS-THE FIRST TERRESTRIAL VERTEBRATES

Amphibias, the first group of terrestrial vertebrates, possess two distinctiv~ features;
externally, two paired lim~s with digits ~hich are the result of gradual chang~· of the 1uin"<i
lobed-fins of crossopteryg,an fishes and internally advanced type of lungs for respiration in
air. Moreover, amphibia's skin, Jacking scales, becomes much thicker thnt prot cts it from e, et:$$
Joss of water in open air.
Fossil and Jiving amphibias are broadly classified into three:! group whi -h ure as foJJm, s-
Cl.ass : Amphibia
Subclass Labyrinthodontia (Dev.-Tris.) : primitive-most nmphibius.
Subclass ; Lepospondyli (Carb.-Perm.) : Lute Palaeozoic umphibius.
Subclass : Lissamphibia (Trias .• Rec.) : modern umphibius.
Labyrinthodonts are the most primitive group of nmphibins chun1ctt1risc.•d hy t Sf.k'Cial r., flt?
of teeth with enamel on tooth-surface 8howing complex sinuous infoldings. Th~ first £'r up of
~
-
~
b
-
~
:::;
!-
-'
~
V,
0
~
~
C
~
(a) Eryops (Labyrinthodont)
EXTINCT --::::
~
::,;,
~
~
to
~
~
V)
(c) Indian Salamander

(b) Diplocaulus
SURVIVING EXTINCT
FIG. 23 - 4: E~TINCT AND SURVIVING AMPHIBIAS ~

0
J02 PALAEONTOLOGY
Supruorbital
Prcorbitnl - -..
L11crym11I
Nasal--.....
Squamosol (SQ) (1) SEYMOURIA

Prcmaxilla
Jugol
Maxilla ~ - -
Eye orbit4----- - ~ Qundratojugal
Anapsida
- - - Temporal
Opening (II) COTYLOSAUR ( Anapsida)
Euryapsidl\
(Ill) PLESIOSAUR ( Euryapsida)
Parnpsidl\ (IV) ICHYOSAUR ( Parapsida)
Synnpsida
I\' 1 SPI II'. .\' ..\( ·1>I>! >Y I' 1.\rn,tpJi tlo1 1
Supratcmporal Fossa
Oiapsid11
(VI) PTEROSAUR (Diopsida)
FIG. 23 - S : SOME EXTINCT REPTILES AND THEIR BASIC SKULL-STRUCTURE (not to scale)
labyrinthodonts arising from lobed-fin fishes are icthyostegids. The earliest fossils lc/1t/zyos1egn
and Acantlzostega are collected from fresh water Upper Devonian rocks of Greenland.
lchthyostega was a small animal having a fish-like tail and vertebral colunrn but limbs like a
tetrapod (Fig. 22-3). In general, icthyostegids may be considered some trnnsitional forms between
the ancestor crossopterygian fish and a true amphibia. They possessed vertebrae and vertebral
columns like their ancestor and also retained a fin on a rather elongated tail. Like their ancestor.
they had external nasal openings far down on the margin of the skull which were separnted
from internal narial openings only by a thin bony bar. But the most significant advancement
was the elongation of rostral portion of the skull in front of eyes and shortening of post-parietal
portion which is just reverse in fish. Acantlwstega and /chthyostega are the oldest tetrnpods
belonging to labyrinthodont amphibias. Skeletal elements of limbs of these two forms are nearly
the same as the later tetrapods. Recently, Coates and Clark ( 1990) have collected Accmtlwstega
with eight digits in hand and ld1thyustega with seven digits in foot. More advanced chmacters
were exhibited by younger labyrinthodonts of Carboniferous-Triassic times. They had external
nostrils shifted on dorsal surface of the skull which were separated from internal narial openings,
located in front of palatal region. There was a well-defined nasal passage leading from externul
nostrils into the throat and this is the basic plan for air-intake, adapted by all subsequent
terrestrial tetrapods. Advanced labyrinthodonts also developed advanced type of vertebrne euch
with the centrum convex on one side and concave on other. This gives interlocking joints of
vertebrae making the vertebral column more strong and flexible. They had also stronger ..md
larger pectoral and pelvic girdle and enlarged limb bones to bear the weight of the body. At
the back side of skull, a tympanic notch for accommodation of eardrum is also a new character
of an amphibia.
In general, labyrinthodonts, are usually grouped into three divisions viz : emblomere group,
rhachitome group and stereospondyl group. The first group. probably evolved from
ichthyostegids, had two discs in each vertebrae one behind the other named inlerce11tr11111 and
pleuroce11/rmn within which located neural arch and neural spine. This group is well represented
by the Carboniferous Eogyri11us. The rhachitome labyrinthodonts, present between Late
Carboniferous and Permian Period, possessed a keel-like intercentrum, slightly larger than
pleurocentrum. The most significant was the Permian form El)ops which had 2-5 meter long
body, a heavy and flattened skull with a wide muzzle, a strong backbone and a nodular skin
(Fig. 23-4a). The other Permian forms were Ard,aegusaurus (having a long snout)
T,.imeror'1ac/1is and Branchiosaurus. This group of labyrinthodonts were considered as the
ancestor of all the modem amphibias. The third group, the stereospondyls, followed a different
line, showing an adaptation mainly to an aquatic life. They had a larger size but with simple
Vertebrae and a relatively weak backbone. One representative genus is the Triassic form
Bueunaria. It can be said that arising from Devonian ichthyostegids, labyrinthodonts followed
two lines of evolution. Along one line appeared anthracosaurs (emblomcres) and rachitomcs
Which were adapted to more successful life in land, while stereospondyls, appeared on the other
line, were adapted for an aquatic life.
Lepospondyls were small-sized amphibias nearly contemporaneous with labyrinthodonts but
adapted to a separate ecological niche not explored by the latter. They mostly lived in swamps
or near-shore zones. Some of them like Permo-Carboniferous Ophiderpeton had small snake-
Palac(Geo)WP-39
- Scanned by CamScanner
304 PALAEONTOLOGY
like body, with suppressed legs. Some developed flattened body and a very broad and flat skull
as shown by the Permian genus Diplocaulus (Fig. 23-4b).
The modern group, the lissamphibias, are first recorded from Triassic rocks. There is a
considerable gap in the fossil record of amphibias for which palaeontologists have failed to
connect the extinct Permo-Carboniferous amphibias with their recent counterpart. However, the
general traits of modern amphibias are (a) reduction of bones in skull and also its increase of
flattening (b) two ossicles in middle ear rather than one found in the extinct group (c) presence
of a zone of weakness between the base and crown of the teeth. (d) possession of four digits
(five in case of fossil groups) in forelimbs.
There are two groups of modern amphibias. The first group are frogs and toads (anuras)
whose fossils are first found in Triassic, represented by the ancestral frog Triadobatrachus;
The second group, represented by salamanders and urodales, are mainly water-living or
burrowi_ng amphibias now living in tropics (Fig. 23-4c) . .
23.S REPTILES-THE RULERS OF MESOZOIC (Fig. 23-5)

A. Amniote egg
Reptiles are the first tetrapods capable of producing shelled eggs (Fig. 22-4). These amniote
eggs, fertilized internally are laid by female on ground in some suitable place. The egg contains
the embryo covered by amnion liquid. The egg also contains a large yolk for nourshing the
embryo and an allantois for storing the waste product produced by the embryo during its
development. The entire structure is enclosed by a hard but porous shell which not only protects
the egg, but also allows air to pass through it for the respiration of the embryo.
Appearance of shelled eggs has made possible for terrestrial tetrapods to colonise in land
over diverse types of conditions and this may be taken as another great leap in the evolution
of vertebrates making the terrestrial life independent of water.
B. Broad subdivisions
Reptiles were more abundant in the geological past especially in the Mesozoic Era. They
may be called the rulers of that period when appeared numerous groups, adapted to variable
types habitats in land, air and also in water but most of which however became extinct abruptly
at the end of Cretaceous. In the classification of the reptiles palaeontologists have to include
all the living as well as the extinct groups. A generalised classification upto the level of order
may be given below mainly following the scheme of Colbert ( l 969) and Romer (1966).
Class : Reptilia
Subclass : Anapsida : Primitive reptiles without temporal opening.
Order : Cotylosauria : (Carb.-Trias.) : stem reptiles. (e.g cotylosaurs)
Order : Chelonia : (Trias.-Rec.) (e.g. turtles).
Subclass : Synapsida : Mammal-like reptiles, skull bearing a single temporal opening below
postorbital and squamosaf bones.
Order : Pelycosauria (Carb.-Perm.) : (e.g. pelycosaurs).
Order : Therapsida : (Perm.-Trias.) : (e.g cynodonts).
GEOLOGICAL HISTORY OF VERTEBRATES
305
Subclass : Eury.apsida : Generali~ marine, aquatic reptiles, skull with a single temporal
openmg above postorb1tal and squamosal bones.
Order : Protosauria (Perm.-Trias.) : (e.g. protosaurs).
* Order : Sauropterigia (Trias.-Cret.) : (e.g. plesiosaurs).
Order : lchthyosauria (Trias.-Cret.) : (e.g. ichthyo·saurs).
Subclass : Diapsida : Reptiles with two skull-openings, one separated from the other by post-
orbital and squamosal bones.
Order : Rhynchocephalia (Trias.-Rec.) : (e.g. sphenodonts).
Order : Eosuchia (Perm.) : (e.g. Petrolacosaurus)
Order : Squamata (Jura.-Rec.) : (e.g. lizards, snake).
Order : Thecodontia (Trias.) : (e.g. archaeosaurs).
Order : Croodilia (Jura.-Rec.) : (e.g. crocodiles).
Order : Pterosauria (Jura.-Cret.) : (e.g. pterosaurs).
t Order : Saurischia : (Trias.-Cret.) : (e.g. sauropods).
t Order : Ornithischia (Trias.-Cret.) : (e.g. omithopods).
C. Geological history
Reptiles evolved from some groups of labyrinthodont amphibias. This transition probably
took place in Carboniferous. The oldest amniotes are Hylonomus and Palaeothyris from Middle
Carboniferous of Nova Scotia (Carrol; 1964, 69). It is appropriate to mention here about the
form Seymouria which shows a morphological transition between an amphibia and a reptile
and its fossils are found in Lower Permian Seymour of Texas (Fig. 23-5). It resembles the early
amphibia in having a similar type of labyrinthodont-teeth and skull structure. But it shows lizard-
like pleurocentrum, wide neural arch and wide ilium and sacrum. Limbs bones were also much
advanced type. As fossils of undoubted reptiles are found well before Permian (Upper
Carboniferous), Seymouria probably was a form very close to ancestral reptiles. There is still
some doubt about its true affinity that is whether it was a reptile or an amphibia. Unfortunately,
there is no record of fossil egg of Seymouria.
(a) Anapsids : Cotylosaurs were the first true reptiles on the earth which appeared towards
the end of Carboniferous. From their primitive but a generalized characters they are supposed
to be stem reptiles from which evolution and diversification of all other groups took place.
Even they could be taken as the distant ancestors of birds and mammals.
From morphology of cotylosaurs it appears that from the ancestral reptile, evolution
proceeded mainly along two lines represented by two groups of cotylosaurs; captorhinomorphs
were small carnivorous group while the other diadectomorphs were large herbivorous forms.
The former group is represented by Hylonomus (Carboniferous of Nova S~oti~) and Limnoscelis
(Lower Permian, New Mexico). The diadectomorphs were much larger m size ( I to 2 meter).
The representative genus Diadecte.v has been reported from Permian of Texas. Its skull was
• Some considered animals of this order belonging to a separate extinct subclass Parapsida.
t Together called 'Dinosaurs'.
-----·
PALAEONTOLOGY
306
large with quadrate bone shifted forward. Frontal teeth were much !arger. Another group. of
cotylosours, the specialized procolophonids, were the only forms ~h1ch were a_ble _to survive
above the Permian boundary. The genus Procolophon attained a wide geographic_ dispersal in
Triassic and their fossils have been reported from Permo-Triassic rocks of Russia, Scotland,
North and South Africa and Central Europe.
Turtles (chelonias), the direct descendants of cotylosaurs have retained a solid skull roof of
primitive reptiles. The most characteristic specialization of the turtle is the development of a
bony dorsal carapace and a soft bony plastron on ventral surface. Their limbs developed into
flipper-like paddles.
True turtles made their first appearance in Middle to Upper Triassic. Proganoche/ys is a
characteristic Triassic genus. Two groups of turtles arose in Mesozoic and are still continuing.
These are: pleurodires (found in South America, Africa, South Asia and Australia) and
cryptodires, the most widespread forms of present day found throughout the world.
A few earliest derivatives from cotylosaurs ancestry in Permo-Carboniferous periods were
synapsids, euryapsids and diapsids.
(b) Euryapsids : One of the earliest known reptile groups are mesosaurs represented by
the Mesosaurus, reported from Carboniferous rocks of South Africa. They were small elongated
aquatic reptiles with long tail and limbs modified to paddles. In many aspects (neural arch)
they resemble cotylosaurs. As skull of this animal is not well preserved, it is difficult to put
them in a particular category. May be they represent some early forms of euryapsids or
transitional forms between cotylosaur and euryapsid. True euryapsids had their beginning at
Permian with their ancestral forms protosaurs. They soon became well adapted to a life in
water and showed all sorts of secondary modifications needed for such life as paddle-like limbs
and a tail or a tail-like structure. Starting from this earliest protosaurian ancestry euryapsids
exhibited some distinct lines of development from Triassic onwards, and the Early Triassic
placodonts probably marked this point of divergence. Placodonts, represented by the genus
Placodus, were adapted to the shallow marine water, feeding upon molluscs. It has a stout body,
paddle-like limbs, a short neck, a tail, a ventral rib-busket and possibly also armours. From
placodonts diverged two groups : long-necked and short-necked forms. The short-necked form
was more li.ke a fish with a large elongated skull, fish like fins (two paired and one median
dorsal) and a reversed heterocercal tail. Some well known fossil-forms are Triassic
Cyrnbosp~nd~/us and Jurassic-Cretaceous ~chthyosaurus. The long-necked forms are represented
by the Tnass1c genus Nothosaurus, Jurassic-Cretaceous genus Plesiosaurus and the Cretaceous
genus E/asmosaurus. They exhibited ~ore re~tile-like body with a small head, a long and
flexible neck and a muscular fin-less tail. P/eswsaurus and /chthyosaurus [Fig. 23-5(iii)-(iv)]
were the two most successful forms of t~e sea in Jurassic-Cretaceous times attaining a huge
size (3- J2m). All these forms became extinct at the end of Cretaceous.
(c) ~ynapsids : Another ~arliest der~vatives _(Pe.rmo-Carboniferous) of cotylosaurs were the
synaps1ds, popularJy cal!ed mammal-l,ke_reptiles . They are also interesting in the sense that
they _are the probable ~ndge between rept1_les and mammals. Synapsids had a lateral temporal
opemng behmd each side of the eyes ~nd m earlier forms this was bounded by the postorbital
and squamosal bones. They were pers1stant quadrupedal and in this respect they differed from
. .,} ' ....
ther reptiles. The other characterist~c features which have made them remarkable are the
0
retention of most of the
. bones,
. and. high degree of dental specialization towards heterodontid
pattern with separate mc1sors, canme and molars. All these features put them in the line of
evolution of mammals. -.
The first synapsids pelycosaurs found in Carboniferous-Permian of Texas, Oklahama, and
New Mexico had primitive and generalised features. They may represent some transitional forms
between cotylosaurs and synapsids. A more advanced Permian pelycosaur is Ophiacodon which
can be regarded as a stem-genus in the evolution and divergence of synapsids. It had a medium-
sized body, a short neck, slender limbs and a long tail. From this, evolution proceeded mainly
in two directions. One line led to carnivorous sphenacodonts and the other gave rise to small
plant-eater edaphosaurs. Sph_enacodonts had strongly differentiated large dagger like teeth in
premaxilla at frontal part and smaller teeth at lateral parts of jaws . .But the most spectacular
feature of their body was a longitudinal sail-like structure all along the mid-dorsal surface which
was composed of elongated spines of vertebrae from the neck back to the sacrum, reaching
their maximum height at the middle. These spines, most certainly had a support of skin, that
produced this sail (Fig. 23.Sv). There are many suggestions about the probable function of this
structure. It might have functioned as a protective device or a feature related to a particular
sex or even it might have functioned in controlling body temperature. The most common
Pennian representatives are Sphenacodon of New Mexico and Dimetrodon of Texas.
Edaphosaurs, in the other line, had smaller skull compared to their size with teeth less
.differentiated but their elongated vertebral spines were more stout having some lateral crossbars.
Therapsids appeared from pelycosaurs in Late Permian-Triassic times in South Africa. Though
derivatives of pelycosaurs, they show a number of distinctive features. They are in the main
line leading to the mammalian evolution. Its skull showed a temporal opening having a tendency
to shift laterally and a separate nasal passage from mouth with the appearance of a secondary
palate below the original reptilian palate. There was a strong differentiation of four types of
~h. Some had two occipital condyles like mammals. Post-cranial skeletons were also distinctly
differentiated into neck, thorax, and tail. Elbows of forelimbs point backward and knees of hind
limbs point forward. That might have increased its efficiency in locomotion .
. Permian therapsids gave rise to two groups of Fermo-Triassic synapsids, one much larger
dmoeephalians and the other, the small-sized .theriodonts. Dinocephalians were a specialized
group with large size (several meters), massive head and neck and a short tail. Except a pair
of. canine other teeth were similar and small-sized. A few Permo:.Triassic genera are Moschops,
Dtcynodon and Lystrosaurus. The last genus was very much widesprend during Lower Triassic
Period and was a common inhabitant of Gondwanaland.
The more significant is the theriodonts which were inhabitants of Middle Triassic of many
Gondwana continents. The animal had small size like a dog with comparatively large skull, a
large temporal opening behind eyes, a parietal bone and well-differentiated teeth. It showed
complete separation of respiratory tract and buccal cavity, large ribs, well developed vertebral
column and a tail. Pectoral and pelvic bones resembled those of mammals. One of the most
:n:1mon theriodonts was Cynognathus of Triassic. Some forms known us tritylodonts. seemingly
enved from cynodont ancestry, was very much mammalian in features like advanced type of
308 PALAEONTOLOGY
zygomatic arches, secondary palate and specializ d t cth. Th· g "nus Trityl~do~ is .known from
South Africa. The most advanced group of theriodonts. th~ ictidosaurs ot Tn llss1c to Middle
Jurassic age, probably represent the conne ting bridg ,· between m:immuls and r~ptiles. In
ictidosaurs. the most of the chan1cters of the th riodont m ntion d ·arher r ·ached n high degree
of perfection. But the most characteristic features wer th· loss of posto.r~il, prefrontal,
postfrontal bones, and a double jaw articulation like mammals, although. the JOmts were non-
functional.
(d) Diapsids : The diapsid has two temporal openings on ach side of the skull. Earliest
diapsid eosuchians, derived from cotylosaurs. are reported from the Permo-Carboniferous rocks.
The two common genera are Petralocosaurus from Carboniferous of Kansas and the other one
is Youngia from Pem1ian of South Africa. Yo1111gia were probably the founder of all other diapsids
of Mesozoic period, some of which are still surviving such as crocodiles. lizards, snakes and
rhynchocephalians.
The earliest crocodiles, evolved in Penno-Triassic, were the protosuchians which in Mesozoic
gave rise to the mesosuchians. From Cretaceous two lines diverged from mesosuchian : the
extinct group is sebesuchian, represented by Sebecus (Eocene) of Patagonia and the Cretaceous
Baurusuchus of South America. The line towards the modern crocodiles is formed by the
eusuchians which made their first appearance in Cretaceous and became diversified in Cenozoic.
The Cretaceous Plwbosuchus was the largest known crocodile fossil, collected from the Rio
Grande river of Texas which had a skull of six feet length. A short-lived gigantic group was
the Mesozoic phytosaurs.
Rhynchocephalians made their first appearance at the beginning of Triassic but might be
present before that. They were common forms of Triassic represented bY. numerous fossils of
rhynchosaurs reported from various parts of the world including Africa, India, South America
and also from other parts of the Gondwanaland. Their size was fairly large often 5 to 6 feet in
length. After Triassic they became very much restricted. Now they are inhabiting on a few islands
of New Zealand and are named Sphenodon, popularly called tuatara.
The order Squamata includes the lizards and snakes which are by far the most numerous
and diverse among the surviving reptiles. The earliest report of their fossils comes from Lower
Triassic of South Africa. The genus Prolacerta, classified by some authors as an eosuchian,
had various lizard-like characters. It may represent an intermediate form between the ancestral
eosuchians an~ the descendant squa~atas. A typical squamata is characterised by a single upper
temporal opening above the postorbttal-squamosal bar but below this the cheek region is open
as there is no lower bar produced by ~uadratojugal as in other reptiles. This gives more mobility
of joint between skull and the_ lower Jaw. These are quadrupedal reptiles with hind limbs highly
elongated that act as a propulsive force so that the animal can stand on hind limbs while running
or before jum.ping: Lizards be~ome we!I establishe~ since Jurassic onwards and they have
evolved and d1vers1fied al~ng dtffere~t hne~ of adaptive radiation. Among all reptiles, snakes
appeared last. They are highly modified hzards where limbs are absent and locomotion is
accompanied by various movements of the body. The tail and body are undifferentiuted and
they together become greatly elongated. The skull is also somewhat specialized with the double-
joint jaws which give them greater mobility so that they can spread apart their jaws to a
309
considerable degree. Unfo~unately, the fossil history of the snake is rather poor) known for
Jack of well-preserved fossils. y
. Among t~e groups of interesting reptiles of the Mesozoic, are the flying reptiles pterosau~s.
[Fig. 23-5 (v1)]. In fact~ t~o g~oups of tetrapods, pterosaurs and early birds, quite independently
~came adapted to a hfe m atr. However, the mechanism of flight shown by the living birds
ts very compl.ex a~d whether pter~saurs which became extinct after Cretaceous, adapted the
sai:ne mechamsm 1s a moot quest10n, especially when reptiles are, in general, cold-blooded
ammals.
Rhamphorhychus is possibly the oldest known pterosaur of Jurassic age which had a body
of.about 2 feet length with p(imitive skull having two temporal openings behind the large eyes.
The jaws were elongated with long pointed teeth, adapted for fish-catching. It had also a long
flexible neck and a tail and their forelimbs were modified into two large leathery wings. The
highly elongated fourth-finger along with a' ~pike-like pteroid bone from wrist gave the major
support to the wings. Other fingers were reduced to hooks helping it hanging from tree-branch
or rock cliff. Coracoid and scapula were strong with enlarged sternum. Hind limbs were stout
having digits with sharp claws for catching the preys. This was more or less a generalized view
of these Mesozoic reptiles wih few modifications iri advanced forms. Pterosaurs reached their
culmination in Cretaceous represented by the form Pteranodon. This was a giant reptile with
•
a wing-spread of more than twentyfive feet. The jaws were long with tooth-less beaks. The
back portion of skull was extended posteriorly into a crest. This was the last member of the
group which became extinct at end of Mesozoic along with many other reptiles.
Triassic thecodonts are considered direct ancestors of dinosaurs but compared to dinosaurs
they were small reptiles with narrow skull lacking pineal opening but had diapsid-like two
temporal openings on either side. The most striking feature of these animals was their bipedal
mode of locomotion running on hind limbs which were much elongated upon which the body
was balanced as a fulcrum. The height of the body was counterbalanced by a long tail which
also partially bore the weight of the body. The forelimbs were mainly used for grasping food
or preys. For obvious reason, the attachment of pelvic girdle with vertebral column became
considerably strong. This was essentially the basic body plan of dinosaurs which they i~herit~d
from thecodont ancestry in Upper Triassic. The ~ormer never completely gave up this basic
structure throughout their long history although, many of them returned to a quadrupedal
locomotion. Lower Triassic Erythrosuchus of South Africa, l.Jpper Triassic Ornithosuchus and
Hesperosuchus are few fossil genera of thecodonts.
. ·f th odonts 1•0 the later part of Triassic and they continued as the most
D mosaurs arose rom ec . .
· 1 d ·1 th
dommant an rept1 es roug o h ut the Mesozoic. Two groups of dmosaurs have been recogmsed;
. h' h' h · b" d rk h'
one saurischia with a lizard-like pelvic joint and the other omit isc ia s owmg a tr - 1 e tp-
joint.
· h' 11 bl d thei·r thecodont ancestors. .They .include several groups showing
Saunc aans overa resem e . .
a varied size and habitats. Some were small carnivoras, hke Tna.ssac Coelophys,s; some were
· rn rus of Jurassic-Cretaceous age; some were
large aggressive carnivoras hke Allosaurus, ,yranosau . ) d the others
. d b'pedals
1 (mostly carmvorous group an
giant herbivors hke sauropo s; some were , ninety feet as found
quadrupedals. Size of dinosaurs is ranging from about a foot to as many as
PALAEONTOLOGY
110
Cenozoic Turtle
Lizard t
Crocodile
Sphenodon
Crctaccou
Jurassic
I
I
I
Therapsida
Triassic
Permian
Up
Carboniferous
t
Labyrinthodont amphibia
FIG. 23 -6 : DIVERGENCE OF REPTILES IN MESOZOIC
Recent
e
C
>
j
u
e
~ ~> c• !! 5
u ·g ]C
«I
:i
bl) -~
-~ C
=
5
....~ u
C
u ><
Condylartha 1 ; r ~ -
~
~ --f---,
lnscctivora
Mcsomic Early Mesozoic M111n111nts

I
Syn11psid Reptiles
FIG. 23 - 7 : DIVERGENCE OF EUTHERIAN MAMMALS JN CENOZOIC
GEOLOGICAL HISTORY OF VISlffEIJIU'J'l·:S 311
within many herbivorous samoµod s ul' Junt 1' i ·-('r ·t; • •ou tin · , uch a Brachio.wurus,
Brontosaurus, Diplodocus, Gi>f(mlOsouru.v t ·. ( 'amivorou. ~1r ,u1 had c mparativcly a large head
and very sharp teeth on their 'jaws. Muny di110Nau r. w ·re a mo11r d uch as Ste~o auru and
Ankylosaurus, others bore horn wlik • structu r · on th ·ir h ·ad " . hown by Crctaccou
ceratopsians.
However, this large group of reptiles, with a Ion , ·ok ic~al hi. tory (a.bout 170 million year. )
showing varied structures and habitats and a wid · , ·u •raphic di. pcn,al, . uffcrcd an abrupt extinction
after Mesozoic that needs a special att ·ntion, and hcnc.:t: "re di su, ~. ·din detail in Chapter 24. A
diagramatic sketch of divergence of reptiles durin M ·, ozoic i shown in fi g. 23-6.
23.6 HISTORY OF BIRDS

Bird arose in Jurassic possibly from some hipedal thccodonts or some thcropod dinosaurs.
Birds are very similar to reptiles except. that they take more advanced methods of adaptation
for flight which are very different from that of pterosaurs. Birds arc warm-blooded animals,
with feathers covering the body. Besides a pair of feathery wings, birds also possess, light bones,
number of air sacs and a special type of sternum hone. All these features have helped birds to
eliminate their Mesozoic rivals, the pterosaurs and have made them the most successful flying
creatures of the modern earth.
Fortunately, the preservation of the fossil of the first bird is good enough which gives us a
fairly complete picture of its morphology. The fossil is named Archaeopteryx found from Late
Jurassic Solenhofen Limestone of Bavaria, Germany (Fig. 22-3). The skeleton is remarkably
complete with impression of feathers also. The animal, of the size of a crow, had a primitive
thecondontid skull, a long neck, a compact body, strong hind limbs and a long tail. The skull
had two temporal openings bearing eyes with sclerotic plate, and beak provided with teeth . The
pelvic joint was quadruradiate (characteristic of bird) and the tail was more reptilian type. This
is truly an intermediate form where skeletons are more reptilian but other features are bird-like.
Recently Chatterjee (1991, 1995) has reported a still older (Upper Triassic) fossil bird
Protoavis from Dockum Group of Texas. Although, details of morphologic description has been
given by the author of this form, some palaeontologists declined to accept his claim firstly
because the specimens are much fragmentary and secondly it is a fossil out of time. If it is
accepted, Protoavis extends the fossil record of birds and theropod dinosaurs (probable ancestor
of birds) back more than 80 m.y. thus creating a long gap in a single lineage leading to Late
Jurassic Archaeopteryx.
By Cretaceous times birds became more advanced with the fusion of skull bones
(characteristic of modern birds) and by the development of high degree of pneumaticity in the
bones of skeleton. Pelvis and sacrum were fused to form a single structure serving a strong
joint between the hind limb and the body. With the beginning of Tertiary, birds attained all its
modern morphologic features. The radiation of ralites (ground-living birds) probably took place
in Cretaceous. Most of the modern birds are remarkably similar in their internal structures and
their great diversity is the result of their adaptation to various habitats. Unfortunately, the fossil
history of birds is rather incomplete for lack. of suitable f~ssils in Cenoz~ic. ~ wel.l known
Cretaceous bird fossil is Hesperornis, found m Kansas. D,atryma, an ostrich-like bird from
Eocene of Wxoming, grew to as much as seven feet tall.
Palae(Geo)WP-40
' 312
23.7 MAMMALS-RULERS OF THE PRESENT
PALAEONTOLocy
Mammals are the descendent of some mammal-like reptiles, probably the therapsids. The
transition took place in Late Triassic, but these early mammals remained subdued under the
dominant reptiles throughout Mesozoic from Palaeocene o~wards they exhibited explosion of
evolution with complex and varied patterns of adaptive radiation. Starting with only five orders
in early Palaeocene, as many as seventeen groups of mammals appeared by the end this period.
Before going into their geological history, we have to have some idea about these different
groups of mammals all of which made their appearance by Quaternary. The classification given
below has broadly followed that of Colbert ( 1961 ).
Class : Mammalia
Subclass : Eotheria : Very primitive Mesozoic mammals.
Order : Docodonta (Trias.-Cret.) : e.g. docodonts.
Order : Triconodonta (Trias.~Cret.) : e.g. triconodonts.
Subclass : Prototheria (Jur-Rec.) : Egg-laying monotremes.
Subclass : Allotheria : Late Mesozoic-Early Tertiary mammals.
Order : Multituberculata (Jur.-Eoc.) : e.g. kamtobaator.
Subclass : Theria : Placental mammals.
Infraclass : Metatheria : Pouched mammals.
Order : Marsupialia (Cret.-Eoc.) : e.g. kangaroos.
Infraclass : Pantotheria : Ancestral placental mammals.
Infraclass : Eutheria : Advanced placental mammals.
Order : Insectivora (Cret.-Rec.) : e.g. shrews, moles.
Order : Dermoptera (Palaeo.-Rec.) : e.g. colugo.
Order : Taeniodonta (Palaeo.-Eoc.) : e.g. taenidonts.
Order : Tillodontia (Palaeo.-Eoc.) : e.g. tillodonts.
Order : Chiroptera (Eoc.-Rec.) : e.g. bats.
Order : ~entata (Palaeo.-Ref) : e.g. ground sloths.
Order : Pnmate (Palaeo.-Rec.) : e.g. man, monkey and ape.
Order : Rodentia (Palaeo.-Rec.) : e.g. squirrels, rats.
Order : Lagomorpha (Palaeo.-Rec.) : e.g. rabbits.
Order : Cetacea (Eoc.-Rec.) : e.g. whales.
Order : Creodonta (Cret.-Oligo.) : e.g. primitive camivoras.
Order : Carnivora (Palaeo.-Rec.) : e.g. cats, dogs.
Order : Condylarthra (Palaeo.-Eoc.) : e.g. ancestral ungulates.
Order : Litoptema (Palaeo.-Pleis.) : e.g. camel and horse-like ungulates.
Order : Notoungulata (Palaeo.-Pleis.) : South American harbivore ungulates.
Order : Astrapotheria (Eoc.-Mio.) : South American harbivore ungulates.
Order : Tubulidentata (Plio.-Rec.) : e.g. aardvarks.
GEOLOGICAL }{/STORY OF VERTEBRATES 3l3
Order : Pyrotheria (Eoc.-Oligo.) : large South American tusk-bearing ungulates.

Order : Pantodonta (Palaeo.-Oligo) : large ungulates of South America.
Order : Embrithopoda (Oligo). : gigantic rhinoceros-like mammals.
Order : Proboscidea (Eoc.-Rec.) : e.g. elephants.
Order : Perissodactyla (Eoc.-Rec.) : e.g. horse, rhinoceros.
Order : Artiodactyla (Eoc.-Rec.) : e.g. catties.
Order : Sirenia (Eoc.-Rec.) : e.g. dugong, seacows.
Order : Hyracoida (Oligo.-Rec.) : e.g. hyrax.
Order : Dinocerata (Palaeo.-Eoc.) : e.g. archaic herbivores.
A typical mammal should have following skeletal characters, (i) dentary hinged to squamosal,
(ii) double occipital condyles. (iii) A palate bone separating nasal passage from mouth,
(iv) comparatively a large brain case, (v) completely differentiated four types of teeth.
About 20 groups of primitive mammals have been reported from Mesozoic (Kemp, 1982,
Szalay et al., 1993) but most of these fossil materials are poorly preserved and incomplete and
their relationship is difficult to establish. Among these, morganucodonts, triconodonts,
symmetrodonts, pantotherids are rather well known. However, the two primitive-most presumed
mammal fossils are Adelobasileus reported from Late Triassic of Texas (Lucus and Luo, 1993)
and Sinocodon from Early Jurassic of China. Brain case of both of them shows a number of
features characteristic of early mammals but their teeth are rather undifferentiated like their
cynodont ancestor.
Morganucodonts (Upper Triassic-Middle Jurassic), reported from Europe, China, South Africa
and England and represented by the genus Morganucodon and Megazostrodon, exhibited
multicuspidate molars with double root and the cusps arranged in a single series on upper surface
of tooth in upper jaw and two series in lower jaw (Fig. 23-8a). Triconodonts known from
Europe, North America and Asia exhibited a clear differentiation between molars and premolars
(J~nkins and Crompton, 1979) which had three principal cusps arranged in a line (trituberculine)
(Fig. 23-Sa). Earliest pantotherids (kuehneotherids), represented by the fossil Kuehneotherium
fr~m Early Jurassic of South Wales (Kermack et al., 1968) exhibited primitive type of
tnbosphenic molars with three pointed cusps forming a broad triangle (Fig. 23-8a). However,
the typical trituberculine type of molars were first exhibited by the symmetrodonts (Late Jurassic-
Earl.y .cretaceous) represented by Spalacotherium and dryolestids like Crusafonia. The molar
exhibited development of three prominent cusps arranged at one side of the tooth as a
symmetrical triangle (Fig. 23-8a). In advanced Upper Jurassic pantotherid, appeared more
:mplexity in this type of molar tooth. The upper molar developed three cusps, one larger
.drotocone) on inner apex of the t~iangle and two smaller cusps and two cuspules on the outer
si
in ue named. as p~acone, metacone, paraconu.le an d metac?nu le. Th ese ~usps of molar teeth
.d Pper Jaw formed a series of triangles or tngons each with an apex pointing towards inner
1
: e. ~e lower molar similarly developed a three-cusped triangular pattern caJled trigonid at
m:~r side. o! the tooth with a low heel called talonid for the fitting of larger protocone of upper
and ~•s 1s called trituberculosectorial or tribosphenic or trilobosphenic type of molar (Bown
se, 1979). The trigonid cusps are named as protoconid, metaconul and paraconul while
,.
I
J 14 PALAEONTOLOGY
.
u~-3 g~
~
u
u o-~
Kuchneothcrid
L ~--·O-~
L
Triconodont
u \I)
u
L~
u Symmctrodont {tribosphcnic)
Multituberculata
Pantotherid (tribosphenic)
(a) UPPER SURFACE OF MOLAR TOOTH OF SOME MESOZOIC MAMMALS
L =Lower molar U =Uppe'r molar
Upper Molar Upper

Molar
Trigon
Trigonid TaJonid Trigon,d Talon1ll

Part - ~ Part
Entoconid
a.-;i111-- J-lypoconulid
Lower Molar Lower
~~~~~,-.'---Hypoconid Mohar
Prot.ocone - - - - Mesoconid
Top View Of Tooth Side View
(b} TYPICAL TRIBOSPHENIC MOLAR TOOTH (basic molar stmcturc) OF PLACENTAL MAMMAL
FIG. 23. 8: PAITERN OF MOLAR TOOTH OF EARLY MAMMALS
GEOLOGICAL 11/STORl' OF \ll•.' R1'b.'IJIU'Jlli:s , I~
talonid basin have thr e to fom sm 111 ·r cusps th slK11n1t~d hy cmlo<.·maicl, l1,y11m;m1itJ, mc,Yoc:miid,
and Jiypocomdid (Fig. 23 8b). This i~ ·onsid 'I' d 11s lh tns · n1olm-st1u ·1111· • of th· n1111t1tt1al
from which grndunlly hnv' cl v hp d nil oth r ·oiupli ·ut~tJ typ~s found i11 udvu11 · ·d rou1rn
by addition of now cusps, con · s, connul s nnd ridgt•,s ~lophs , t11T1111g d i11 vnriuhl cotnpli ··at ··I
pattems. Multituberculate mnmmnls (Lutt) .h11·11ssk to 1,nt • Uo · · 11 ) ·xh1blt ·d molurs with parallel
rows of cusps arrang"d in two s •ri s hrondly similnr to thmH of pn·N ·nt duy rc:>d •nts (Fig.
23-Sa).
The surviving mnmmals may b includ •d hronclly Into thn ·• groups, th· 1nonotrcmcs, the
marsupials and the plocentuls.
Monotremes ar~ specialized but hnsi nlly pl'imltiv · m1unmuls. now survivin in Aui,tralia.
They possibly aroso from docodont an stry in l .. ot Jurnssi .. _ r tuc"ous tim N, Munrnpiuls,
another group of primitive mnmmals npp nr din pp r r ·toe ous in North A1nericu. while the
first placental mammals, insectivorn nlso mnd th ir d- but, 1n Upp·,,r r~tn<.:"OUS and their fossils
are found in Mongolia and North Am •ricn. Both th so fonns wcr considered derived from
pantotherids. As it is almost impossibl to d scrib the history of nit th ·sc numer us groups of
placental mammals within this limited spnc •, th~ nuthor will try t'Cl touch only some key points.
lnsectivoras may be discussed in purticulnr for th y possibly occupy u central point of ull other
subsequent mammalian radiations (Kielnn-foworowsku, 1979) (Fig. 2 -7). nc of the generalized
Cretaceous insectivoru is Zalambdalestes of Mongolia, which hud n primitive skull, tribosphcnic
molars and a ring-shaped tympnnic bon~ (n chnruct·er of "uth(irids). From such a gencruliscd group,
eutherian radiation took place into vnrinblt, directions in Pnlnt,oc •1,c-Eoc ne times. The different
groups of mammals radiated out are cd ntutes, chiro1 t~rus, dcrmoptcrns, taeniodonts, • tillod nt.s,
primates, rodents, lugomorphs (rabbits), nrchuic cnrnivorcs (crcodonts)• uncl urchiac ungulate
condylarths*. The last two are so close us to indicut.c an immcdiute common uncestor and they
subsequently became the ancestors of most of the oth •r r,roups of n-rnmmuls with two distinct
food habits (carnivorns from creodonts und hcrbivoro hoof d mnmmuls from candylarths).
Bats or chiropteras are flying mummnls whose front lirnbs nrc modifcd to form wings. The
limb bones are elongated ns arc all the fingers oxCC[>l the thumb supporting the membrane of the
wings. Hind limbs are weak but have clnwed digits by which th •y hung themselves upside down
when they rest. The history of these mnmmnls is innd qunt ly kn )Wn due to luck of fossil. Another
descendant of insectivores arc colugos or flyins I •murs ((formopt •rids) which are tree~living
squirrel-like animals. Edentutes urc anothc1· primitiv • d .. rivntives from insoctivorns huving highly
simplified or subpressed teeth adapted to v •ry sp ·cinlize<l clicts. However, t.hey evolved in South
America, remained isolated from the rest of th world und m· still so upt'l"l th present day. Initially,
they evolved in two lines. One group includ s th· ground sloths (now ~xtinct), the tree sloths
and the ant-eaters. The other includ ·s unnndillos uml lyptcukmts. This s • ·ond group of cdentatcs
have developed excessive nrmour of h"nvy bony pint s for th ir protection. round slot.hs
developed sometimes to giguntic size. M<1Ratl1t1rium wns ns ll\l'g • ns n smnll el phunt. Another
small group of unteuters urc pungolins (pholidontid) found 111 Asin und AMcu, possibly evolved
·n,
from the primitive ed "'ntnte pnlnconodonts. ~nlodonts nnd tlllodonts ua·c nlso two smnll extinct
groups of mamm•ds of Pulococene-f!oc •ni., d riv d fr )Ill ins ct.lvorns with n f •w fossil records.
. . -
extinct grou,,s
PALAEONTOLOGY
~16
Skull
Skull
Hyracotherium
Palaeodonta
(Stem-perissodactyl of Lower Eocene)
(stem-artiodactyl of lower Eocene)
Skull
Condlylarth
(Upper Cretaceous-Palaeocene-Eocene)
(Stem ungulata and ancestor to all other ungulates)
lnsectivora
(Up Cretaceous-Recent)
(Ancestor to other euthcrids)
CrcodontJ
(Ancestor of other carnivores)
(Up Crctnccous-Plio-Pleistoccnc)
FIG. 23 • 9: SOME ANCESTRAL GROUPS WITHIN MAMMALS
317
Creodonts (Fig. 23-9), appeared in Late Cretaceous from · t' ·
. d . msec 1vore-ancestry an North
Amenca, Europe an Asta. The Upper ~retaceous Deltatheridium may be a transitional form
between them. Creodonts had sm~ll bram-case with tribosphenic molars provided with cutting
blades and a large and sharp _canme. They started with two groups. Palaeocene oxyaenids led
to hyenodonts of Eocene-Oligocene and the other the Eocene-Oligocene miacids were the
probable ~ncest~rs of all the modern carnivores. There are two groups of modern carnivores :
fissipeds, 1~cl~dmg mo~t of_ the land carnivores represented by canids (dog-like) and felids (cat-
like) and pmmpeds which, include aquatic groups like sea lions, walruses, 'seals etc. Although,
the last group appeared in Eocene times but they have no fossil record prior to Miocene. They
are adapted for aquatic life and their other morphologic structures are accordingly modified
such as, a streamlined body, paddle-like limbs but no propulsive tail. Teeth are also modified
for catching fishes and become conical and pointed. Some common fossil carnivore genera are
Cynodictis (Eocene), Hesperocyon (Oligocene), Hemicyon (Miocene), Amphicyon (Pliocene)
Ursus (Miocene), Indractos (Pliocene), Smilodon (Pliocene) etc.
The earliest most rodent is Paramys reported from Palaeocene-Eocene rocks of North
America. It was like a large squirrel with clawed feet for grasping and climbing and a long
tail for balancing. Incisors were large chisel-shaped and their cheek-teeth were low crowned
with blunt cusps. From this generalised fonn, rodents diverged along numerous trends of adaptive
radiation during Cenozoic times giving rise to such fonns like squirrels, rats, beavers, porcupines,
etc.
Rabbits are formerly considered as a suborder of rodents but at present are placed in a
separate order lagomorpha. This is because the very primitive fossils of rabbits and rodents
both found in Palaeocene-Eocene show that they are two separate distinct groups of mammals
and their resemblance is more superficial than basic. The cheek-teeth of rabbits are taJI prism
with transverse ridges. There are also differences in their post-cranial skeleton. A rabbit is
specialised for hopping by its larger and stronger hind limbs while its tail is highly suppressed.
One of the earliest and generalized forms of rabbits, is Eurymylus collected from Palaeocene
rocks of Mongolia. From very Early Tertiary, lagomorph exhibited two separate groups, one
represented by pikas and the other by rabbits and hares.
Cetaceans that include whales and porpoises are highly specialized, and most unlike the other
mammals. Thus it appears that they were separated from the basic eutherian ancestor at a very
early date and they have passed through a seri_es of extra-ordin~rily rapid evolutionary changes
making them completely adapted for the li~e m water almost hke a fish . The r:t~rn of whales
to water may be a fine example of evolut1?na~y convergence and wha_les exh1b1t many such
characters shown by the aquatic extinct repttl: 1chthyosaur~ ~nd fishes hke the modern skarks.
They have a typical fish-like body with no hair and no vanatlo~ of ~eeth. They have nu~ero~s
vertebrae, a dorsal fin, paddle-like forelim~s ~ut suppressed hmd hmbs and ~ propulsive tail
(unlike fish, extends horizontally). Their fossils are rather ab~uptly present. m Eocene rocks
although, they were deviated from the ancestral group much earlier. Some fossils are th~ Eocene
Protocetus and Miocene Prosqualodon. The modern forms are represented by doJphms. blue
whales, sperm whales etc.
The ancestor of all the hoofed mammals and some other related forms is_ t~e c~n~ylarr~
group which appeared in Early Palaeocene and exhibited evidences of early d1tlerent1armn ot
318 PALA EONTOt.ocy
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herbivoras from carnivore mammals. Condylarths (Fig. 23-9) were small creatures with 44 teeth
3143
, ") Th ey ha d tn·bosphemc
· molars with
· blunt .cusps, short legs, a long tail and feet
( O.F. 3143 x - ·
with flat nails. From th~ very beginning they showed a tendency to become larger in size and
to develop more cusps m molar tooth. Phenacodus is one of the most primitive condylarths of
Eocene of North America.
A larg~ number of ungulates were radiated out from condylarth-ancestry during Palaeocene-
Eocene times (Pr?thero et al., 1989). These are amblypods,* xenungulates,* pyrotherids,*
notoungulates, * htopterus, * astrapotherids, * tubulidontids, perissodactyls, artiodactyls and
proboscideans. Evolution of these groups of herbivore ungulates took place almost independently
and sometimes they remained confined to one particular continent. Many of them became extinct
at the end of Pleistocene and some surviving groups are represented at present only by one or
two genera.
Some earliest (Palaeocene-Eocene) large ungulates are Pantolambda (Palaeocene),
Uintatherium (Eocene) and Coryphodon (Eocene), all belonging to the group amblypods.
Tubulidentid are first reported from Miocene rocks and at present they are represented in Africa
by a single genus Orycteropus popularly known as aardvark. All South American ungulates such
as xenungulates, pyrotherids, notoungulates, astrapotherids, litopteru$ mostly appeared in Palaeo-
Eocene, attained climax in Mio-Pliocene but became extinct by Pleistocene. From an ancestral
form approximated by Hyracotherium (Fig. 23-9), perissodactyls evolved along various paths
of adaptive radiation reaching at climax in Mid-Tertiary times after which they have declined
in number. At present, they are represented by three genera Equus, Rhinoceros and Tapirus.
From the very beginning perissodactyls evolved along three independent lines. One of them
includes equids; the second line leads to rhinoceratids and tapirs and the third includes the extinct
chalicotheres and brontotheres. The last group often attained a g_igantic size. One such form
was Oligocene genus Brontotherium showing approximately eightJeet height. Miocene forms
like Moropus of North America and Macrotherium of Euresia were as large as modern horse.
Most of them became extinct in Pleistocene.
Evolutionary development of rhinocer~tid~ too~ plac~ in Af~ca. They ~ppeared in Middle-
Upper Eocene and attained their culmination m Ohgo-M1ocene times. Earher forms were .small
slender-limbed animals. The animals within this group show a general increase of size,
molarization of premolars, elongation of cheek teeth with continuation of pattern of strong crest
(established in the primitive rhinoceratid) and ~nally d~velopm~nt of horns and three toe~ feet.
Baluchitherium was one of the largest extinct rhmocerat1ds of M10cene age often became sixteen
to eighteen feet in height. This was a hor~less animal. The modern form Rhinoceros appeared
in Pliocene and exhibits either two or a single horn.
Tapirs in South America and Malaya first appeared in Eocene. Modern form Tapirus exhibits
many primitive characters compared to other perissodactrls, such as greate_r number of toe~ (~).
possession of all fortyfour teeth in jaws etc. The evolution of horse .remamed confined w1thm
North America at least during Eocene-Miocene time~. They_ evolved m Lower Eocene fro~ the
earliest form Hyracotherium which was a dog-like ammal with fortyfour teeth and 5-toed animal
* Extinct
320 PALAEONTOLOcy
.k their ancestor condylarth. With progress of evolution equids exhibited gradual increase f
l ie .. f I I . o
· decrease of number of digit in feet (5 to I), molanzat1on o premo ars, e ongat1on of cheek
size. d · N
teeth, and more complexity of brain. The modern ge~us Equus a~peare m orth America in
Upper Pliocene. But all American equi~s became ~x!mct after Pleistocene. ~hose forms which
were able to migrate to Euresia and Africa are surviving at present, all of which are represented
by a single genus Equus (for details see chapter 25).
Artiodactyls, the even-toed mammals (4 or 2 toes), starting almost at the same time as did
by perissodactyls, have emerged in the present earth as the most successful among the ungulates.
At present, the artiodactyls are represented by a diverse group of animals such as pigs, precarries,
mppopotamuses, tragulids, deers, giraffs, sheeps, goats, antelopes, catties and so on. Inspite of
their wide radiation they exhibit some common characters in their skeleton and soft-part anatomy.
Primitive forms had a full set of fortyfour teeth but advanced forms show reduction and/or loss
of upper incisors and lower canines. The cheek teeth become bunoselenodont to selenodont with
high crown showing cresentic cusps. But the most characteristic feature of this group is their
possession of a four-chambered stomach which gives the animal a complex mechanism of
digestion of food and this probably gives the animal more success and advantage over the other
ungulates.
The first artiodactyls appeared in Lower Oligocene from some archaic condylarths having
even-toed limbs and omnivorous food habit (astragulus). Diacodexis, a Lower Eocene palaeodont
of North America, Asia and Europe is considered as the earliest-most form (Fig. 23-9) (Rose,
I 982). These were non-ruminant plant-eaters. Some of the subsequent groups of artiodactyles
such as entelodonts, oreodonts, dichobunoids, attained quite giant size but they mostly became
extinct by Mio-Pliocene. From them, radiated different groups of surviving artiodactyls like
non-ruminants suids (pigs) hippopotamuses and tylopods (camels and Lamas) and ruminants
like giraffids, bovids (antelopes and catties) tragulids etc. Some of the common fossil forms of
different artiodactyls are as follows :
Hippopotamids : Merycopotamus (Pleistocene), Hydaspitherium (Plio-Pleistocene).
Anthracotherids : Anthracotherium (Plio-Pleistocene).
Tylopods : Protylopus (Eocene), Protomeryx (Oligocene) Syndyoceras (Pliocene),
Dicroceras (Miocene), Megaloceras (Pleistocene).
Tragulids : Archaeomeryx (Eocene), Eumeryx (Oligocene), Palaeomeryx (Miocene).
Giraffids : Pa/aeotragus (Mio-Pliocene), Giraffakeryx (Pliocene), Sivatherium
(Pleistocene).
Bovids : Merycodus (Miocene).
Most of the recent artiodactyls appeared in Plio-Pleistocene times. These are : Sus, Tragu/us,
Cervus, Hippopotamus, Gira.ffa, Capra, Bucapra, Bos, Bison, Came/us etc.
~ot_her group of ungulates ~re proboscideans which include elephants and their allies such
as saremans (sea-cows), hyr_ac?1ds, and the extinct groups like desmostylia and embrithopods.
They took a very early devaataon from ancestral form, the condylarth.
Sea-cows are adapted to aquatic marine life and were deviated out from other ancestral
proboscids at a very early dat.e of geological history (Eocene). They have their teeth character
GEOLOGICAL HISTORY OF VERTEBRATES '.\2 1
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FIG. 23 - 11 A SIMPLE PHYLOGENETIC TREE OF VERTEBRATES SHOWING EVOLUTION OF

DIFFERENT SURVIVING GROUPS
PALAEONTOLOGY
3~~:r=:::::::::::~~:::::::::::::::================~~~~
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FIO. 23 - 12 : GEOLOGICAL RANGE AND RELATIVE ABUNDANCE OF MAJOR
GROUPS OF VERTEBRATES
323
similar to pr~bos~ideans. They ~ossess a blunt head (unlike fish}, flipper like forelimbs and
suppressed hmd l_imbs. Two fossil forms are Protosiren (Eocene) and Halitherium (Miocene).
The recent form 1s dugong.
Elephants are a unique group of animals possessing a mobile trunk or probosis, a pair of
rusks and a huge body. They are still living in Africa and Asia but represented only by two
genera. The African form is Loxodonta and the Asian form is Elephas. The earliest elephants
are represented by Moeritherium of Late Eocene of Africa, from which diverged four different
groups, each showing independent line of development. These are dinotheres, trilophodontids,
mastodonts and elephantids. All of them except the last one got extinct by Pleistocene. Modern
elephants of Africa and Asia are derivatives from elephantids in Pleistocene (for details see
chapter 25).
The last but the most important group of mammals are the primates to which we belong.
Primates are, by no means, structurally an advanced group. In fact, they retain many primitive
characters inherited from their insectivore-ancestor. Tupaia is a modern form, morphologically
lying close to the demarcation line between insectivore and primate. Primates can be called a
primitive but highly specialised group of mammals. They have comparatively large brains,
bionocular vision of eyes and pre-hensile limbs with brachiating digit and a tail. All these
features have proved exceptionally suitable for an arboreal life ventured by them. They have
primitive jaws with all types of teeth suitable for variable types of food. Man exhibits further
specialization by developing an erect bipedal locomotion. This enable him to use their hands
for preparing tools, that ultimately has made him supreme ruler of the present earth.
Primates are represented by two groups : prosimians or lower monkeys and simians or higher
monkeys. The former are more primitive creatures and mostly became extinct except a few
surviving forms such as lemurs, lorises, tarsiers etc. Higher group of monkeys developed in
two separate continental areas. The new world monkeys, appeared in America in Eocene times
(often called ceboids), have a very poor fossil record. The old world monkey, also called
anthropoids, appeared in Africa and Asia in Upper Eocene-Oligocene times (slightly younger
than the former), include three different types : monkeys (cercopthecids), pongids (apes), and
hominids (man). Parapithecus of Egypt is one of the earliest form of this group of Early
Oligocene age from which possibly diverged cercopithecids and hominoids (ape and man) along
. two different lines. Cercopithecids are represented by such forms as Mesopitheccus,
Palaeopithecus, Macacus etc. The recent survi\'ing forms are African baboons, Indian monkeys
etc. Propliopithecus is probably the forms from which ape and man began to diverge in Lower
Miocene. Some Mio-Pliocene apes are Dryopithecus, Sivapithecus, and Proconsul. Dryopithecus
were possibly the ancestor of most of the surviving apes of Africa like chimpanzee and gorilla.
The actual point of divergence between hominids (man) and pongids (ape) remains confused
for lack of fossils. The first true hominid form was Australopithecus of Lower Pleistocene of
African which led to Homo habilis and Homo erectus in Middle Pleistocene. Modem man or
Homo sapiens probably appeared about 200,000 year ago from some H. erectus lineage (for
details see chapter 26).
A diagrammatic sketch of phylogeny of major groups of vertebrates and geological range
of major groups of fishes, amphibias, reptiles and mammals are shown in fig. 23-11 and
fig. 23-12 respectively. The relative abundance of major groups of mammals and probable
phylogenetic relation among them are shown in fig. 23-10.
Chapter 24
DINOSAURS
24.1 INTRODUCTION . . , . , J. JJ
Dinosaurs are a group of extinct land reptiles of Mesozoic. The name d10nosaur, 1tera Y_
meaning 'terrible lizard', was given by Richard Owen in 1841. It is rather an unfortunate name
for these animals which were mostly slow-moving plant eaters. Only one or two groups of
dinosaurs were flesh eaters but whether they were active hunters or simple scavangers remains
still unknown. Thus it seems that there was nothing terrible about these animals except their
gigantic size as exhibited by most of the members. The animal appeared towards the end of
Triassic from thecodont reptiles, attained their climax in Jurassic-Cretaceous but suddenly
became extinct at the end of Mesozoic along with many other groups of reptiles of Jand and
sea. They were the supreme rulers of the earth throughout the Mesozoic for about 170 m.y.
Their fossils are mainly represented by skulls, vertebral bones, limb bones, foot prints, coprolites,
gastroliths and fossil eggs. Fossils comprising of almost complete but disarticulated skeletal
materials have been collected from several sites of the world which have been reconstructed in
many cases in laboratory and museums to get the complete picture of the skeletal structure of
these anim~s. Their modes of locomotio~ hav~ been interpreted from their foot-posture, foot
bones,. pelvic structure and also from their fossil foot-prints. The food habits of there animals
are known from their teeth-characters, movement pattern and also from the analys ·s f f te
& ·1 Th · f · h b" . 1 o copro 1
,oss1 s. e1r nature o m a 1tants 1s known from the rock types of the fossils sites.
24.2 CLASSIFICATION
S D~nos~urs, within. th: su~class Diap~ida of ~eptile group, are divided into two orders viz :
aur1SChia and Ormthisch1a. The basis of this subdivison is th · . . . .
24-1 ). The former had a triradiate type of pelvic joint like most o::~ nature of ~1p-.,om~ (Fig.
latter was characterised by a bird-like quadruradiat type of . . _e other reptiJes whiJe the
pe1VIC JOint.
Order : Saurischia (Fig. 24-2a) : Mostly allied to th
made up of ten cervicals and thirteen t k e ancestr~ thecodonts; back-bone
quadrupedal; pelvic typical to that of a~·n ~~rtebra~; ta1I long; both bipedal and
digits with curved claws. izar • teeth m front portion of jaws; five
Section : Theropoda : Advanced biped· earn·
, 1vorous.
(C)* etc.
~= :r
Families : (a) Coelurosaurfa : Relatively s II
tail, skull very small; Examples . for~s (8 to I Oft); long flexible neck and
· e ophys,s (T-J)*, Coelurus (J)*, Omithomimus
(b) Carnosauria : Relatively larg . d" ·
large, thick and massive acti e in amension, skull large, neck very small· tail
ng as an effective s ~ •
• T : Triassic, J : Jurassic c . c tac . . upport ior these animals·' mostly
• · re eous.
324
·~·
DINOSAURS 325
Ilium
Acetabl1hun - .......- - -
Pubic Process
(a) SAURISCHIA
FlG 24 - 1 . ST (b) ORNITHISCHIA
. . RUCTURE OF PELVIS OF DINOSAURS
Coclurosauria (Coeloplrysis,
length: 8- \ 0 feet)
Camosauria (Tyranosaurus, length 40-45 feet)
(a) SAURISCHIAN DINOSAURS
Stcgosauria (S~go.umrus, length 20 feet) ,
Omithopoda (lq"anodon) (length 20-2~ feet)

....
Ceratopsia (Tric•ratops, length 40-SO feet)
(b) ORRITHISCHIAN DINOSAURS
FIG . 24 - 2 : DIFFERENT GROUPS OF DINOSAURS (not to scale)
326 PALAEONTOLOGY
<..:11,,f1111hysi.1·
(An early Triassic Saurischian)
Fibm.w urees
(An early ornithosuchian
of Upper Triassic)
Euparlt.ula
( A lower Triassic lhccodont und
possible nncestor of dinosnurs)
FlG. 24 - 3 SOME EARLIER FORMS OF DINOSAURS AND THEIR PROBABLE ANCESTOR
-
DINOSAURS 327
bipedal, forelimbs very small, hindlimb much larger; carnivorous with numerous
sharp and effective teeth. Examples : Allosaurus (J-C), Tyranosaurus (C) also.
Section : Sauropodomorpha : Mostly larger in dimension; quadrupedal and herbivorous;
forelimbs and hindlimbs well developed.
Families : (a) Prosauropoda : Nearly 20ft; both the limbs effective; herbivorous. Example:
Plateosaurus (T).
(b) Sauropoda : Mostly gigantic in size, quadrupedal, herbivorous; with a long
tail and neck; skull absurdly small. Examples : · Cetiosaurus (J), Brontosaurus
(J-C), Diplodocus (J-C) Brachiosaurus (J), Gigantosaurus (C) etc.
Order : Ornithischia (Fig. 24-2b.) : Bird-like hip-joint, pelvic tetraradiate type; predentary
bones feebly developed, with or without teeth often restricted to the hinder part
of jaws; herbivorous.
Families : (a) Ornithopoda : Bipedal or quadrupedal; skull long and low; beak-like sheath
in front of mouth (duck-billed). Example : Camptosaurus (J-C).
(b) Stegosauria : Armoured dinosaurs. Examples : Stegosaurus (J-C),
Ankylosaurus (C) etc.
(c) Ceratopsia : Armoured and homed dinosaurs; quadrupedal; thick limbs and
a short tail. Examples : Protoceratops (C), Triceratops (C) etc.
24.3 ANCESTRY
It is generally believed that dinosaurs evolved from some thecodont reptiles in earlier part
of Late Triassic times. It is generally suggested tbat· all the different groups of dinosaurs
descended from a common bipedal thecodont similar to Euparkeria. (Fig. 24-3). The earliest
members of the dinosaurs were the saurischians represented by the group coelurosaurs of Upper
Triassic age. Euparkeria of Lower Triassic of Africa was a small form with lizard-like hip-
joint and a relatively smaller forelimbs. Possibly it could be able to run on their hindlimbs. So
the appearance of the first group of bipedal saurichians in Upper Triassic times from such a
thecodont ancestor is almost certain. The three oldest dinosaurs are Eoraptor and Herrera~ai,rus
from Argentina and Coelophysis from North America. It has been assumed that these bipedal
saurischian theropods, evolved from thecodont, were the ultimate anc·e stors of all other
sau~schians and ornithischians. Quadrapedal dinosaurs of both the groups appeared at a later
per~~: Considering dinosaurs as a monophyletic group was essentially based on the belief that
omitthschians were geologically younger than saurischians, and evolved in Jurassic times, as
f~ssils of these dinosaurs were unknown from any Triassic rock for many days. However,
discovery of fossils of some dinosaurs similar to ornithischians from Up~r Triassic rocks has
f(oLrced the scientists to think differently. These primitive ornithischian fossils are Pisa11osaurus
ate T · · . · .
. nass,c of Argentina) Heterodontosaurus (Upper Tnassic) and Fabrosaurus (Early
1
urasts,c, of ~outhern Africa) which were all small bipedal dinosa~rs with bird-like hip-joints.
No on y this mo h l . II h . . to the Triassic
. . thecodont Hesperosuchus
d . • rp o ogica y ,t ey are very similar
O
: b' ~~t~suchus which were also small bipedal thecodonts showing pelvic joint tending to
ir i e. Thus a thecodont could also be the possible ancestor of ornithischian group. In
PaJae(Gl.!o)WP-42
PALAEONTOLOGY
fact. the nnimals belongin~ to th• two ord rs of dinosaurs were so different in their skuJI, jaw
and pelvic strn ·tun' thut it is difficult to ace pt their origin from a common ancestral stock. It
is rather lo~ical to think. that th se two groups of dinosaurs evolved almost at the same time
Upper Tri t1ssic ind •pend ntly fro m the two differe nt thecodont-stocks and in that case
dinos~mrs ma) he a pol) phyl tic group . Howe ver, rece nt cladistic analysis have indicated
dinosaur a monoph) leti" group, exhibiting wide morphologic and ecologic variations.
24.4 SOME IMPORTANT SITES OF DINOSAUR FOSSILS

Dinosaur fossils have been collected widely across the world. A few important fossil sites
are :
Great Britain Lyme Re gis. Dorsal and Oxfordshire : Fossils include lguanodon,
Megalosaums etc.
U.S.A. : Utah, Wyoming, Colorado and Montana : Fossils include Allosaurus,
T ra1rosa11ms , Stegoceras, Hadrosaurus, Triceratops etc.
India : Jabalpur-Pisdura, Godavari valley and Trichinopoly : Fossils include
Titanosaurus, Laplatosaurus, Barapasaurus, Stegosaurus etc.
Monogolia and East Africa : Tarbosaurus, Brachiosaurus, Protoceratops etc.
24.S GENERAL CHARACTERS, LIFEWAYS AND ADAPTATIONS

Problems of 'big life' : Size of ~inosaurs was quite variable ranging between Jess than one
foot · h. as 90 feet, measunng from head to tail. Most of the sauropod d.mosaurs are
· to· as long
g1gan~1c we1g mg near1y 50 tons. As an animal grows larger, its surface area through which
heat 1s lost and rabsorbed becomes proportionately much smalle r. Fl uctuat1ons · o f bo d y
temperature h
be d t ercaorc
d · occur more slowly.. Thus small variation of temperature, for instance
.
tween ay an mght. would have had httle effect on these hulk· be
of cold or hot weather might have affected such animals. mg ast. Only a long spell
Saurapods exhibited a lizard-like posture and they had massive ii · . .

the huge weight of the middle part of body L. b bo P lar-hke hmbs supporting
. im s nes were verticall d d . d
together with tough ligaments that gave them enormou Y arrange , an tie
differently where less materials were used. There s strength. Upper part was designed
skeleton wit.h hollow bones. Thus they were bottowa_sh a small. head (sk~II) and light axial
adaptation for the ·big life' . Some sauropods had hemd eavy 8!' mals, which was a suitable
1
genera it was not bigger than a kitten's. Vertebra a f even smaller than a horse's. In some
0
pleurococls) on their sides. This involved m.inimum: t:ropods had large cavities (called
the animal's overall weight. Walking for a big anim:; ~a nes and .therefore helped to reduce
a flat feet to support the enormous body. But walk. Y be a ~aJor problem. Elephant has
whi~~ is not too bad for. t.he elephant is provided ~~~ oan fl~t f~et is rather an exhausting work,
pos1taoned at the back sade of foot and that kee , h built-in he~I made up of tough tissues
:d
great sauropod dinosaurs probably had u simri:rt eleph.~t on its toes while walking. The
supported behind by a thick wedge of tissues fu . ~ptat!on. Toes of limbs were possibly
allowed in the quadrupedal locomotion to plod r nctionmg .hke a heel and this specialized feet
. ff rt
C o .
,orwurd using n,·mu~um· amount of energy and
DINOSAURS 329
Other features : Carnivorous dinosaurs like tyranosaurids (largest among the group) had a
bird-like bipedal posture. It had a body of about 45 feet length and standing as tall as a double-
storey building. It had a comparatively large skull more than 3 feet with formidable-looking
teeth, each had about 6 inches length and serrated edge. The back legs were muscular, ending
in three toes. They had also a long and powerful tail acting as a counter-balance while walking
upright but had surprisingly small front legs. The latter were probably used by the animals for
getting up from rest condition (lying on stomach).
Besides the two general patterns mentioned above, many dinosaurs developed several other
additional features. Stegosaurs, for example, were large-sized slow quadrupedals having armour
plates running from neck to the tail all along the back being largest at the middle (about 3 feet
long). The tail end was provided with several sharp spines. Duck-billed ·dinosaurs were probably
grazers having a projected mouth-front in the form of a beak of a bird. Many duck-billed forms
and ceratopsians bore a variety of head-crests. For many days it was thought that these dinosaurs
were aquatic and the head-crests worked as aqualungs or snorkels. But the present idea is that
these helmets acted as identification tags. They were some kind of head-badges or signals
helping male and female of the same species to recognize each other. Ceratopsians had strange
horns and huge backward-pointing neck frills. This gave them sufficient protection from both
sides. This might have also helped the animals keeping the body cool by acting as heat and
light-deflector. Ankylosaursus was built like a small army tank covered from head to tail by
bony plates. Pachycephalosaurus carried a solid mass of bone over IO inches thick on its head
and the area was surrounded by a number of bony studs.
Life ways : In general, dinosaurs, whether quadrupedal or bipedal were slow-moving animals.
If we assume that its body activity was similar to that of a modern crocodile, then, after walking
over a long distance at considerable speed, its bulky body would be in danger of over-heating.
Saurischians like the giant sauropods, and ornithischians like the stegosaurs and ceratopsians
were all quadrupedals, while carnosaurs and omithopods were bipedals. Carnosaurs mostly
possessed very small forelimbs compared to their hind limbs, which might have been used in
other purpose except locomotion. But ornithopods possessed comparatively larger forelimbs and
it is suggested that many of them could have walked on four legs as and when necessary.
Coelurosaurs, often called ostrich dinosaurs for their size like modem ostrich, had a Jong
tail, larger hindlimbs and a pair of small forelimbs. As they were smaller in size they could
run on their long nimble legs and grasped their preys with their flexible hands provided with
clawed fingers. Some of them like Ornithomimus probably survived on a diet of insects, lizards
and eggs of other dinosaurs. Coelurosaurs were the fastest dinosaurs which could move at a
speed of about 55 kph. Large head and tooth character suggest that most of the camosaurs
were undoubtedly flesh-eaters but opinion is divided over the question whether they were active
predator-hunters that caught their prey by sprinting after it or they were merely scavengers
feeding on dead bodies of other animals. The shape of the skeleton and a long powerful tail
counter balancing the trunk, suggest that they were fast running animals over short-distance.
They .lurked in ambush and then pounched. The skull, made up of heavily reinforced bones
su.ggests that it had to withstand an enormous impact when it ran into its prey with its jaw
wi~e open. Some experts. however, believe that such a large carnivore was too big to move
qulckly for catching comparatively smaller sized preys. They also argued that their sharp and
330 PAI.AEONTOLocy
Bonehead
Cresl
Horns
FIG. 24-4: _DIFFERENT TYPES OF CRESTAL STRUCTURES OF HEAD IN DINOSAURS
.Triceratops
!!l
v.,
::> Trachodon =
·;;;
C.
Tyra11osaur11s
0 0
w 1i1rbosa11rus
u ...
«i
u
<
I- :..)
w
a: Ornithomimus
u Protoceratops
lg11anodo11 .
~
Allosaurus
:I
Sre~oceras a Mega/osa11r11s -~
Camptosaurttl 0 ::i
~
$ i
~
u Ornitho/estes g"'
~
o" .:::
4 J
•
Plateosaurus
•Heterodontosuurus
,I
------- _ _._ - ----- ..
Ornithosuchids ~
~
~~
..
'
"'-...._ Euparkcriuns
FIG. 24 - 5 : PHYLOGENY OF DINOSAURS
DINOSAURS
·&
km1e l"k h 331
t e teet would have been shattered if the . . .
large dinosaur. More probably they d h . Y wer~ driven m force mto the body of another
lump of flesh from the b d 'f d use t e1r teeth hke a set of steak knives to slice off large
scavengers. o y o a ead animal or in other words these animals were essentially
It is almost certain that most 0 f th d" .

t d . e mosaurs hke prosauropods sauropods ornithopods
~eg~s:~;s da~. ceratopsians were vegetarian, feeding on soft plant, 'with their ;eg-like teeth,
uc - 11 e mosaurs were perfect grazers with their numerous small teeth (as many as t 000).
on the back-half of .the J. aws and th e f ront part of Jaws
· ·
were projected out to form the beak.
These. teeth
. were bemg .constantly l d D · f · ·
rep ace . ommance o duck-billed dmosaurs in Cretaceous
may t~dicate that battenes of teeth in them were adapted to deal harder shoots of the incoming
~owe~mg plants of that time. Orinthopods like lguanodon were also herbivorous, generally
hved ~n herds. ~ herd of about twenty lguanodon fossils, presumably buried by a gigantic
landshde, was discovered from Belgium.
Sau~opods also developed peg-like teeth on the front half of the jaws, suitable for soft
vegetatt~n and they ate soft plants of marshes and lakes. Although, the creature's legs could
support tt on dry land, a sauropod usually spent a lot of time wading in marshes and lakes.
This would help its legs to bear its heavy body. In addition, as the fierce meat-eating dinosaurs
did not like to encroach the water, sauropods were less their targets. Their long necks allowed
them to stand with their bodies submerged in deep water keeping their head above the water
surface. Their enormous weight was buoyed up by the water .around them. It has been also
suggested that the long neck with the nostril positioned on top of the head was an adaptation
to life in deep water, allowing the animal to breath when standing under water. However, many
workers have argued that submergence of such a huge body (about 12 meter high) in water
would create water pressure so high that the animal's lungs would have collapsed and it would
be unable to breath. Even increase of blood-pressure at such stage could have caused its heart-
failure. So, it seems that they were normal animals and their skull, bone-structures and foot-
posture were more suitable for walking on land. The advantage of long neck was that it enabled
the animals to feed to topmost branches of long gymnosperm and fem group of plants as modem
giraffids do.
Stegosaurus and their allied forms like Ankylosaurus, Polacanthus were all terrestrial slow-
moving dinosaurs and eaters of land plants. This is also true for the ceratopsians, the horn
dinosaurs.
Structures for defence (Fig. 24-4) : As most of the vegetarian dinosaurs ~ere the, main targ~ts
of the carnivorus group, many of them developed severa'l structures_ on theu body as protect_ive
devices. Duck-billed dinosaurs developed bony crests on their skulls; boneheads, like
Pachycephalosaurus had a thick bony mass on their skulls; ceratopsians developed armour or
.· th ssessed
bony plates covering the body and also sometimes large shields. In add1ttoo, ey po
one to many sharp horns on their skulls. Stegosaurus had also thick bony armours on the back
and some long bony spikes on the tail.
Control of body temperature : Birds and mammals are warm-blooded animals and they ~~n
· . . · d d b b king down of food materia s
mamtam their body temperature internally. Heat ts pro uce Y_ rea . . . . b an
and their life activities. This heat in a warm-blooded animal ts retained wathm its body Y
PALAEONTOLOGY
332 . b . 1 st by sweating or panting.. Mo dern rept1·1 es
insulatory cover of fur or feather and it ~ay he o. ·tant body temperature by their behaviour.
· I b t th "Y can achieve t e cons
are cold bto?ded a~1ma s u ·.e . as much solar heat as possible. Later during day they
In the morning their body surface absorb . h ti ocks to avoid overheating. In winter, when
shun the shearing heat and tak~ shelter benea~ d .1e r ·or~ of activity, even consuming food and
sufficient solar heat is not available they avo1 any s
some go for hibernation for the entire cold season. .
• . 1, Thus the animal would take a very long time to
Dinosaurs, except a few, were large amma_s. . . . d' hiding beneath a rock or went
warm u or cool down. It is also difficult to 1magme a mosaur . .
for hi:mation for several months. The question of hibernation does. not ansel iff whe acceptldthbat
· · h · M oic But a great dea o eat wou e
there was no seasonal fluctuation durmg t e entire esoz . · . . .
generated by these animals by their daily activities like hunting, walking or eating and that h_eat
would have to be lost by a prolonged rest. They were unable to loose their bod~ he~t by sweating
as they possessed impervious thick skin like that of modem r~ptil~s. So the question is that whet~er
these animals were warm-blooded or possessed any other device, internal or external, for regulating
their body temperature. Bakker ( 1972, 75) argued that erect gait and high speed of dinosaurs (not
applicable to giant sauropods), long neck of sauropods through which they had to pump blood
upto the brain by a powerful four chambered heart (crocodiles have powerful four chamb.ered
heart but are cold blooded reptiles), bone histology (resembling those of mammals and birds)
and occurrence of fossil dinosaurs in polar region are some evidences indicating dinosaurs as
endothermic animals. But none of these observations may be taken as a conclusive evidence. Most
of the authors have come to the view that dinosaurs had some intermediate condition (neither
reptilian nor mammalian type). They probably achieved thermal constancy (homeothermy) by
making the rate of internal temperature change extremely slow during normal subtropical
condition. Some authors consider that armours, plates, spines, horns and such other skeletal
fe.atures found in many groups of dinosaurs might have played some role in controlling the internal
body temperature. Warm blooded animals, of course, have some advantage, as the animals will
be quite independent of environment. But this was surely disadvantageous for such a huge animal
as in that case the internal temperature regulation would require a huge amount of food. It is
questionable if the large sauropods could be able to eat enough food in a day. The gaint flesh-
eater Tyranosaurus would have needed to eat about a tonne of food everyday to keep a warm-
blood body going. Normally, a highly active warm-blooded animal could catch this amount of
food every day. But if they were cold-blooded they could have survived on much less amount of
food. However, it is a known fact that a cold-blooded animal grow more slowly than a warm-
blooded one. S~ how fast did a dinosaur grow ? American scientists are now studying fossilized
nes~~ o~ some dinosaurs and try t_o get the answer of this question and then it will be possible to
decide m future whether some dinosaurs were warm-blooded or not.
Dinosaur egga : Report of dinosaur egg-fossils from various parts ot· the
h . II f · · worId may m · d. t
1ca e
t at. a o them, from
. small theropods Lo large sauropods· , 1u·,d t·hc•1·r e ggs. m
, a nest-
. i·k1 e structure
w h 1ch was dug in the sund or earth and covered over for ·
. · · . I' . mcu bnt·ion as ~een among many
sur~1~mg repu 1cs 1ke crocodiles. Recent work hus also revealed that d. . . · h h·
exh1bued parental earn Skeletons of . I. . . - mosaurs m1g t ave
that t.he juveniles HUl;ed togeth,er f:v~rn J~vena!es found a_ round such nests definitely indicate
sometime uft~r hutching. One of the sites of fossils of
DINOSAURS 333
dinosaur egg is Egg Mountain, Montana where eggs of the hardrosaur Maiasaurus are found
within Late Cretaceous sediments. Dinosaur eggs and nests are equally common in Mongolia.
India has a special place in the study_of dinosaur eggs. In India the largest Cretaceous nesting
sites of dinosaurs extends over a wide areas from Kutch in the west to Nagpur in the east and
further southward to Adilabad, Andhra Pradesh. From the different localities within this belt,
several hundreds of egg-fossils have been collected so far. A lan~e dinosaurian egg is commonly
known scientifically as Megaloolithus. These are cylindrical, pillar like structures with knobbed
outer surface, probably belonging to large sauropod dinosaurs. Another group of eggs are also
found called Elongatoolithus which have been assigned to carnivore dinosaurs exhibiting ridge-
like features on external surface.

Among the saurischians the first appeared group was carnivora theropods represented by
Late Triassic coelurosaurs. Coelophysis is a small, lightly-built biped with a long neck and tail
and probably were the fast moving dinosaurs. From an ancestor more or less like this, the
t11eropods evolved mainly along two lines : some remained small retaining more or less the
ancestral characters except some increase of size. These are represented by Late Jurassic
Ornitholestes which exhibited a secondary four-footed locomotion. A further increase of size
was marked by the Cretaceous coelurosaur Ornithomimus.
In the second line of theropod evolution, camosaurs appeared for the first time in Jurassic
represented by such forms like gigantic carnivores Allosaurus and Megalosaurus which had large
skull, thick and short neck, very small forelimbs, massive hindlimbs and a thick massive tail.
They were typically bipedals and might be active hunters, killing their preys by their sharp
and large teeth. Tyranosaurus, a gaint of about forty feet came in Cretaceous. The other
Cretaceous carnivores were Tarbosaurus and Gorgosaurus. Most of them persisted upto the end
of Cretaceous.
Another group of saurischias, the prosauropod, could be a descendant of coelurosaurs. They
appeared in Upper Triassic represented by the genus Plateosaurus having considerably large
size compared to the theropod Coelophysis. It had solid hindlimbs and forelimbs, comparatively
larger and perhaps were used in locomotion. They were plant-eater saurischias with small skull
and flattened teeth. The trend foreshadowing by plateosaurs, was culminated in Jurassic-
Cretaceous times with the appearance of gigantic sauropods like Diplodocus, Brontosaurus,
Brachiosaurus, Cetiosaurus, _Apatosaurus, Melanosaurus etc. They were all ranging in size
between 40ft-90ft, having small skull, four massive limbs for locomotion, a flexible narrow
but long neck, and a long thick tail. They were typical slow-moving vegetarian dinosaurs and
they persisted upto the end of Cretaceous.
First omithischians were represented by the Upper Triassic Heterodontosaurus and Early
Jurassic Fabrosaurus belonging to the group orinithopods. But a more common form of
omithopods is represented by Camptosaurus of Upper Jurassic. It was a medium-sized dinosaur,
with well-developed forelimbs that possibly were used frequently in locomotion. especially at
the time of feeding. From this, camptosaurid-like ancestor evolved in three lines. In one line
camptosaurid type of structure continued with increase of size. Such a group was represented
334 PALAEONTOLOGY
in Cretaceous by plant-eater Jguanodon. Its thumbs were enlarged into sharp spikes, might be
used for defence. Another line of adaptation was shown by the dinosaurs with bony-head-crests
represented by Stegoceras and Pachycephalosaurus. Another group of spectacular ornithopods
of Late Cretaceous were the so called duck-billed group having frontal extension of jaws in
the form of beak of a bird. These vegetarian dinosaurs were represented by Trachodon and
Hadrosaurus having a large size with occasional qua,drupedal locomotion. lambeosaurus had
a hatchet shaped crest while Parasaurolophs developed a long tubular crest extending backward
behind the occipital region. Careful observation has shown that premaxillary and nasal bones
of these animals were pulled back over the top of the skull to form these hollow crests. But
about the function of such long nasal passage there are different opinions of the experts. Possibly
these structures had increased the smelling power of the animal or they acted as resonating
chambers increasing the power of voice. Possibly they were adapted for semi-aquatic life.
Another group of ornithischias possibly, evolved from camptosarian ancestry were stegosaurs
or armoured dinosaurs. They also appeared in Jurassic and became extinct at early Cretaceous.
They possessed various types of bony armours and spikes on the body that gave protection to
these slow-moving moderate-sized plant eaters. The common Jurassic forms are Stegosaurus.
Ankylosaurus etc. Stegosaurus developed an enlargement of the spinal nerve cord near the
sacrum which was previously thought as a 'second brain' of the animal.
The last group of ornithischians were the horned ceratopsians appearing in Late Cretaceous
times. Their relation to other ornithischians remains confused. They were moderately large-
sized, Rhinoceros-like animals, having bony plates covering the body and two large bony neck-
frills. They also showed development of one or more horns on nasal part of the skull. They
were geographically limited forms found only in North America and Northea.s t Asia. Some of
the common forms were Protoceratops, Monoclonius, Triceratops, etc. The former was of the
earliest hornless form. Monoclonius was a single-horned and Triceratops, a tri-homed form.
Protoceratops is famous because it was the first dinosaur to be discovered in Mongolia together
with its eggs.
24.7 CAUSES OF EXTINCTION

It is very difficult to giv~ a satisfactory explanation of the sudden extinction of dinosaurs at
the end of Cretaceous when they were practically at their culmination. Various hypotheses have
been put forward by various workers in different times to explain their sudden death but from
critical evaluation of these hypotheses it appears that none of the reasons could be taken as the
sole responsible for this unusual event. It has to be remembered that the end of Cretaecous
wa~ marked not only by the extinction of dinosaurs but various other organic groups were
ser~ously affected also .. Abou.t 90~ of the Mesozoic reptiles both terrestrial and aquatic made
their last appearance (mcludmg dinosaurs, pterosaurs, ichthyosaurs and plesiosaurs). Among
the invertebrates, the ammonities, a dominant group of marine cephalopods also became extinct
at the same time. A significant change is also noted in the plant-kingdom too. Most of the
dom~nanti~g gym~osper~s and large ferns became extinct at the same ti1i1e giving room for
the mcommg fru1t-bearmg flowering plants that ultimately become dominant in Cenozoic.
However, the small amphibias, small reptiles and rising birds and mammals of Mesozoic could
be able to cross the Mesozoic-Tertiary boundary and the last two groups become the dominant
forms of Cenozoic.
--
DINOSAURS 335
Some probable causes resulting in such abrupt extinction of dinosaurs and others organic
groups at the end of Mesozoic are discussed below :
External causes
Competition : Dinosaurs persisted through rivality and war-fare. Carnivor dinosaurs were
the killers of many vegetarian groups. Other sources of competition came from the
contemporeneous reptiles, mammals and birds. Some have suggested that dinosaurs were the
unfortunate victims of egg-eating mammals. But this type competition is always present among
animals and it is impossible to believe that such activity could cause abrupt extinction of such
a dominating group.
Food : Most of the dinosaurs were herbivorous feeding on large gigantic gymnosperms and
fems of contemporeneous times. It has been suggested that there was a significant change of
plant kingdom towards the end of Cretaceous .with gradual extinction of most of the Mesozoic
gymnosperms and ferns with the introduction of angiosperms, many of which are deciduous ·in
habit. Plant-eating dinosaurs failed to adapt with this new type of plant food and died. This
might have an indirect effect on carnivore dinosaurs too as most of them Jived on the flesh of
herbivores. Other suggested that some plants of this time developed poisonous alkaloids. Some
flowering plants developed new types of wind-blown pollens. Some plants became poisnous
for greater concentration of selenium in soil derived from contemporeneous large scale volcanic
activity. All these had ultimately poisoned the herbivorous dinosaurs resulting in various diseases
causing their death and this had also affected carnivorous forms due to shattering of the food-
chain. At best, all these can explain extinc_tion of terrestral forms of dinosaurs.
Environmental and climatic changes : According to other group of scientists, the end of
Mesozoic was marked by some catastrophic events affecting the whole world. There was a
significant change in distribution of land and sea due to breakdown of southern and northern
supercontinents; there was beginning of the Alpine-Himalayan orogeny causing earth movements;
world wide volcanic eruption broke out both in continental and oceanic regions; there was
reversal of earth's magnetic field and marine transgression and regression. All these had serious
effect on the earth's climate and environments.
The critical evaluation of the various effects of these geologic events have indicated that
change of climate and temperature was brought out both in land and sea as a result of ~ontinental
drift. As the continents moved apart, sea-floor spread out with volcanism in mid-oceanis ridges,
and island arcs increasing the temperature of sea water. As ridges grew and expanded, the
capacity of ocean basins was reduced and oceanic water overflew on the low lying land causing
flood and a general fall of temperature of land areas. Dinosaurs and other giant reptiles were
especially affected as they were habituated with an un.changed mild temperature persisting
th~oughout the Mesozoic. They failed to adapt themselves with any such drastic change of
climate especially for want of any type of insulation-cover on their skin (as they were apparently
cold-blooded animals).
. Alternatively, a reversal of earth's magnetic poles might have caused a temporary breach.
10
the ozone-shield of the earth which otherwise protects the organisms from harmful cosmic
rays. Change in oxygen percentage in atmosphere has been also suggested. Analysis of 'ai.r-
bubbles' trapped in fossil resins also indicates high percentage of 0 2 gas (35% to 40%) m
.... _.
336 PALAEONTOLOGY
Cretaceous atmosphere that might have some bad effects on organisms. A startling theory has
been proposed by some suggesting that the dinosaurs killed themselves off with their own
flatulence (wind). The methane, they produced caused earth's atmospheric temperature to warm
up, creating a kind of greenhouse effect. This might have increased the global warming to a
point where dinosaurs were unable to stand the heat.
At present, three models are gaining ground to explain the mass extinction at. the K-T
boundary which are : catastrophist extra-terrestrial model (Alvarez, 1990), catastrophist
vulcanological model (Courtillot, 1990) and the gradualistic ecological succession model
(Archibald and Bryant, 1990). The first model proposes the disappearance of dinosaurs as a
result of a major extra-terrestrial impact on the earth. The second version explains the same
event by means of major volcanic eruptions. The gradualistic model sees a decline caused by
a long-term climatic changes in which dinosaurs gave ways to more efficient mammals. There
are several evidences of the impact hypothesis such as (i) presence of a large and deep crater
(about 180 km diameter) has been identified within the Late Cretaceous sediments in the Yucatan
Peninsula near Gulf of Mexico, (ii) presence of several sites (more than I00) on the earth surface
from where abnormal content of irridium has been reported (irridium is an uncommon element
on the earth but common within meteorites), (iii) presence of shocked quartz grains and glassy
spherules (microspheric molten droplets of rocks) both are supposed to be the effect of meteorite-
impact, (iv) presence of some peculiar minerals like microdiomonds and spinels which require
high temperature and pressure for their formation, (v) sudden loss of angiosperm taxa and their
replacement by. ferns and then a progressive return of normal flora. All the features (ii to v)
are found at several places near K-T boundary and they together strongly suggest the meteorite
impact theory. The immediate and subsequent effect of this massive impact was similar to that
of a world after a hypothetical nuclear bomb explosion.
The vulcanological model has also tried to explain some of the abnormalities at K-T boundary
(mentioned above) by volcanic activity. Large scale volcanic eruptions have been known from
India (Deccan traps) and several other continents at K-T boundary that could have caused
formation of dust cloud through explosion and resulted in subsequent effects similar to those
found in case of an extraterrestrial impact. But shocked quartz and glassy spherules, as argued
by petrologists and geochemists can hardly be produced by any volcanic explosion.
The gradualistic model sees gradual decline and time to time disappearance of many groups
of animals and plants right from the beginning of Mesozoic caused by long-term change of
climate cum ecosystem in which subtropical dinosaurs gradually gave way to more efficient
mammals.
However, there is every possibility that a combined effect of all the three models mentioned
above might have caused mass extinction at K-T boundary. There was long time decrease of
flora and fauna as explained by gradualistic model and the final extinction at K-T boundary
was the result of combined effect of impact-induced stress and large scale volcanism.
Internal causes
Dinosaurs were the supreme rulers on the earth throughout the Mesozoic. They lived in
favourable climatic condition for a long time with abundant supply of food materials but without
any serious competition coming from other contemporeneous animals. Such a condition would
DINOSAURS '.B 7
cause gradual loss of adaptibility of the animals, gradually making them.very much conservative
with that particular condition. Such a condition would also lead to the production of
progressively larger size and a longer life of .dinosaurs which could be correlated with their
late sexual maturity and ultimate loss of fertility. This stage is called 'racial senility' of the
animal group. A sudden change of climate and/or environment at this stage, however mild it
might be, would be fatal to that animal group. This was actually happened in case of dinosaurs
and other gigantic reptiles at end of Cretaceous. Loss of fertility of dinosaurs, especially in
females is sometimes attributed to some diseases of pituitary gland which is yet to be proved
by any conclusive evidence. However, observation on some Cretaceous dinosaurian egg-fossils
has shown some peculiar features. These egg-shells are multilayered and relatively thick. Youngs
within these eggs might have found difficulty in coming out by breaking them and many of
them became dead within_the egg shells, failing to come out of the shells on maturity. Moreover,
female dinosaurs had to supply more and more calcium from their body for making such thick
egg-shells which might have resulted in alarming fall of their blood-calcium level leading to
some. sort of diseases causing their death or loss of fertility.
Conclusion
There are evidences of several geologic events throughout the Mesozoic times. There was
splitting of continents, spreading of sea-floor, large scale volcanism both in marine and
continental sectors, submergence of some continental coasts below the sea, emergence of new
lands due to retreat of sea in other parts. Most of them must have caused a gradual change of
world-climate and this appears to be the most plausible explanation of mass extinction of many
dominant groups of organisms, both plants and animals at the end of Cretaceous. Within this
time there was elimination of many gigantic reptiles both from land and sea. Along with them
gradually disappeared many gymnosperms and ferns of Mesozoic times. There was also large
scale reduction of phytoplanktons in sea. Phytoplanktons form the basis of the marine food-
chain and their sudden fall certainly had set off a chain reaction in the entire marine ecosystem
causing disappearances many fishes, alJ sea-monsters (reptiles) and also extinction of the most
dominating cephalopods, the amononites. Final -blow came at the K-T boundary when the earth
experienced a large scale volcanism and impact with large extra-terrestrial bodies resulting in
several sweeping environmental cum climatic changes. This was enough to eliminate dinosaurs.
Such sweeping environmental changes should normally be expected to affect all animals and
plants but the fact is that end-Cretaceous extinctions were rather selective. Certain animals
survived because they have warm-blood with insulating covers (the hairs in mammals and
feathers in birds) that helped them to adjust their body temperature to cope with the changing
environment. Smaller reptiles survived possibly because they fled away easily from the disturbed
areas and stayed passively in some other suitable places (caves, holes etc.) as done by most of
the ruling terrestrial reptiles and amphibias during the winter season. Dinosaurs, so much
conservative to a particular environment, failed completely to adjust themselves with these
changing conditions at the end of Cretaceous. Their gigantic size, poorly organised brain, lack
of any kind of insulation on their skin, and overall their racial senility badly affected them
causing their total extinction.
Chapter 25
EVOLUTION OF HORSE AND ELEPHANT
25.1 EVOLUTION OF HJ)RSE

A~ Best documentation of evolution
Toe horse belongs to order .Perissodactyla and family Equidae. It is claimed that origin
and evolutionary history of the horse is known most clearly and completely. The reasons behind
this are as follows :
In North America the entire evolutionary sequence of the horse is recorded in the form
(a)
of fossils within some undisturbed and uninterrupted series of sedimentary deposits
ranging in age from early Eocene to Recent.
Because of the grazing habit of these animals, their adaptation to life on savannas and
(b)
lains and their tendency to live in large herds, fossils of horse wherever they are
~reserved, are found in large number.
The pattern of evolution of the horse is found remarkably simple showing a single line
(c) of adaptation instead of showing diversification through multiple radiation along different
lines. Such an evolution exhibiting progressive change of characters in some
predetermined. directions is_ oft~n called orthogen~sis. But ~het~er the evolutionary
pattern. exhibited by the horse 1s truly orthogenetlc or not, ts qmte debatable. Many
researchers have the opinion that orthogenesis is common in case of evolution and
development of different organs in an animal's body, but it is rather an unusual
phenomenon as regards the evolution of an entire animal group. It is argued that the
pattern of evolution of horse appears apparently orthogenetic possibly because we yet
to discover the fossils of many other offshoots of this group diverging from the main
line. Moreover. it is seen that in Middle and Upper Tertiary times, there are of course,
cases of lateral offshooting from the main line of evolution of horse which resulted in
some forms that exhibited both progressive and conservative characters.
B. Adaptation of the modern horse (Fig. 25-1)

The horse lives in grassland or savanna, fed mainly on grasses. Its body and other associated
structures are adapted to this type of life and food habit. Animals living in open grassland,
must have the capa~ity .of quick movement and running to save themselves from aggressive
predators. Thus, with increase of size the horse develops elongated slender limbs with
unguligrade posture (walking on toes). But in contrast to other ungulates evolution of the horse
h~s. gone to an extreme poi.nt resulting in suppression of all digits except the middle one (Ill
digit) s~ that the. horse ultimately becomes an one-toed animal (Fig. 25-1 a,b). To make the
mechanism of quick movement most effective and perfect, the III-digit of the limbs becomes
greatly elongated a~d. the metatarsal/metacarpal of the leg forms the powerful cannon bone.
One of the most stnkmg adaptations of the horse's foot for rapid running consists of a set of
EVOLUTION OF HORSE AND ELEPHANT
339
----Carpus
r + - - - - Enlarged Metacarpus
Forming Cannon Bone
(b) SPRING MECHANISM OF HORSE'S FOOT

~r----- Hoof
(a) ADAPTIVE MODIFICATIONS OF SKELETAL
ELEMENTS OF FOOT OF THE HORSE
FIG. 25 - l : ADAPTIVE MODIFICATIONS OF STRUCTURE OF HORSE'S FOOT
Equus Hyracotherium
Equus
Leli Hindloot
Left Forefoot
I t7 ~
n
IV
Ill Ill 111 Ill
(a) CHANGE OF DIGITAL NUMBER IN HORSE-LIMB
Top Vil!W Of lipper And

A Molar TooJh Top View Of Upper Lower Molar
(Arachydont) And Lower Molar A Molar Tooth
Hyracolherium (Hypsodont) Equus
(b) CHANGE OF STRUCTURE OF MOLAR TOOTH IN HORSE
FIG. 25 - 2: CHANGE OF SOME SKELETAL FEATURES IN THE EVOLUTION OF HORSE
FROM HYRACOTHERIUM TO EQUUS
340 PALAEONTOLOG'(
spring ligaments lying on the posterior surface of foot connecting cannon bone with sesamoid
bone and the latter with phalanges of the hoof. The arrangement is elastic like a rubber band.
The impact of the weight of the body upon the foot and latter upon the hard ground is translated
into an upward and forward propulsion. This gives horse a very rapid movement. For making
the movement smooth across the air the horse also develops a streamlined body.
Along with the increase of length of limbs, the horse might have faced difficulty in eating
grass from the ground. The problem has been solved by developing a flexible and elongated
neck. To avoid extreme lengthening of neck, as it would be disadvantageous for a large headed
animal like the horse, growth of the neck has been coupled with elongation of frontal portion
of the head anterior to eyes, thus developing an elongated muzzle. One result of such an
elongation of jaw is the formation of a gap between anterior and posterior set of teeth which
is called diastema.
As a grazing animal uses its front teeth cutting grasses, incisors of horse become chisel
-shaped with very sharp edges. Canines are of no use and hence highly suppressed. But the
animal has to give the maximum stress upon its cheek teeth, for massive chewing of harsh
grasses with grits and silicified cuticles. Such food materials are so abrasive that they cause
rapid wearing of the teeth. The problem has been solved by the horse in two ways; firstly
transforming all premolars into molars; secondly, developing high crowned hypsodont teeth
which grow continuously as they are eroded throughout the life of the horse. On the upper
surface of the cheek teeth, crests of cusps are variously conjoined and the enamel of these crests
is folded enclosing the space covered by softer cement. Thus as the cement wears away more
rapidly, the complex and folded enamel band remains projected slightly above the softer dentine
and cement-covered area keeping always a roughened condition of the tooth-surface. In fact,
these teeth become self-sharpening, self-renewing and highly efficient grinding tools.
C. Basic ancestral characters

Fossil evidences indicate that condylarths (Fig. 23-9) which were ancestral to so many groups
of hoofed mammals, were also the probable progenitors of the perissodactyls.
Lower Eocene Hyracotherium, the most primitive known horse exhibits a close morphologic
similarity with a North American Palaeocene condylarth, Tetraclaenodon, especially as regards
it structure of limbs and cheek teeth. The cusps of Tetraclaenodon were found transformed into
transverse ridges or lophids. The changes from Tetraclaenodon to Hyracotherium points to a
shift in adaptation for efficient browsing and quick running over the ground in response to
pressure from contemporaneous aggressive predators.
Hyracotherium, (Fig. 23-9), also designated as Eohippus can be called as a prototype of
perissodactyls. It was a small lightly built animal about the size of a fox-terrier ( 12 inches
height). It had a low skull, short neck, small but slender limbs. The ankle and the wrist were
raised. There were four digits in .forelimbs and three in hindlimbs, but the middle digit was
comparatively larger, clearly indicating an. adaptation for running. Skull had a small brain case
but no postorbital bone. Cheek teeth were brachydont low-crowned with four rounded cusps in
the upper molar namely protocone, paracone, metacone and hypocone. There were also two
small intermediate accessory cusps, the protoconule and metaconules, connected with two inner
cusps by two low oblique ridges (protoloph and metaloph). The structure of forefoot and hindfoot
and molar tooth of Hyracotherium and Equus is shown in fig. 25-2.
J, VO l 1'/0N W IIONS F \ NI l'I /;' l'IIAN1' 341
D. P1"t>gn·ssh·t~ trmuls of twolution

A)lllp u'ing lh~ ·h.um:tcrs of moll ·rn ,~·quus with those of .. artiest ancestral form Hyrac:o-
thrrium it is pl_)Ssiblc to tral:(;'I out th t' lin "' s of ·hangr of morphologic churncters during
volutio1ury <k l'lt pn,c.,11t from l ' lll'li ·st t,l modern hors•. Thes • trends muy be summarised as
foll ows-
i) lm:1\'HSl' ~,f si'lc with lwi rht im: 1'l·nsing from t 2 inch "' S to above 72 inches,
(ii) Length ·ning of limbs with grndunl shiftin , of ankl · and wrist uwuy from the toe.
(iii) ~ngth ·ning c1f ne ·k nnd straightening uncl stiffening of buck producing a stream-lined
body.
(iv) R •du ·ti n of numbers of to •s in limbs making all the limbs one-toed.
(v) L "11 'th nin " th " portion of h ·ad in front of eyes forming a muzzle.
(vi Wid ~ning of incisors with v ry sharp edges.
(vii) Molnrizatit)11 of premolars.
(viii) Development of high-crowned cheek teeth (hypsodont) from original low-crowned
brnchydont type with consequent deepening of frontal part of skull and lower jaw to
accommodate those teeth.
(ix) Development of complex fold-pattern of enamel on the upper surface of the crown of
cheek teeth.
(x) Gradual increase of complexity and size of brain.
The first five changes have been brought about to acquire fast running on the plains while
the remaining changes are in response to their change of food-habit from browsing to grazing
type. The change of food habit has been attributed to a great environmental change at Late
Oligocene in North America causing retreat of humid forests and spread of grasslands. However,
most of these changes were initiated from the very beginning of the Eocene times and continued
upto the upper part of Cenozoic when appeared the modern horse.
E. Phylogenetic history (Table-15)

Although, a complete history of horse is known from the fossils preserved within the
Cenozoic rocks of North America, the fossils of the earliest member Hyracotherium/Eohippus
have been also reported from Europe (London Cl<\}' Formation) and Asia. From fossils, it appears
that at least during Eocene-Oligocene times the horse exhibited an orthogenetic pattern of
evolution proceeding along a single line of progressive development. The successive forms
appeared are : Eohippus (Lower Eocene)-+ Orohippus (Middle Eocene)-+ Epihippus (Upper
Eocene)-+ Mesohippus (Lower Oligocene)-+ Miohippus (Upper Oligocene). All the characters
mentioned earlier showed a gradual change in these successive forms. Miohippus, thus at the
end of Oligocene attained a height of about 24 inches. It exhibited suppression of fifth digit in
forelimbs producing all limbs with three functional digits of which middle one was larger. The
last three premolars were molarised and the cheek teeth became strongly crested with one in
upper teeth became W-shaped but two in lower teeth V-shaped with points directed outward.
These sharp crested teeth efficiently functioned as choppers for cutting leaves and other plant
materials taken as food by this browsing horse.
342 PALAEONTOLOGY
With the advent of Miocene three lines diverged out from Miohippus. In the main line it
gave rise to Parahippus of Lower Miocene age. The first lateral offshoot from Miohippus led
to Miocene Archaeohippus which remained conservative as regards the structure of its skull
•
teeth and feet and they subsequently became extinct. At other line of decent was Mio-Pliocene
form Anchitherium which showed an increase of size but it retained the teeth and functional
three toed-limbs like Miohippus. They were probably forest-dwellers that browsed deep wood-
lands. Anchitherium migrated into the Old World towards Late Miocene showing a wide dispersal·
there and became extinct after Pliocene. In North Aml"!rica evolved Hypohippus from
Anchitherium in Pliocene which sometimes attained a large size somewhat like the modem
horse, but ultimately it failed to cross Pliocene-Pleistocene boundary.
Merychippus, evolved from Parahippus in Upper Miocene in the main line, was a horse
showing some distinct advances over the preceding forms. It had a size of a pony, having three
toed limbs, but walking on larger middle toe which was provided with a rounded hoof. The
high-crowned cheek teeth were covered with cement. Crests of crown became variously
conjoin~d with protruded enamel bands showing complex folding making the teeth efficient
grinders. This was a clear indication of change of food habit in Merychippus from browsing
to grazing type. Merychippus also developed a postorbital bar behind the eye opening in
between orbit and temporal region which is a typical feature of all the later horses.
Two groups of horses emerged out from Merychippus in Pliocene. A lateral offshort leads
to the form, represented by the cosmopolitan genus Hipparion which migrated to all countries
except South America by Plio-Pleistocene times. It was a lightly built horse with enlarged skull
and complex enamel-folds on crown surface of molar tooth like advanced horse but it remained
conservative in the retention of three-toed feet. Hipparian fossils are so abundantly found within
Pliocene rocks that Pliocene mammalian faunas are often deginated as Hipparion faunas. There
was a sharp decline of Hipparion after Pliocene and it became extinct towards the end of
Pleistocene.
Pliohippus, appeared in Pliocene from Merychippus in the main line, was a progressive horse
in all respect and truly an one-toed equid. The side toes are reduced to splint bones concealed
below the skin at upper part of limbs and the same structure is now maintained by .. the modem
horse.
From Pliohippus, towards the end of Pliocene, diverged two lines. One offshoot line Jed to
the Pleistocene form Hippidium which originated in South America. h was a short-legged but
large sized horse which became extinct towards the close of ice-age. From Pliohippus in the
main line, appeared the modern horse Equus in Upper Pliocene. Equus, first appeared in North
America, survived there for the entire Pleistocene but became extinct a few thousand years ago
towards the close of ice-age. Equus that migrated into South America met the same fate.
Another group which had migrated into Euresia and Africa ultimately becomes the horse of
the present day with worldwide distribution. At present, it is found with a number of species
represented by such forms like horses, zebras, asses etc.
F. A few more points

Many of the evolutionary changes with time found within a particular animal group are
measurable quantitatively. In some cases, it is often found that the change of two such characters
EVOLUTION OF HORSE AND ELEPHANT 343
may show interdependence. Such phenomena have been observed in the evolution of horse.
The increase of length of facial part of the skull and the increase of size of the body went on
side by side but maintaining always a definite relation between them. These two characters
furnish an instance of positive allometry and they bear a close parallelism. Again, the increase
of size, if correlated with decrease of number of toes in the limbs, becomes an example of
negative allometry.
Although the history of evolution of horse is known more or less completely, some of the
events related to its evolution are yet to be explained. Why different groups of horse, evolved
in North America remained more or less confined to that continent instead of showing migration
TABLE-15
OUTLINE OF PHYLOGENY OF HORSE
RECENT
PLEISTOCENE Equus (S. America) (Ext.) Equ~s (Euresia, Africa)
(Ext.) t
'~ Equus (N. Amer.) (Ext.) Hippidium (Ext)
t (S. Amer.)
PLIOCENE
UPPER
Hipparion
·~(N. Amer
Euresia)
l
Equus (N. Amer.) Hypohippus (Ext.)
' (N. Amer.)
LOWER Pliofppus (N. Amer.)
MIOCENE Merychippus (N. Amer.) Anchitherium (N. Amer.)

UPPER (Euresia, Africa)
LOWER
1 Arr:haeohippus (Ext.)
~(N.Amer.)
Parahippus {!l· Amer.)
OLIGOCENE Miohippu/. ~
UPPER (N. Amer;)
Mesohippus (N. Amer.)

LOWER
i
I
EOCENE Epihippus (N. Amer.)

UPPER
i
I
Omhlppus (N. Amer.)
MIDDLE
r
HyracotheriumlEohippus
LOWER (N. Amer.• Europe. Asia)
a• I ' ,.,.
PALAEONTOLOGY
,\ molar tooth
Mocri1hcrium
(height 2 feet. ancesuul elephant)
Molnr \OOlh
EV LUTlON OF DIFFERENT GROUPS OF ELEPHANTS

FIG, 25 • 3 (SH WING TIIEIR RESPECTIVE SKULL AND MOLAR TOOTH)
...
~

to other countries. Absence of many intermediate forms in the evolutionary line of horse outside
North America sometimes has made many European palaeontologists to believe that evolution
occurred by occasional jumps. Cause of extinction of Equus at the end of Pleistocene in North
America, inspite of the change of climate in a favourable direction also remains unexplained.
25.2 EVOLUTION OF ELEPHANT

A. Major traits of elephants ,
Elephants belong to the order Proboscidea within the class Mammalia. Although, surviving
in considerable numbers in Asia and Africa, elephants are now representing the last members
of a dying group. At present, they are represented by only two genera each with a single species;
one is African elephant Loxodonta africanus and the other is the asiatic elephant, Elephas
maximtts. However, fossil evidences have proved that various types of elephants, some
exhibiting gigantic forms inhabited the world quite abundantly at least during middle and upper
part of Cenozoic Era. Unfortunately all, but two, failed to cross the Pleistocene-Holocene
boundary .
The most striking features of this fascinating creature are its gigantic size, a huge bulky
l?<><IY, pillar like legs, a long flexible probosis, a pair of tusks and very peculiar type of molars.
The African elephant differs from its Asian counterpart in having great palm-leaf like ears and
somewhat larger and more curved tusks.
Many of the adaptations of the elephant are obviously related with its huge size. To support
the weight of the body, limbs of the animal aquire a strong pillar-like shape. The feet have
retained all the five toes, but much of the weight of the body is supported by a pad of elastic
tissue that forms a sole like a rubber-heel below each broad foot. As the height of the animal
is considerable, it is faced with the problem of feeding from the ground. The same problem is
solved by the horse by increasing the length of its neck and skull. But as the head of the elephant
is quite large and heavy, lengthening of neck or head would be mechanically disadvantageous.
For this, they have developed an organ like probosis or trunk. The development of the probosis
is the result of continuous elongation of snout and upper lip with the support of upper jaw.
The two external nostrils are situated at the tip of the trunk but the bony openings of the internal
narcs have receeded to be located on the middle of forehead. This recession of nares helps
firm anchorage of the powerful musculature of the probosis. Another peculiarity of the elephant
is related to its teeth. A pair of tusks have developed by elongation of its second pair of incisors
of the upper jaw. The ivory, comprising them consists of dentine elements of the teeth. A large
open pulp cavity at the base each tusk provides their continuous growth. The molar tooth of
the elephant is also unique as regards, its structure and replacement. Each molar is large, much
elongated, and consists of some parallel flattened plate-like ridges or lamellae, lying transversely
side by side. Each ridge has an outer margin of enamel, that en~loses softer dentine inside and
cement fills the space between the successive ridges. This molar has a broad grinding upper
surface on which ridges of enamels protrude out owing to more rapid wearing of softer dentine
and cement occurring in between the ridges. More surprising is the mode of replacement of
the molars. In most mammals there are two sets of teeth, a set of milk teeth, replaced by u set
of permWlent teeth Wld the replacement takes pluce mostly in a vertical manner. But in elephant,
the t~ec molars replace one another successively rather than replacement of all the three molars
at a hme. Thus at any time of life only four molars of an elephant remain functional one in
346
PALAEONTOLocy
each side of the jaw. As these molars wear out, they are replaced by the next set of
. a Iong1tu
bu t m . d ma
" I manner, t he new tooth pushes out the weared one in a forward ct·molars
. •
.
from the rear side. rrect1on
B. Ancestral proboscidean 'Moeritheres' (Fig. 25-3)

All the different forms of elephants living or extinct probably arose from a common ancestral
stock often called moeritheres. Their fossils are obtained from Middle to Late Eocene rocks of
No~h . Africa. From this, evolution of elephants diverged along a number of lines of adaptive
ra~1at1on. T~e descen~ants spread out from Africa all over the world except Australia by
M10cene-Ple1stocene times. But the ·overall phylogenetic history of this group of mammals
becomes somewhat complex, partly because of several gaps in fossil record and partly for the
existence of parallel evolution among the different independent groups.
Moeritheres were represented· by several forms but the most well known one was
Moeritherium ·w hose fossils are fou~d from Late Eocene rocks of Egypt. It was a heavily built
animal of the size of a tapir. It had short probosis, elongated limbs and spreading feet with flat
hoofs at the end of the toes. The skull was rather long, with eyes set far forward. The forward
shifting of eyes resulted in the elongation of zygomatic arch. The second incisors of both the
jaws were slightly elongated and projected forward giving indieation of development of the
initial tusks. Canine and third incisor of the· lower jaw were suppressed. The molars were low
crowned with two transverse crest~. each being formed by two large cusps placed side by side.
The longitudinal method of .replacement of teeth however came at a much later period. The
dental formula was 3133 x 2 = 36. This morphology of Moeritherium may give an idea of the
. 2003 . ·
generalized and basic proboscidean characters.
C. Trends of evolution
Starting from . this basic proboscidean form, evolution proceeded along different lines ~f
adaptive radiation. Although each line shows independent. and varied development, som:
dominant trend_s running through the wide range of probosc1dean forms can be traced out a
follows :
(i) Increase of size.
(ii) Lengthening and thickening of limbs with the development broad feet.
(iii) Extraordinary increase of skull size.
(iv) Shortening of neck for keeping a balance between bulky body and heavy skull.
(v) Gradual elongation of lower jaw, although secondary shortening may be found.
(vi) Growth of trunk by elongation of upper lip and nose.
· . of the second pair of incisors to form the tusks used for d n·11·mg ground.
(vii) Over-mcrease
and also in defence. . h · and
Specialization of cheek teeth in various ways as adaptation for better c ewm8
(viii)
grinding plant foods.
..
"!::" · •.

D. Phylogenetic history (Fig. 25-3) (Table 16)
The relationship among the various groups of proboscidea in Early Tertiary times remains
somewhat uncertain. According to some, the evolution started from Palaeomastodon which
arose from moeritheres and became the key point of radiation at the beginning of Oligocene
while moeritheres remained as . an isolated offshoot. Other considered the moeritheres as the
earliest and central forms from which diverged two distinct lines, one included the dinotheres
and the other one was the elephantoids.
(a) Dinotheres
Dinotheres were distinct from the very beginning evolving along a narrow path throughout
the Cenozoic times before its final extinction in Pleistocene. Their fossils are found first from
Miocene rocks of Africa. Although, they were considered as derivatives of moeritheres, there
is no fossil-evidence showing a transition between them. This group includes a large number
of forms in its long history but they are morphologically so steriotyped (except showing increase
of size) that they are all placed with a single genus Dinotherium. This is a fine example of an
jnitial rapid evolutionary development followed by a long period of evolutionary stability. Late
Cenozoic forms of Dinotherium were giant, long-legged animals with highly flattened skull
(unlike other proboscideas), and without any tusk from upper jaw. On the other hand, two large
tusks were projected out from lower jaw which curved backward toward the body like a pair
of huge hoods. Such tusks were probably used in digging the ground for uprooting trees. The
crown surface of cheek teeth showed two/three cross-crests each formed by 3-4 small cones.
Trunk was well developed. Dinotheres first appeared in Arica and soon dispersed over Euresia
but failed to reach the New World.
(b) Elephantoids
Elephantoids on the other hand, diverged at least into three directions after Miocene and
formed three independent groups which are : the gomphotheres or long-jawed mastodonts, short-
jawed mastodonts (true mastodonts) and the elephantids (true elephants). Modem elephants are
the derivatives of the last group while the other two group became extinct after Pleistocene.
Long-jawed mastodonts or gomphotheres are also sometimes called trilophodontids. They
appeared first in Oligocene from moeritheres and were represented by two genera in Africa,
Palaeomastodon and Phiomia. They were comparatively larger than moiretherids and exhibited
larger skulls, increased height, much elongated jaws (especially the lower jaw) and a well
developed trunk. The most characteristic were their two pairs of tusks; the upper pair were
projected forward and downward while the lower pair projected horizontally. The cheek teeth
were low crowned with three transverse pairs of blunt cusps. In Phiomia, the trunk and the
lower tusks were more pronounced. There is a considerable gap in the record of proboscidean
evolution for most of the parts of Oligocene and the next group of long-jawed mastodonts are
found in Middle-Upper Miocene represented by the genus Gomphotherium, also called
Trilophodon. It possessed much elongated lower jaw with a pair of large horizontal tusks and
a well developed probosis. The molars possessed three transverse cross-crests. each with three
larger cones and a few smaller accessory connules. Trilophodon was a widely distributed form
of Mio-Pliocene whose fossils are found in Africa, Asia, Europe and North America. A little
younger forms of Pliocene in the same line are Serridentinus. Synchonolophus and
Tetralophodon. Serridentinus exhibited a sinuous enamel pattern produced by partial wearing
348 PALAEONTOLo
Cy
of crests of cones and connules on molars. Such complexity of molars reached their cul . .
within Synchonolophus and in the Lower Pleistocene form Stegomastodon. l.n the lac.it h:•~ation
the lower jaw became shortened with disappearance of the lower tu sks while their upper ;rrns
became larger and curving upward. The Pliocene Rhynchotherium exhibited downwardly cuUsks
lower pair of tusks. The molars became much more complicated in the Plio-Pleistocene g;ed
Dibelodon. Tetralophodon were nearly similar to Trilophodon but with four transverse cus us
on each molar with smaller accessory connules. A specialized form of gomphotheres wast~:
Pliocene Amebelodon of North America and Platybeledon of Asia showing shovel-like flattened
lower tusks . All long-jawed mastodonts are found for the last time in Pleistocene.
Short-jawed mastodonts or true mastodonts followed another independent line in the evolution
of the elephant. They were characterised by a lower jaw without tusk. They possibly arose from
Palaeomastodon in Oligo-Miocene times. The Miocene form is called Miomastodon which was
followed successively by Pliomastodon in Pliocene and Mastodon in Pleistocene. The most well
known Pleistocene form is Mastodon americanus of North America. It was a large heavily built
form with a well developed trunk and a pair of strongly curved upper tusks. It had sharply
cross-crested molar as seen in all other members of this line. They persisted upto the end of
Pleistocene and were contemporaneous with the early man in North America.
Stegolophodontids, the first members of elephantids appeared in Pliocene in Africa possibly
emerging out from some trilophodontid-ancestors. They were represented by the genus
Stegolophodon which was a medium-sized elephant with shortened tusk on lower jaw. The
upper tusks were long and much curved. The molar became elongated, cross-crested; each crest
with few large cones and many smaller connules. It was a common form of Pliocene of Africa
and Euresia. From stegolophondonts arose the representatives of the earliest group of modem
elephants, the stegodonts. They appeared in the Old World in Late Pliocene and continued upto
the end of Pleistocene. The Pleistocene genus Stegodon was a gigantic elephant, long legged
and with a large and deep skull. The upper tusks were very long and curved upward but the
lower jaw was short and tusksless. Molar teeth became considerably elongated, upper surface
of each with many low cross-crests made up of number of connules. To accommodate such
elongated molars, elephants take an adaptation of using one molar at a time and replacing it
by the next molar in a longitudinal manner. The step from stegodont, first to mammoths and
finally to the modem elephants showed a V(?ry rapid change of characters of the cheek teeth.
Stegodonts of Lower Pleistocene had low-crowned teeth but towards the end of Pleistocene
molar teeth of mammoths and modern elephants exhibit highly compressed ridges on the er?~"
making them as tall, parallel plates or lamellae rather than V-shaped ridges. From the giganuc1ty
and abundance of mammoths. Pleistocene or the great ice age is sometimes called the age of
mammoths (name commonly applied to extinct elephants). During Pleistocene they dispersed
throughout the world except South America. There were various species of mammoth~. one
group led to modern asiatic elephant Elephas maximus and the other group gave nse to
Loxodonta africanus, the present african elephant. Mammoths so~e!imes a~!ained I 0-15 fe::
height. Perhaps the best known forms are the wooly mammoths hvmg u~t" the very en:u 5
Pleistocene in North America and Euresia. They were contemporaneous with the early gr P8
of modem man and the latter often left their pictures on the cave walls. Moreover, we have .
omplete idea about the morphology of wooly mammoth owing to the good fortune that_ theifr
C • • the P Ie1stocene
comolete body fossils are found frozen w1thm · • of s i·beria · These foss1 1s 0
ice
N OF /ION. E r\NI> £/.1.,'}->JIANT
£' oiuno · 349
ooly nmnu1101hs •xhihit Iheir hoJ :skin co~cr •d wi1h lark hrown coloured hairs, few inches
0
" fo,•1 in k ng1 h· The c"'"'' s ol c Xt met 1011 of 111ammot hs is however di !'lieu It 10 predict
;~':rrcct of great cl imal il'-l' han ' ' following Ih ' end ,if Pl 'isl nee n · ,1ac ial ion can 1101 be ruled
our. Many r •searchers
. . .. b ' \1·<' 1ha1 I con1c-mpor.111 ·ons man 111igh1 ha, · had so111ell:ing lo .do with
hclt
. But ii is d It lieu It to c t 've t tal Ihey :1!011 • run Id had bronght abou1 the end ol such a
it. . >rotis• and gigantic gr up of
nlllllC . crcatur,·s.
. Possihly lh 'i. r ext incl ion was the result of a complex
factors of which we have a little hmt at thl! prl!~Cnt 11111,
TABLE-16
OUTLINE OF PHYLOGENY OF ELEPHANT
Recent
t
Elephas (Eures1a)
. f
Loxodonta (Africa)
(Ext.) (Ext.) (Ext.) (Ext.)
Pleistocene
t t t
Dibelodmz Mammoths Mastodon
Pliocene (Ext.) Tetra/ophodon Stegodon Pliomastodon
(Ext.)
t
Stegoloplwdon
Miocene Miomastodon
Dinotherium Triloplwdorz
Oliogocene (Ext.) Palaeomastodon
Eocene Moeritherium
Chapter 26
PRIMATES AND ANCESTRY OF MAN
26.1 INSECTIVORA-PRlMATE TRANSITION
At
. the. end
. of Mesozoic
. ' with. th e f.a 11 f mighty
? · ·
reptiles, mammals began to diversify and
rad1~te 10 different Imes occupymg the different habitats vacated by the reptiles. One of the
earliest groups of mammals, the insectivores ascended the forest trees which offered them
refuge zone where competition was less intense and the supply of food was adequate. But lif:
became complicated as it was a new florid, a new dimension of life among the dense foliage
and branches of trees of the forest. Successful adaptation to this new arboreal life led this early
group of generalized insectivores to a new line of development terminating with the evolution
of primates. The transition, although not evident from any fossil document, probably took place
· n Palaeocene-Eocene times. But a glimpse of the ancient most primate may be visualized by
~ king at the modem tree-shrew 'tupaia', (Ptiloceras) and its relatives, living in the Orient at
00
ent (Fig. 26-1 ). Tupaia is a small animal of about the size of a squirrel having a long snout
pr~ imilar tail. They are mostly adapted for climbing and living on branches of trees in tropical
an st by their great toes, slightly apart from other digits, somewhat similar to primates. They
foresd minantly fruit-eating animals rather than feeding on insects. They also have a
are ~atively large brain and large eyes and the latter like primates separated from temporal
co~P by a postorbital bony bar. Thus morphologically and habituaJly they are very close to
region
. e of demarcatton . b etween msect1vores
. . an d primates.
. T hese features also suggest a
the Itn . . . . .
·t·on from insectivores to pnmates m response to a new type of hfe habit.
trans• t
_ BASIC LINES OF ADAPTATION FOR AN ARB?REAL LIFE .
26 2The most important character adopted by the early pnmates for a successful arboreal hfe
related to their locomotion. Hand and foot of a primate are characterised by five digits,
w~tadactyly with grasping thumb and big toe. These are necessary not only for climbing trees
pe d moving along the branches but also in helping the baby to cling tightly to the stomach of
:::e mother at the time when his mother is moving or jumping. Fossil foot-bones of primate of
about 52 m.y. old (Eocene) provide the evidence of existence of such grasping type of digits.
Another important trait of arboreal adaptation is the retention of two separate bones in forelimbs,
the ulna on outer side and the radius on the thumb-side. This structure allows a primate to
have prehensility of its limbs giving their greater mobility and rotation in a desirable direction
which becomes useful in a successful arboreal life.
In this new habitat, ancestral primates had to change their food habit also. The diet of
insectivores includes mainly soft bodied invertebrates which are quickly sliced and swallowed
by their tall teeth provided with sharp conical cusps. Such teeth are unsuiLable for chewing
rough and tough seeds, fruits and leaves which are mainly available as foods for the arbor~als.
As a result, ancestral primates developed short teeth with bulbous cusps suitable for crushing,
grinding and chewing.
350
PRIMATES AND ANCESTRY OF MAN 351
As fruits, seeds, soft leaves and associated insects are usually found at the terminals of the
branches, the early primate as an opportunistic omnivore found itself quite safe on such slender
branches by its grapsing digits. Not only this, the early primate was able to lift both of its
hands and could sat erect on a branch anchoring himself firmly by its grasping digits of hind
limbs. At this position it was possible for him to pick up vegetable foods or insects by its free
hands. This erect posture of sitting on hind limbs may indicate an early manifestation of erect
bipedal locomotion achieved by some higher primates like man.
Perfect vision is another basic requirement for arboreal life. From a very initial phase,
primates have developed their greater reliance on sense organs related to vision. Eyes become
large and are shifted towards the frontal part of the skull to achieve a stereoscopic binocular
vision. Such a vision is essential to make accurate judgement of distance before jumping from
one branch of the tree to the other as a misjudge leap may result a fatal fall. Eyes are further
protected by a postorbital bony bar separating them from the temporal region (a characteristic
feature of all primates).
There is a lot of controversy about which one of these basic traits were acquired by the
primates first. It might be possible that adaptation to arboreal life resulted in all other features.
Or, the change of diet and vision-pattern, developed among some ancestral groups, gradually
have made them successful arboreals.
However, most of the other traits found in primates are secondary achievements such as their
less reliance on olfactory sense which is emphasized by the loss of the naked rhinarium (the
moistened, hairless and tactile sensitive skin surrounding nostril of many mammals). Less use
of mouth and jaw for the purpose of picking foods-stuff causes shortening of snout, shortening
of jaw-bones and crowding of teeth on the jaw. Their dental formula is ~:!! x 2 or ~:~; x 2.
The zygomatic arches become laterally broad and strong increasing the activity of masseter
muscles attached to the arches. It is presumed that postorbital ridge of the primate originally served
to prevent deformation of eye-orbit caused by contraction of chewing muscles at the time of
crushing and chewing of food. Primates have also developed internal ear enclosed by a hollow
bony-structure called auditory bulla. Finally, they have developed comparatively a larger brain
reflected by their more agility, activity, curiosity and intelligence which have ultimately made
them superior to other mammals.
26.3 POSITION OF MAN IN THE ORDER PRIMATE

Order Primate
Division Prosimii Primitive or lower group of monkeys.
Suborder Lemuroidea Lemurs

Suborder Lorisoidea Lorises
Suborder Tarsiioidea Tarsiers
Division 2 Anthropoidea Higher group of monkeys, apes and man.
Suborder Platyrrhini New World monkeys, cebids

Suborder Catarrhini Old World 111Q.nkeys
/
PALAEONTOLOGY
g\ Jaw with teeth

Sku\\
Jaw with teeth
Anccslral inscctivora
f\G . 26 - \ : FAM\L Y MEMBERS OF PRIMATES SHOWING v ·AR\ATION OF SKULL AND

JAW (not to sca\e) ·
353
Superfamily Cercopithecoidea Monkeys

Superfamily Hominoidea Ape and Man
Family Pongidae Ape

Family Hominidae Man
26 4 KINS OF MAN (Fig. 26-1) .

· Lemurs : Lemurs representing very early and primitive forms of primate evolution, are n?w
inhabiting Asia and Africa. The common lemurs are small animals with. very long tails,
elongated flexible limbs and a much elongated muzzle giving them a fox-hke face . Le~urs
are widely distributed as fossils such as the Eocene form Notharctus from North Amenca,
Pleistocene Megaladapis from Madagasgar etc.
Lorises : The lorises, now living in India and Africa are separated as a group equal in rank
to the lemurs. This primitive monkey is also represented by two forms Loris, the sle~der ~orm
of India and Galago, the bush baby of Africa. The muzzle of the loris is reduced hke higher
primates and its eyes are more forward than those of lemurs. Indraloris is the only fossil
recovered form the Pliocene rocks of the Siwalik Group.
Tarsiers : Tarsiers, another group of primitive monkeys, appeared in Palaeocene and
continuing upto the present day, are found only in the East Indies and Philipines. It is a small
squirrel-type animal with a large pair of. circular eyes in front of face: It has a long tail and
long hind limbs for prolongation of calcaneum and navicular bone, which give its ability to
make tremendous leaps through the top of the trees. They are nocturnal animals. The Eocene
fossil Tectonius had enormous orbit and short jaws.
Monkeys : Monkeys, apes and man constitute a group called simians or anthropoids. Most
of them are able to sit in an upright position making their hands free. New World monkeys
(ceboids) appear to be more older and are found in Central and South America where they live
in tropical forest. Characteristically it has a very effective prehensile tail mainly for grasping
tree branches. The several living groups are squirrel-like marmosets, large-sized cebids, spider
monkeys, howling monkeys etc. Their fossil history is scanty. A Miocene genus Cebupithecia
is reported from Argentina and Columbia.
. O_ld w~rld monkeys or cercopithecids are ~ore represented in the fossil record especially
withm Oligocene and younger rocks. The earliest known genus is ·Parapithecus of Egypt of
Early ~ligocene age. Among the various forms of Old World monkeys the common groups
are Afncan rhesus monkey, guenons and the baboons. They were mostly omnivorous. Baboons
are grou~d living animals having elongated snouts and large canines thus often resembling
dogs. Asian monkeys are mostly ~epresented by macaques and semnopithecids. Fossils of Papio
(baboon), Macaque and Semnoptthecus are found in Plio-Pleistocene rocks of Siwalik Group.
~pes : Parapithecus was probably an ancestral cercopithecid monkey (Old World) from
;:•ch evo\v~d ~he apes. :he most primitive ape fossil is Propliopithecus recovered from the
. me rock y1eldmg Parapithecus. It showed comparatively a shorter and deeper lower jaw than
its ancestor an~ molars with five low cusps. O\igo-Miocene rocks of Africa have yielded several
other ape-fossils such as Aegyptopithecus .
.
\
PALAEONTOLOGY
f,_ (a) ARM DIGITS.,,,.
Man
(b) FOOT DIGITS
1,-.--llium1--~ ~u--Pubis
.,...___ lschium
~---sacrun1"1o--~f-
•
~~---Pubi " '..
\
I
Side View
Side View
Ape
Dorsal View
Hind Limb
Man
L
' I
(c)° PELVIC GIRDLE
Spinal Chord
Sigmoidal
I
Spinal----lia I
Chord I '~ _,/)--.._
I • l '
I
I I :' J ' .- - - - --\ (
'I \ I : ' \ I
'\ ' ' I : I I •
I ' ·-· I
l_
-.....;
1• \I
L-,
I
I Q11adrupedal Vertebrate
I
I • f
I
I : I
l .{
MaJ! ~- I
Cd) TYPE OF POSTURE AND VERTEBRAL COLUMN

FIG. 26 - 2 : STRUCTURAL CHANGES FROM APE TO MAN
p !I . . . ' ~ --·
355
PR/MAT SANO AN< 'F.S1'N)' 01· MAN
Tt • most common Mioc ·n • np ·s w ~r Proc:011.rnl nnd Kenyapithecus (Afr_ica) and
Dr ;0 ;;,lr<·.,;.,.
Sil'npith<· 'II.\' of Afri ·u, Asiu and Europe. The latter is generally considered as
. ~t·stor of orang-utan. Dry, pitlw,·w· is ·onsidcr ·d by some us the distant ~ncestor of t.he modern
111
~frican gr •at apes like l:him1 nnzc~ and gorilla. The other forms of surviving apes are the ora~g
ut.m of ust lndi ·s Islands and gibbons of lndin. Modern apes are comparatively small, tail-
less animuls. with long for •limbs. ,orillns ar • th· lurgcst nrnong the modern apes. Presence of
a gigantic ape. igm,wpithecu.,· is known from the fossils collected from Plio-Pleistocene rocks
of hina und lndin.
26.S ANCESTRY OF MAN

A. Paucity of humun fossils .
The gr"'at evolutionary success hns made higher primates to surpass all other mammals m
their mental developm"'nt by Miocene-Pliocene times. But the climax of this evolutionary story
was the diverg nee of hominids from some primitive apes in Oligocene-Miocene times and from
them, appearance of the modern mun in Pleistocene-Recent transitional period. We are yet to
get a complete picture of the fascinating story of human evolution mainly due to lack of adequate
fossils. The exact reason of scarcity of human fossil is difficult to predict. Human fossils may
be rare due to following reasons :
(a) Man in its primitive stage lived in plainlands, within caves near the forest. Fossils of such
organisms living in terrestrial environment are usually rare for the lack of suitable condition
for preservation.
(b) As primitive men showed a heavy reliance on animal and plant food, they required a large
territory to ~-·t suficient food and it is quite likely that they lived in small groups.
(c) Self-destruction of dead bodies may be another factor. At early stage, cannibalism might
have persisted among the early hominids. At later stage, dead bodies might have been buried
in some secret, localized places and there is every possibility that we are yet to locate many
such important sites of human cemetery.
(d) The remains of human fossils discovered in the last 20 years exceed those recovered in
the preceding I00 years and it is possible that the remains which will be recovered in the
coming years may exceed those of the last 20 years, with availability of more trained
workers, and by application of technically advunoed methods of fossil collection.
8. Trends of evolution
The process of human evolution started as early as from Oligocene with the appearance of
th~ first ancestral ape from the Old World monkey. The evolutionary development of human
bcmg cannot be treated as an event of great magnitude; it wns rnther n mutter of perfection of
so~e structures and related habits attained by their ancestral npes from the very beginning of
!heir appearance. Erect posture while sitting attained by monkeys and upes gradually led to
~mpe~e~t to perfect bi.~edol locomotion. Fr~c. fore limbs ?f upes and monkeys. used mainly
0
~ packing food
matenuls, were gruduully utilized for makang tools and weapons from natural
~bJects. by the early men. These two, coupled with continued increase of bruin size (resulting
an the increase of mental ability) und chunging food-hubit have cnused all sorts of structural
356 PALAEONTOLOGY
modifications leading to the evolution of modern Homo sapiens. The major trends of evolution
of man may be summarised as follows :
(i) Evolution of bipedalism and erect posture.
(ii) Increase of brain-size and its gradual complexity, and delaying of post-natal development.
(iii) Effective adjustment of the terrestrial habitats.
(iv) Extensive manipulation of natural objects to facilitate tool-making.
(v) Increased acquisition of meat-protein.
The (ii), (iii) and (iv) trends together lead to the development of human civilization.
(a) Trend - 1
Bipedalism, leading to upright posture is the result of an anatomical restructuring of the
pelvis and lower limbs. It is one of the major traits differentiating early human from the
common ape-human ancestor. Consistent bipedalism characterised by erect posture with
straightened-knees is found only in Man. Bipedal-walking is common among many apes but
that is a bent-knee gait in contrast to a straight-knee gait of modern man which is their normal
mode of locomotion (Fig. 26-2d). It is interesting to know how and why this was performed.
Among the different explanations the Washburn's scheme is as follows :
(a) There were major changes in human lower limbs to accompany the shift to habitual
bipedalism, not only in general skeletal proportion but also in the form of muscle and in
general limb-functioning (Fig. 26-2c). The structural changes in lower limb include an
elongation of femur bone and a reconstructing of the foot including its digits. Lower limb
bones become larger than upper (opposite to the great apes). The human foot bones and
digits have shifted from the ancestral grasping to a weight-bearing platform. The essential
points are that the body weight is borne on the great toes and that while walking foot-toes
lie parallel to one another. (Fig. 26-2a-b)
(b) The pelvis of the human and ape (Fig. 26-2c) can also be distinguished. Pelvis is essentially
composed of three bones ilium ischium and pubis. The structural basis of bipedalism
involves a reorganization of the pelvic bones. This is caused by a shortening and broadening
of ilium and then its tilting backward. This bending of ilium has allowed the trunk to be
held verticalJy. It is followed by a rotation of sacral vertebra (last vertebra at the end of
backbone) which is compensated by a S-shaped (sigmoidal) curving (lumber curve) of the
spinal cord. Along with this, there is also change of musculature of the thigh and overaJJ
structure of pelvis girdle (basin-shaped).
Theories of origin of bipedalism is speculative. Darwin argued that bipedalism arose when
our ancestors came to live somewhat Jess on trees and more on ground. Shipman ()984)
suggested that bipedalism was an adaptation to an early human pattern of scavenging meat.
Although, bipedal running is neither fast nor efficient compared to quadrupedal gait, but bipedal
gait, and bipedal walking is energetically more efficient than quadrupedal walking. This
increased efficiency was an important factor in the origin of bipedalism. It is an efficient means
of slowly covering a large areas and hence it was an appropriate adaptation for the primitive
scavanging man. Bjpedalism also increases the area of vision caused by elevation of head,
5000 Modern npc (Gorilla) ...
UJ
z
UJ
Neanderthal
~
V)
Homo erectus
ui
..J
c..
3-8
M.Y. I
I
I
I
I
I I
I ____
L f ____ ,r-- 1---,:
tJ.J L--!.-J
zUl I
8:J I
I
c.. I
I
I
I
I
I
I
I
7 My I Dryopithecus
I
26 l'n1p/i<1pi lhe, ·11.1·

M.Y.
Unknown oligoccnc}anlhropoid ancestor

I
FIG . 26-3 : CHANGE OF SKULL PAlTERN IN HOMINID EVOLUTION SHOWING PRi>BABLE

RELATION AMONG THE GROUPS
358 PALAEONTOLocy
Glacial Epochs Human Fossils Cultu~al J Tools

Age (Yrs, Oeol. Time Remains
4SOO Metal
Recent
~(r=~
10000 Neolithic
IV Interglacial UJ
,::ii::
::, ~ Metallic tools
30000 Homo sapiens ·~
~"~
f-.. -;;
..J -g,
e:
·~ "'
~
~
::,
a ...
E
:,
:J u .....
:!
V Pol;sh<d tools
A, ~
70000
u
,,I
u
C
u Homo
- C
9 neanderthalensis :I: ... "'u

·.: /!
"'
·u Ill Interglacial K. ;;; I·
C. :,
0:: Carefully chipped pointed tools
... f-, p~
~
C.
::i
-
100000
Tyrrhc Archaic
nian Homowpiens ...l
=
·u·
••'
~ •
~ a
~
~
"'
·u 0
~ Mila u
~ zcan
II Interglacial Homo erectus =6
:E "3
.!
:g :i ~ ~ Shaped hand axes and scrappclS
i ·i. w u
a ii <
Siciliar I Interglacial
•j §
<
0
1800000 -
0
Homo /,obi/is
...l
.I
G,~ (j
i
~
I
Jt Australopithocus
g. < .3
! > a..
a
!
,=i
Crude pebble tools

3IOOOOO
FIG. 26 - 4 : HUMAN FOSSILS AND CULTURAL EVOLUTION
thereby, improving the ability to locate items at a distance. Bipedalism with agile-climbing ability
made the early human-form like Australopithecus afarensis further to improve the opportunities
to exploit the environment. Bipedalism frees the hands and makes them available for manufac-
turing and carrying tools and arms. In fact, bipedalism pre-dates the oldest known tools by
more than a million year. Sinclair and other ( 1986) suggested that bipedalism developed along
with long distance migration because migratory scavangers have to access to an abundant and
constant supply of food. A human mother can, however take more care for her infant/s provided
she would be supplied with necessary food for her own and her infants. One possible food-
source could be her male-partner who in search for larger quantity of food roamed over a large
area and found bipedalism as an effective adaptation.
Most of these structural changes of foot, pelvis and spinal cord took place at the earlier
stages of human evolution.
(b) Trend - 2
There has been a trend towards increase in brain-size and its complexity during the course
of human evolution. Brain capacity of Australipithecus and H. habilis is around 300-650 cc.
For H. erectus it is ranging between 800-1200 cc and for sapients it is 1200-2000 cc. However,
the rate of increase of brain-size was not consistent throughout the geological time. Its rate of
increase was slow for the first 3 m.y. from Australopithecus to early hominid (H. habilis) (300
cc to 600 cc) but later the rate became quicker from Middle Pleistocene onwards from H. erectus
to H. sapiens stage (650 cc to 2000 cc). The increased brain size is probably related to such
factors as tool use, tool-manufacture, ability to speech, increasing ability to tackle environmental
challenges and appearance of more complex social life. It may be also related to infants slower
rate of maturation which in turns requires greater period of their parental care. This extending
period of parental care on infants has led to a more perfect family and social life by which
infants can learn more from their parents and like to associate with them even after their
maturity. All these led to a cultural development as pre-requisite for such development are
adequate memory storage and development of a perfect society.
The increase of brain size also causes some secondary changes in the overall skull pattern
such as (i) shifting of foramen magnum towards anterior, (ii) total facial plate including forehead
becomes high, upright and nearly vertical, (from low slanting type in ape), (iii) loss of heavy
eye-brow ridge, (iv) decrease of jaw-length from U-shaped in ape to parabolic in man and
consequent decrease of teeth size, (v) development of chin. (vi) enlargement of dorsal side of
skull (bunshaped) to accommodate the large-sized brain. (Fig. 26-1, 3)
Most of these above changes took place to a lower extent at different stages from
Australopithecus to modern man; some were mainly attained during sapient stage, more
precisely during the change from Homo erectus to Homo sapiens.
(c) Trend - 3
Behavioural adjustment was essential to cope with the change of habitat from ancestral
arboreal to terrestrial. Major adjustment was reflected in the social groups each of which was
matrifocal (mother-centred) genealogical unit with an old mother, her daughters and the latter's
one or more infants. Soon a division of labour came in between the two sexes where
Palac(Oco)WP-46
PALAEONTOLOGY
360
(sapient)
(Sapient)
(d) NEANDERTHAL MAN

{e) CRO-MAGNON MAN
(Australopithccinc)
(b) AUSTRALOPITHECUS
.\ (a) PROPLIOPITHECUS
FIG . 26- 5: STAGES OF EVOLUTION OF MAN
adventurous wandering males became hunters and food searchers while the females became
less mobile acting the role of infant-feeders. Concomitant with the division of labour based on
sex was increasing parental care. Tool-use and terrestriality probably placed a premium on the
development of an effective signalling system. Language acquisition might be a response to
communicate effectively in a mor~ complex life, for example, how to make a tool, where to
find food, how to trap a prey etc. Many, however have tried to link an increasing brain-size for
.
the onset of language. \\'..hich was supposedly tied in co-ordinating hunting-behaviour, food-storing
etc.
(d) Trend - 4
Extensive manipulatory behaviour of human was facilitated by his hands made freed of
locomotive function, perfect stereoscopic binocular vision, increasing brain-size (where. a lot
of information and acquired knowledge are stored) and a more effective hand-eye co-ordination.
The selective pressure for extensive environmental manipulation grew out of increasing tool-
use which was in turn related to increasing problems for survival. A major advantage of
primate's grasping hand is that it permits detachment of the object from environment. They
can pick up objects and examine them. In course of time this leads to tool-use and manufacture
of new type of tools.
(e) Trend - 5
The exact time of ' meat' acquisition by man is not known but probably it occurred in Middle
Pleistocene, at the last stage of human evolution. Consistent meat-eating had caused changes
in dentition and jaw-masculature. Once meat became the basic food in the diet, they had to
develop means whereby it could be efficiently obtained, butchered, cooked and then shared
the food to other: members of the family. Sometimes analysis of upper-surface of tooth can
tell the role of certain food in an animal's diet. Tooth· surface of Australopithcus robustus
suggests that it was basically a vegetarian. A shift in the enamel-wear pattern on the tooth-
surface of H. erectus suggests that a large amount of grit was incorporated in their diet which
came from root or tuber and rare-meat eating, but there is no sign of bone-crushing. According
to Shipman the evidence of cut-marks, tooth-wear and bipedalism together with a knowledge
of scavenger-adaptation is consistent with the hypothesis that about 2 m.y. ago men were
scavengers rather than accomplished hunters. Hunting as a way of life appeared about t .5 m.y
ago within H. erectus.
26.6 STAGES OF EVOLUTION OF MAN (Fig. 26-5,6)

From the very early days, a close m~rph~logica~ resemblance between man and anthropoid
apes f~cinated more than one natural sc1ent1sts. Keith proposed a plausible explanation of how
brachiating (arm-swinging) apes could have evolved into bipedal man. The human-chimpanzee-
gorilla relationship is closest not only in terms of morphology but also in DNA-hybridization
and. nature of protein-c~ains in amino acids so that it can be inferred th~t human and great
Afncan apes are genetically close to each other and hence they are placed into a common
group 'Hominoidea'. But the question is wheather man was evolved directly from ape or not.
The answer is not easy, especially in the absense of fossil of any such transitional form between
ape and man. It is also possible that man and ape arose independently from some common
ape-like ancestor. No doubt, many aspects of history of the evolution of man remain stiJJ
. .....
-,,........-
;!__... - .
.1
- J ·. A
• ".1 · ,, :_ .. .... ' ·
PALAEONTOLOGY
362
· d th · ture has been modified at different times with the discovery of new
controvers1a1 an e p1c . . . . .
. d f ·1 ·tes At present, it is possible. to subd1v1de
foss1 1s an new oss1 s s1 . .
the enttre history mto the
... . . .
following stages : (i) Pre-Man stage, (ii) Austral_op1thecme stage, (111) Hab1lme sta_ge, _(iv)
Pithecanthropine stage and (v) Sapient stage. A bnef account of each of these stages 1s given
below:
A. Pre-Man Stage
The earliest common ancestor of ape and man, arising from some Old World monkey stocks
probably appeared about 40 m.y. ago in Eocene of Burma. The we~l-k?own forms are
Amphipithecus and Panduangia (D. F. 2133). Both of them show a combmat1on of lower and
higher primate traits with latter having an approximate size of the small ground ape gibbon.
The next group of fossils have been collected form Fayum of Egypt which are about
35 m.y. old (Early Oligocene) represented by Parapithecus and Propliopithecus. There is
somewhat doubt about ape-affinity of Parapithecus. Propliopithecus, represented by a short and
deep lower jaw fossil, containing cheek teeth with five low cusps (D. F. 2133), is considered
as the primitive-most ape of Africa. A slightly younger form is Aegyptopithecus, reported from
the same locality (age 34-33 m.y.) represented by almost a complete skull. Aegyptopithecus,
evidently of a size of modem gibbon, had expanded cranium, forwardly directed eyes, and a
jaw with a continuous series of teeth showing two spatulate incisors, one large canine, and five
low-crowned cheek teeth with low cusps. Morphologically and chronologically such an ape could
be derived from Propliopithecus and could be the ancestor to the extinct Miocene apes of Africa,
the 'dryopithecines' . ·
~ng ~he Mi°':ene times~ hominoids _were dominantly represented by one fossil group, the
dryop1thecmes, ~h1ch were m ~um, led do~inantly by three genera viz; Limnopithecus and
Proconsul of Afnca _and Dryop,thecus of Afnca and Euresia. They were very much like apes
~d showed clear d~vergenc.e from monkey. Limnopithecus could ~ the ancestor of modem
gibbon and Proconsul and might have led to Dryopithecus and other M" · D ·h
is the most widely distributed form found in Africa Europe and A . iocde~e ape~d· rydop,t ~cbuJs
. . , s1a an 1s cons1 ere poss1 e
ancestor ofall other apes hvmg and extinct. A slightly younger M" v ·h
of Africa d s· · h fA · . iocene apes are nenyap,t ecus
l h.ai;. h;ri" ecus o s1a. S1vapithecus exhibited specialized molar often calied Y-5
mo ar w 1c a 1ve cusps separated by a deep Y-shaped groove.
For many years it was suggested that one of th M' ·
to the earliest hominid This view b d ese iocene apes was probably the ancestor
. was ase on the doubtful t t
Ramapithecus, first described by Lewis f:0 L s· . . s a us o f t he Phocene
. &
aorm
form refers to a fragmental part of m ~~bl ower iw~hk (Phocene) of India. Originally, the
(parabolic}. Chronologically it was foun::'~cue s:;;rficially res~~bli?g a human jaw-pattern
and Australopithecus to be a human an t JY g a proper position m between Dryopithecus
trait of this form and have called f:or pcles ?r. . an~ others have however, stated the non-hominid
f
~m. Indaa-Pakastan
. . acmg 1t with the ge
and China. The fate of Ra .
s· · h
nus 1vap1t. e_cus, an undoubted ape
watha~ the ge~us Sivapithecus was final with ~~?ecus as a non-h?mm1d ape and its inclusion
of thas form, including a complete skull th asc_ov~ry of multiple of new f!)ssil-materials
undoubtedly ape-like. at c Iearly mdacate that the jaws of Ramapithecus is
. Primat~ fossils of Gigantopithec (F' 2 .
Gcgantopuhecus have been called us ·~·. 6-6) have had a colourful history. At different times
everyt mg from an ape to a giant human. Sometimes it was
363
designated as the ancestor of illusive yeti or snow-man; at times it was linked to China's illusive
wild h~iry-man. First menti~n of Gigantopithecus was made of by Koeningswald in 1935 on
the bas.is of some teeth fossils found in a Chinese drug store. These bones were collected by
the Chmese as dragon-bones for the use of herbal medicines. Since then, more fossils have
been collected fr~m India and also from China. At present, the genus is represented by a large
number of lower Jaws and thousands of isolated teeth fossils. It appears that Giagantopithecus
might have a weight of 400-600 lbs and it stood about 6 feet tall. They range in age from
9-5 m.y. in India but are found in rocks as recent as 500,000 years in China where these
materials have been collected from the rocks yielding bones of giant panda and fossils of Homo
erectus. The Indian version of Gigantopithecus belonging to the species Gigantopithecus
bilaspuransis was collected from Bilaspur, Simla hills, Himachal Pradesh in 1968. Later in 1984
fossils of Gigantopitecus have been also reported from North Vietnam.
Teeth and mandibles of Gigantopithecus are very large. Teeth are diagnostic as they are
largely composed of enamel, possibly an adaptation to heavy chewing stress. lolly's seed-eating
hypothesis has been called upon to explain Gigantopithecus trait. This seed-eating hypothesis
of Jolly is ·however based on feeding mechanism, behaviour and dental pattern of an African
monkey (Theropithecus gelada) . The existence of similar dental trait in Gigantopithecus
suggests that they fed on dietary items similar to those of gelada which according to Jolly were
composed of large quantities of tough morsel-like grasses, seeds, stems and rhizomes mixed
with grits. Such foods, a~ailable in the open savannas, need a powerful ability of chewing with
strong premolars and molars.
In formulating the seed-eating hypothesis, Jolly also noted the existence of a number of
morphological and functional parallelism between early human and gelada (and hence
Gigantopithecus) as both of them exhibited the following similarities :
(i) open grassland and open country habitats,
(ii) reduced incisors and canines placed more vertically,
(iii) crowding of molars and sign of wear and tear in the molar teeth,
(iv) both with robust and thick mandible,
(v) maximization of chewing ability and more efficiency of chewing muscles.
Not all agree with the lolly's interpretation of ~iving excess stress on dietary adaptation for
understanding the major factors of human evolution. Wolpoff ( 1980) noted that all powerful
chewing mammals are not nece,ssarily seed-eaters and such dental traits are of no help in
understanding other human adaptations like bipedalism and tool-use.
Gigantopithecus probably evolved in Indi~ a! first an~ then .spread !o north and east Asia. It
seems to be a long-lived side-branch of hommo.'d evolu~1on. G1gan~o~11he~us and early human.
whatever the final resolution may be about their evolutionary affihat1on. 1t may be concluded
that Gigantopithecus may represent a tread towards non-arboreal ape.
B. Australopithecine stage (Fig. 26-Sb)

(a) Fossil record . .
Most of the people believed that the early human population is. record~d w~th,". the
· · · · - - - .. "' Th"' fir.1:;t fossil member of this group was found m I 925 m a limestone
-~
PALAEONTOLOGY
364
. . ) It was a skull of child showing upper and lower
quarry of Tuang, South Afnca (Thang m;n h·. Tuang child as Australopithecus africanus and
ja~ wi~h all tee~h. Raymond Dart r~ari;;rmt ~; human. Three species of Australopithecus have
mamtamed-t~at it represents an ea ;d South African Australopitheus africanus and Austra-
been recogm~ed. so far. The East h African robust form Australopithecus robustus. Of these,
lopithecus bo,s~, and another Sou~bl l d t h man line and the other two are now considered
th former one is smaller and poss1 y e o u d 'b d f
e . (W lk· t 1 l986) Another form of Australopithecus was escn e rom
as the extmct apes a er e a · · ·h ,I'. · It ·
Tanzania East Afnca · and Hadar, Eh' · w h'ich is named as Australopit ecus a;arens,s.
t iopia f h f ·11s
' f ·1 · k ed 'Lucy' It shows some undisputable features o t e amt y
a skeleton o a young g1r ~,c nam 1 . . 1 cavity (450 cc) and sexual dimorphism.
hominidae regarding its size, denta pattern, crama . l .h,
Johansen and White (1979) considered it as a basal human stock from which Austra optt e.cus
africanus and Australopithecus robustus diverged in one di~ecti~il and eventually became extm~t
while Homo habilis diverged from it in the main line, to give nse to Homo ~r~ct~s. A a[arens,s
is thus considered as the oldest undisputable evidence of existence of hommt~s ~n ~fnca. The
major traits of A. afarensis are : considerable size-variation a~ong adu~ts md1catl~g _sexual
dimorphism; primitive cranial and post-cranial structure; bones thick, heavily ~u~c.led, btp~~al,
arms slightly larger than the modern man; brain capacity 400-500 cc; most pnm1t1ve dentition
with comparatively larger molars and premolars and a more sharp canine, no evidence ~f t~?l-
manufacturing; age between 2.8 to 3.8 m.y. White et al. (1994) discovered a still more pn~t~ve
hominid fossil named Ardipithecus ramidus dated about 4.4 m.y. Its teeth were more hommme
than found in any of the great apes. Its age also coincides with the presumed data of splitting
between ape and man about 5 m.y. ago. Subsequently, another primitive form of hominid
Australopithecus anamensis (4.1-3.9 m.y.) has been reported by Leaky et al. ( 1995), which is
considered to be an intermediate form between Ardipithecus and A. afarensis.
Downfall of the argument that the members of hominidae might be found among some Miocene
hominoids began with the wide acceptance of an idea of late divergence of human and ape which
is substantiated with fossil evidences from China, Pakistan and Africa. Middle Miocene hominoid
group known as Sivapithecus which was splitted off from Lower Miocene Dryopithecus may be
the ancestors to organg utan rather than to gorrila, chimpanzee and man. Dental pattern of
Sivapithecus and that of the first hominid Australopithecus is not indicative of an descendent-
ancestral relationship but is the result of parallelism. More probable form in the human-ape line
could be Kenyapithecus (14 m.y.) from Kenya. Thus the gap between Miocene and Pliestocene
as regard hominid fossils is still there and we are yet to get a 'missing link' between the ape and
the man. Important sites of some early hominid fossils are shown in Table-17 A.
(b) Llfeways
Despite the fact that most have argued that the earliest tool user was Homo habilis, there is
a possibility that at least one group of australopithecines knew the use of tools. The stone tools
appeared first about 2 m.y. ag? roughly contemporaneous with the first appearance of the genus
Ho"w or. some late austra!o~ithecines. Napier ( 1962) suggested two types of use of fingers :
pow~r gnp (the type of gnp an holding smaller objects) und precisio11 grip (the type of grip in ·
holding a hammer). Presence of power grip indicates their ability to manufacture · stone tools
which are found lo ~ present associated with the specimens of some Attrtrc,/o••ithecur and Homo
habilis of Olduvai Gorge. · ' ·
PRIMATES AND ANCESTRY OF MAN ~65
TABLE-17A
SITES OF AUSTRALOPITHECINE AND EARLY HOMINID FOSS1LS
South African sites :
Sterkfontein Australopithecus africanus/ robustus
Makapansgat Do
Thang Do
Swarkrans Homo habilis I Homo erectus
Kromdraal Australopithecus robustus I boisei
East African sites :
Olduvai gorge. Tanzania Australopithecus boisei. Homo habilis.
Australopithecus africanus I afarensis (3.7-3.5 m.y.)
Hader Australopithecus afarensis (2.5-3 m.y.)
Ethiopia Omo valley Australopithecus africanus I boisei
Middle-Awash A. afarensis, Homo habilis
Ardipithecus ramidus (4.4 m.y.)
Lake Tarkana Austra.lopithecus anamensis (4.1-3.9 m.y.)
Australopithecus africanus
Australopithecus boisei
Homo habilis
Lake Baringo Do
There are variations in tool manufacturing technique and the types of raw materials used.
There are two basic categories of tools. The core tools and the flake tools. To make a core
tool, chips were knocked off from a lump of stone until the lump attained the desired shape
and size. A flake tool is a chip struck off from a core. A chopper is a tool typically found in
Lower Pliestocene sites. It is a smooth rounded cobble stone or block to which was given rough
cutting edges by knocking flakes from all sides.
Australopithecines were probably hunters but surely scavengers. Food materials were
predominantly vegetables consisting of herbaceous plants. There may be a relationship of this
type of food habit with such tools like chopper. scraper. flake knife etc. Large cutting tools
may be associated with meat eating and other hunting pursuits. Again hunting in open savanas
means a cohesive social organisation, co-ordination and communication system, but not
necessarily speaking ability. They might had taken such techniques as relay races to wear down
the prey by driving it into the members lying in ambush, encircling and attacking it from many
directions. Lacking technically advanced toors early man must have hunted from ambush or
driving the prey into some low land, or within deep mud. Many others argue that these early
humans were poor hunters due to lack of suitable tools and they relied mainly on scavenging
366
PALAEONTOLOGY
to ~btain meat, skin and other products. Australopithecines probably shared some of the
ava1lable food among themselves. The male probably scavenged and hunted animals and
obtained the choice pieces. The remainders were divided among the olds, infants, females and
the youngs. Some of the social rules regulating human behaviour stemmed from this food
sharing and this may be one of the basis selecting for language.
(c) Morphology
The average body weight of australopithecines was about 66 lbs and for robustus it might
be 160 lbs. In terms of height it ranged around 4-5 feet. Size variation may be related either to
sexual dimorphism or to the ontogenic stages. Australopithecine's dental trait is complex, not
conforming with that of higher hominids. Upper incisors become smaller like ours but the lower
incisors and canines become much larger than those of modern man (but smaJJer than those of
the apes); the premolars and the molars indicate a primitive human trait. The changes that follow
in astralopithecines are as follows :
(i) evolution of visual sense with reduction of nasal area;
(ii) assumption of an erect posture and bipedalism;
(iii) recession of chewing apparatus with reduction of power and extend of tooth-grinding
surface leading to restructuring of jaw, face and skull.
Largely because of shape and dimension of skull, scientists were at first inclined to believe
that australopithecines were apes alike to gorrila-chimpanzee rather than early member of
hominid. The relatively smaller brain; combined with massive and projecting jaws gave them
a superficial ape-like appearance, but their brain was unlike from that of an ape in having the
following features-
(i) expansion of posterior area (controlling the auditory, somatic and visual zones
concerning hearing, touch and vision);
(ii) greater complexity of the frontal lobe of the skull (control area of speech);
(iii) expansion of temporal Jobe (control area of visual analysis and auditory memory).
The pelvic structure of ape and human is significantly different in response to their type of
locomotion for different mode of life. Human pelvis is modified to accept an upright bipedal
posture and in this respect australopithecine's pelvis ~tro~~ly differs from that of an ape a~d
approaches closely to that of a modern man. One s1gmf1cant change in pelvic-structure 1s
broadening of ilium. Besides this, the two lower leg bones tibia and fibula indicate the
adaptation of the body for bipedalism.
There is controversy about the two morphologic varieties of australipithecines. A. africanus
is a smaller, lightly built gracile form whereas A. robustus-boisei represent a larger, robust
form with coarse skeletal structures. Many have suggested that there existed habitual as well
as anatomical difference between these two groups of aurstalopithecines and they should be
viewed as two separate genera. Robinson suggested that the gracile fon.a should be included
within the genus Australopithecus (A. africanus) and the robust form which was slightly younger
should be treated as a separate ape-genus Paranthropus. The former was a generalized form
leading to the advanced human while the robust form was a specialized offshoot that changed
little during its evolutionary history and finally became extinct.
----
(d) Olduvai gorge . . .

The Olduvai gorge of East Africa (Tanzania) is one of the best sites of the world y1eld1~g
human fossils. Most of the early human fossils belonging to australopithecines as well as fossils
of H. habilis and H. erectus have been collected from this area. This site was investigated for
years by Prof. Leaky and his family members. The Olduvai g~rge is about ?O m deep cut by
water through a series of old lake deposit of volcanic dust which covers a _t1m~ ~pan _of about
2 m.y. Lithostratigraphically, this horizontal sequence of Pleistocene rock~ 1s d1v1ded mto fo~r
beds (From Bed I - IV). Succession and important fossils of the Olduvai gorge are shown m
Table-17B. ·
TABLE-17B
SUCCESSION AND IMPORTANT FOSSILS OF OLDUVAI GORGE
BED AGE FOSSILS
Upper bed 100,000 - 400,000 yrs. Homo sapiens I tools
Bed (iv) 400,000 - 700,000 yrs. Homo erectus (Acheulian hand axe, cleevers)
Bed (iii) 700,000 yrs. - 1.2 m.y. Few stone-artifacts
Bed (ii) 1.2 - 1.65 m.y. Homo erectus I Homo habilis
Bed (i) 1.65 - 1.8 m.y. Australopithecus boisei I Homo habilis
Volcanics 1.9 m.y. (Sterile) .
C. Habiline stage (Fig. 26-6)
Habiline stage is represented by the fossil Homo habilis (litterally meaning 'handy man')
collected by Leaky from Bed-I of the Olduvai gorge from a site positioned slightly lower than
that of A. robustus I boisei. This form appears to be more advanced than australopithecines and
is referred to the genus Homo suggesting that it was the first record of true hominid. The age
of the fossil is dated as 2-1.8 m.y. The fossils include two parietal bones, one jaw, one clavicle,
two fragmental hands, and almost a complete leg bone. More materials have been collected
from the Omo valley and Lake Tarkana, East Africa and also from Swarkran, South Africa.
Some research workers maintain that Homo habilis is nothing other than a gracile
australop.ithecine. ~hey consider H. ltabilis as an advanced mem~f'J of the gracile
Australoptthecus on its way to produce the next advanced member of human that is Homo
e~ectus : But careful observation of several fossil materials has pointed out the following traits
diagnostic of Homo habilis.
(a) thinner skull with brain capacity 600-670 cc, about 15% increase over the australo-
pithecines;
(b) although, facial pattern and mandible still exhibit adaptation to powerful chewing, the
dentition shows canine smaller than that of australopithecines and cheek teeth elongated
posteriosly;
(c) an almost erect and bipedal motion indicated by the leg bones;
(d) most possibly makers of primitive tools which includes small chopping tools und very
sharp flakes, called 'Oldowan tools'.
Patae(Geo)WP-47
/
PALAEONTOLOGY
368
Absolute Geol.
Culture
Time Time .
,, C
-
0,_
1
..- 2500 B.C. --
r
G)
N
...J C
._
~~
0
,_
,_ m z
MODERN i- 3500 B.C. - w
GREAT APES
G)
Q. u
MONKEYS Q. w
I (Gorilla) g cc
MODERN 4500 B.C. -
'' GROUND APES u
(Gibbon) ·~ ._:i: >,
,_
:J
0
w
-
a.
G)
0
-10,000 B.C. -
------------~-- - -
Extinct
- H. sapitm
z
C
as
·.::::
CD
H. n,andtr tl1al1nsis : Ch
~
I
'' Archaic H•.rnpitns

0
w
' ', u ::\? 80,000 yrs. z
w
~...J
/ C
,_ I as
-.... -'
(.)
1.8 My
H. crtetus 0
:,
G) ~
Cl)
s::.
w
<
u w
II. habilts ~ ...J
a..
\
~
A. robust11s
A. a/rica11as
' C
al
,__
3 .2 My -
' ' ~ 3:
... ,, /
J.--•-•------a++i~
"C
: ..... .' 0
-
• Australopithteus a/artnsis
t--~---- - --- - - _1 _ - - - - - - - - - ; ·- - - - - - -
3 .8 My -
I I W
I
I
Ardipithteus anamtnsis ffi
I (.)
Gigantopithec11s 0
A.rdipithtcus ramldus :J
~-~ - --
I . - - - . - - - - 7'- - - - - - -
, - - - - ~- - - -
: - - - - - - - - - -
a..
I I · 5.5 My
I I
I
I
Sivapithuus Ktnyapithtcus wicktrii
'II ,,, /
,,, .,
I
I 7.0 My w
I /
,, ;
z
w
\
\ I
I
. 0
' I 0
- Dryopltl11cu1 ~
15 My
' .... , ....
Umnopltlt1cu1 -- - - - ',
- - - - - - --. Proconsul
.. ----- - -------- -- -- - ----------1- -- - - -- - - - - - - -
A.1gyptopitl11cus 30-22 My
'I w
Propllopllh1c111
I
zw
35-30 My 0
0
I
'
Paraplth1cu1 CJ
:J
MONKEYS
--- --- -- '
I
I
40-35 My
0
I
Unknown anthropoid ancestor
FIG. 26-6 : AN OUTLINE OF PHYLOG ENY OF MAN
D. Pithecanthropine stage (Fig. 26-Sc)
(a) Stratigraphic distribution
The representative form of this stage is Homo erectus. Though, it is the only representative
form of this stage, the species is found widely distributed in Middle Pleistocene times and
reported from Asia, Africa and Europe. In 1891 Eugene Dubois, a Dutch army doctor discovered
' a skull cap from an excavation near the Solo river in Java. The specimen was identified as an
early human form and named as Pithecanthropus erectus, (Java man), an apeman who could
walk upright. More such specimens were discovered from near Peking, China (Peking man)
named Sinanthropus pekinensis. Similar type of fossils have been also reported from South
Africa (Swarkrans), Olduvai gorge (Bed-II and Bed-IV), East Africa, Morrocco and Algeria,
North Africa. A close analysis of all these specimens and the associated fossil sites indicates
that they had much larger brain and mor~ advanced culture and hence they are grouped into a
single species Homo erectus. African specimens are considered slightly older and the total range
of the form appears to be 1.8 to 1.4 m.y.
(b) Major traits

Homo erectus had a cranial capacity ranging between 700-1200 cc (average 1000 cc). There
was much flattening of skull vault with a sagittal ridge running across the mid line of the· skull
and the skull bones were somewhat thick. The skull appears to be more primitive compared
to modem man where the width along skull vault is much higher due to reduction of the mass
of chewing muscles. There w·a s also a heavy and large eyebrow ridge above the eye orbit and
behind this was a postorbital constriction. Nasal bones were broad, flat and the jaw was more
prognathic than that of modern man. Mandible was also heavily constructed lacking a chin.
Teeth were larger than that of H. sapiens; canines was sharp and projecting; cheek teeth were
cusped with enamel wrinkles. Limbs were essentially like modem man representing a true erect,
bipedal posture.
Although there are a number of traits separating H. erectus from H. sapiens, there is a general
consensus that former is the ancestor to the latter. The idea is based on the following facts.
(i) The morphological traits of H. erec_tus conform very well with the theoretical postulation
for being an intermediate stage in the evolution of H. sapiens.
(ii) The existence of H. erectus from the earlier part of Pleistocene upto the later Middle
Pleistocene illustrates a graded series of morphological changes that led H. erectus to
H. sapiens.
(c) Life ways and culture

The control of fire is one of the most momentous discoveries in the history of human
evolution. The earliest evidence showing use of fire by H. erectus has been found from some
East African sites of these human fossils. The Swarkrans site exhibits the presence of burnt
clay and burnt bones which is dated as 1.5-1.4 m.y. The materials are associated with skeletons
of H. erectus. Natural sources of fire were possibly the contemporaneous volcanoes. the fire
associated with natural gas/oil and the natural forest fire. By using fire H. erectus men were
not only able to remove the darkness of night and caves. but also protected themselves from
cold climate, as that was a periods of gradual initiation of Pleistocene glaciation. Discovery of
fire proved highly effective for subsequent ice-age man (neanderthals) as fire was their main
--
370
PALAEONTOLOcy
we-apon against the glacial climate of Pleistocene. Use of fire made human a regular cave.
dwellers. Fire was also possibly used in burning food, especially meat, that had also caused a
change of their teeth characters. The European fossil Homo heidelbergensis (780000-500000 yr.)
which are much younger that H. erectus could be an advanced H. erectus but more probably
are representating some archaic H. sapiens in Europe.
Tools associated with Homo erectus belong to Acheulian culture which seem to be more
advanced type over the preceding Oldowan tools associated with H. habilis. Acheulian tools
are made from a wide range of raw materials. In contrast to all-purpose Oldowan tools, different
types of tools were prepared for different purposes showing a versatile range of cutting,
scrapping, piercing, chopping and pounding tools. The precision, evident in the manufacture of
acheulian handaxe apparently resulted from a new method of flaking, the soft hammer technique
in which stone flakes were detached off with a wood, bone or an antler-made hammer. Use of
such soft hammer allowed more control over the length, width, thickness and shape of the final
products. In fossil sites, several kinds of activities may be noted such as :
(i) several unmodified rock-wastes resulting from tool-manufacture;
(ii) butchering sites where disarticulated animal bones found associated with cutting tools;
(iii) sites of workshops with varying types of rocks lying, adjacent to raw materials.
H. erectus was mainly a cave dwellers or might have Jived within built-shelter. They were
big-game hunters and were roughly contemporaneous with the first and second glacial epoch
(Mindel) in Europe. Culturally they belonged to Early Palaeolithic type.
E. Sapient stage (Fig. 26-5d, e)

(a) Neanderthals and Homo sapiens
Sapient stage, the final phase of human evolution is represented by two separate forms,
neanderthaJs and modern man. There is however a lot of confusion about the origin of modem
man and its relation with neanderthals. Some workers believe that neanderthals is an earlier
subspecies of H. sapiens (H. sapiens neanderthalensis) from which evolved the subspecies of
modem man (H. sapiens sapiens). Many other have regarded neanderthals, a distinct species
(H. neandertha/ensis) and would place them as an offshoot in the evolution-line of modem
man suggesting that there is no ancestral-descendant relationship between them. Moreover. there
is no evidence from any part of the world. showing morphological transition from neanderthals
to modern man and there was possibly no sufficient time for such transformation. Co-existence
of both these forms at least for sometimes is evident from fossils. The former group of workers
however, have argued that in Europe there were two variants of neanderthals. one more gracile
forms of East and Central Europe and Middle East. often designated as 'progressive
neandertl,als'. and the others were robust-built 'classical nea11dertl1als' of Western Europe.
They have suggested that progressive neanderthuls probably represent the intermediate forms
between earlier neanderthals and later H. sapie11s and the 'classical' forms are considered outside
our direct evolutionary lineage.
If _n~derthals are kept outside the main line of human evolution, then H. sapiens was surely
~~•tatav~ of earlier H. erectttJ'. Recent findings of hominid fossils from some parts of Africa,
1
e East and Europe t.irongly suggest the presence of some forms in a time span between
• J>
1()()()()0-300000 year showing an intermediate morphological traits, some belonging to H. erectus

and some to H. sapien-s and such transitionals forms are expected in case H. erectus gave rise
to H. sapiens. These forms are designated by some as early H. sapiens or archaic H. sapiens.
Most researchers now agree that Homo sapiens evolved from a H. erectus lineage. Now it is
considered that modem H. sapiens appeared first in Africa and Middle East, sometimes between
100000-120000 year ago and migrated to Europe (40,000-30,000 years ago) prior to which
Europe was inhabited by neanderthals (120000-35000 years ago). They coexisted for sometime
after which neanderthals were replaced by the more advanced and efficient H. spaiens. Even
some authors have not ruled out the possibility of hybridization between them resulting in the
evolution of H. sapiens sapiens of the prese·nt day.
(b) Archaic Homo sapiens

Fossils of early Homo sapiens have been discovered from Africa (Kabwe; Zambia, Rodesia),
Asia (Solo river, Indonesia) and Europe (Swanscoinbe, London; Steinheim, Germany; Greece
and France). The rodesian specimen is named as H. sapiens rodesiensis while the Indonesian
form is named as H. sapiens soloensis. The fossils include several skulls (mostly broken), limb
bones, jaws and teeth. The cranial capacity of these forms is ranging between 1100-1400 cc,
average 1166 cc. For H. erectus this value is 800-1200 cc. So these specimens exhibit about
11 % increase of brain size over that of H. erectus. These fossils also show gracility of leg-
bones. Structures, supporting the facial musculature are somewhat similar to those of modem
man. Existence of chin is found in Arago specimens. On the other hand, they retained such
primitive characters as strong eye-brow ridge, and thick cranial wall. The time span of these
forms is calculated as 300000-120000 years before the present.
Archaic H. sapiens possibly lived in ·caves and built dwellings. Lumley (I 969, 1973)
discovered from Nice, France even some oval huts which might have housed 10-20 people
and each hut is characterised by a central hearth and large quantity of charcoal and ash.
Occurrence of large quantity of bones of severai vertebrates (bird, turtles, several iarge mammals)
and invertebrates shells (mussels and oysters) may indicate that the inhabitants were hunters.
The presence of fish bones indicates that inhabitants also knew the art of fishing. Such big-
game hunting, fishing and large scale cooking are never found among any pre-existing H. erectus
group. Some impressions of wooden bowls on sand dunes, may indicate the earliest trace of
container (wooden). Lumps of red ochre in nearby areas also indicate earliest use of colour for
decoration.
Evidence of cultural advance is also known from the occurrence of tools and tool making
areas. More than 100000 artifacts were recovered from two sites of France. Overall the stone
implements associated with these skeletal remains have been a~signed to Middle Acheulian
handaxe tradition.
(c) Neanderthal Man (Fig. 26-5d)

Neanderthals were roughly contemporaneous with the third glacial, third interglacial and
fourth glacial epoch of Europe and can truly be referred to as 'Ice-age men'. They were mainly
inhabitants of Europe, although a few forms are known to occur in the Middle East, The chief
fossil-sites of Europe are : Belgium (La Naule&Je, Spycave) France (La Chapellu and Le
Montiere), Germany (Neanderthal Valley near DusseldroO, Yogoslovia (Krapina), Hungary
PALAEONTOLOGY
372
irceo) The general time span of these forms is perhaps 12000().
(Subalyuk) and Ita1Y (Mo nte C ·
35000 years.
Although, there is a huge collection of fossils and sto~e tools of neand_erthals from different
parts of Europe yet the problem regarding thi~ group 1s the m~st nettmg one. Ne~n.denhal
population lived in a variety of habitats and ~nviron~~nts for which they ha~ to take different
adaptations biologically and culturally with mmor vanatt?ns. Mos~ anthropologists have accepted
the following traits for neande~hals : They had a cramal capac1~y of 152_4- I~~ cc for. m_ales
and 1270-1425 cc for females. Although, the cranial capacity of the form 1s w1thm the hm1t of
modem man H. sapiens sapiens, it was archaic-looking with a roh~st bun-shaped post~ranial
part. It showed a skull, with a flat crown surface, large eyebrow ndge, n~sal prognath1sm, a
large robust jaw without chin and broader anterior teeth. Their average height was 1.5 meter.
Neanderthals were big game hunters and animal-meat provided them a large part of their
caloric intake. Wooly mammoth and wooly Rhinoceros were successfully hunted as the caves
inhabited by them were found to contain broken skeletons of these animals. Fire was their chief
weapon in their fight against the severe cold climate of that period. They used to burn wood
and animal bones and also used animal furs and hides to cover their bodies. Climate also forced
them to adopt the techniques of storing food materials.
The cultural assemblage corresponding to neanderthals is known as Mousterian and the type
of tools are the main clues about their possible life habit. Mousterian culture gradually replaced
the earlier Acheulian tradition of H. erectus'. The most notable feature of this new culture is
the increase of varieties of tools indicating their specialized use. More than 63 types of tools
of neanderthals have been recorded. Tool preparation involved striking off thin and fine flakes
from a pre-determined size and shape of a core with a greater precision. This is called 'core
levallois technique'. Each tool suggested a special purpose rather than a multipurpose use.
Broadly the cultural grade was Middle to Upper Palaeolithic.
Neanderthals present some earliest and clearest evidence of mortuary practice. Several burials
have been discovered in Belgium caves and from excavation of hillside at La Chapelle, France.
But the largest of this type is known from near La Ferrssie, France, described as 'neanderthal
cemetery'.
(d) Appearance of Homo sapiens sapiens

~t is suggested by most o~ the worke~s that H. sapiens appeared about 120000 year ago in
Afnca from some H. erectus hneage passmg through the transitional forms archaic Homo sapiens
(range: 300000-120000 m.y.):From Africa, they migrated to Europe and North America about
40,000 yea~ a~o. But why they took so long time to enter Europe ? One possible reason may
be ~he glaciation that had affected ~everely Europe and North America during a greater part of
Ple1~tocene that prevented them to mvade these continents earlier. Or, they might have entered
earher but we yet to get their fossil record.
The _evolution~ development of H. sapiens sapiens can be best understood in terms of
worldwide changes 10 selection acting on the already differentiated archaic H. sapiens. Selection
changed the gene-pole of the existing archaic H. sapiens in the direction necessary to account
for the appearance of modem subspecies of this genus. Almost all the characteristic features
373
of the later population appeared in lower frequency within the earlier H. sapiens population.
The earlier forms of H. sapiens can be distinguished from archaic forms by a size-reduction of
anterior teeth, reduction of eyebrow ridge, increase of cranial height and appearance of a perfect
chin. However, these changes occurred within the different groups at different times and at
different rates.
One of the most familiar forms of H. sapiens in Europe was found in 1868 in the Cro-
Magnon shelter located in a limestone cliff at the French village Les Eyzies. Six skeletons were
found buried deep within the rocks at the shelter. Three of males, two females and one of an
infant. Reconstruction of the skeletons has led to a man of 5'4" -6' height, with a broad face,
high forehead, projecting chin and a cranial capacity of nearly 1600 cc. These were named as
'Cro-Magnon man' (Fig. 26-Se). These forms were probably survived here upto the end of
last glaciation. Later numerous other fossil skeletons have been recovered from different parts
of the world and the more important sites are given in Table-18.
Homo sapiens sapiens has the following general morphologic characters : it has larger
cranium compared to its height, (range of cranial capacity 1500-2200 cc), upright forehead, an
elongated nasal opening, squarish orbit, rounded occipital region, well-developed chin, and
anterior teeth much smaller than the earlier forms. It also has feebly developed eyebrow-ridge
(which is never continuous), higher skull vault and thinner skull bones compared to the earlier
forms. There is an overall reduction of bone thickness and muscles making the form very much
gracile. The average cranial capacity of neanderthals is less than that of H. sapiens but in a
few cases brain-size of the former is found greater than the average value of the modern man.
This has suggested that brain-size alone is not significant. What matters is the brain's complexity
and of course, the development of some particular area of the brain, such as the frontal lobe,
the fun· development of which has caused change of the facial configuration of modern man.
TABLE-18
SITES OF FOSSILS OF HOMO SAPIENS
Country Fossil sites Approx. age (years) of fossils
Africa .
Omo valley, Ethiopia 13000
Byas, Laetoli, Tanzania 12000
Asia China, Java 20,000-10,000
Europe Yugoslovia 40,000-30,000

Les Eyzies, France, Italy Do
Australia Lake Mungo, Kosipe 25,000-35,000

Kow swamp 14,000-9,500
Tasmania Fraser cave 20,000-15,000
N. America California 30,000-32,000
Brazil 32,0QO
S. America
PALAEONTOLOGY
374
. · were big-game hunters with improved hunting techniques
I01.f 11 Homo sapiens sapmes . . .
,a Y . t f weapons such as spears, jevelmes, harpoons, clubs, stone m1ss1les,
and tools A great vane y o . 1· d . . h
a~ s throwing sticks, snares and pitfalls were used for k1I mg ~n trapping t e games.
boomer g ' . . were chased off the cliff. Some cave-drawings have preserved the
Herds of greganous amma1s . . . .
images of different hunting weapons and tool-patterns, R~indeer and s1m1 1ar other _animals
. d no t only the meet but also such materials like clothings,
supp11e . . tents, threads etc. Their bones
were used for making harpoons, needles and other soph1st1cated tools.
People inhabited a variety of dwellings. Rock shelters and large caves were common!~ used
where they lived within huts and tents. Cave-inside was heated by w~d and _bone-fire. In
Central and Western Europe there are remains of some permanent dwellmg which are made
up of wood or large animal's skeletons covered with animals skin and hid_es . They pre~erred to
live in large social organization. Possibly, they had also developed. nt~al of coming ~nd
remaining together. Mating and kinship rule might have created an intncate and cohesive
relationship among the large number of people.
(e) Cultural development (Fig. 26-4, 6)

Modern man started with the Upper Palaeolithic culture that far exceeded anything known
to their predecessors both in variety and perfection. Finer stone-tools and delicately worked
out bone implements were produced. There are abundance of one type of tools called bla4es
with long, smooth and parallel surface. Chisel-shaped blades were probably used for engraving
and working on woods. bones of antlers for using them as handles or shafts. Scrappers were
used for hollowing out wood or bone. Hollowed bones, organic shells were used as ornaments.
Upper Palaeolithic population, for the first time, left numerous traces of their artistic works
in several cave-walls showing many features of their daily life and ritual practices. Although,
paintings were confined mainly inside of caves, sculptured or engraved rock-blocks were found
from rock shelters or at the foot of the cliffs. Broadly, the Upper Palaeolithic arts are divided
into two types : mobile arts and cave arts. The former include arts on small pieces of stones
or bones while the latter are restricted to wall of caves and rock-shelters. So far more than
200 sites of prehistoric arts have been discovered, 90% of which are located in France and
Spain, greatest concentration being within 20 miles radius around the villago Les Eyzies,
southwest France. Some of these paintings were drawn in such an inaccessible portion of the
cave that might have been dangerous for both the artists and the viewers. Some researchers
have tried to explain such phenomena offering a relationship between the art and the magic.
Some arts are seen as being related to ritual before going out for hunting. Most of the arts are
zoomorphic, concerning sketches of different animals, wild or domestic. There are paintings
showing group-hunting of such animals like wooly Rhinoceros, mammoth, buffalo, bison,
reindeer etc. Herds of deers, horses, mammoths, bisons were also drawn. Some scientists believe
that in the absence of any scripts, these were the only ways for early man to store the knowledge
and memorize it. Sometimes it is indicated that cave arts might have served as an educational
means which is evident from the presence of lot of children's foot-prints near the sites of arts.
Probably, the children were trained for hunting and made familiar with the common animals
to~ hunted. Vari?u~ natural rocks and minerals were used as colours for these paintings. Long
continued Palaeohth1c culture came to an end around 8000-9000 B.C. when ice lost its grip in
Europe and America and the climate became warmer. Much of the ice melted and sea level
~"' .
.
' , .. L.:-. J
... 375
S AND ANCESTRY OF MAN
PRIMATE .
. d once more covering many land bridges and low lying areas. North Amenca and
w;; rais.:ecame separated and Britain was cut off from the rest of Europe. Th.e plant life. also
0
E ped 1·th the appearance of great forests spreading over Europe. Many Pleistocene ammals
change w · · 1 l' k
including mammoths, wooly rhinoceros, got extinct. In their place came smaller amma s ~ e
wild catties, smaller carnivors and othe~s. Man invented newer types of tools for huntmg
smaller-sized games. More sophisticated, precised and polished stone weapons are the
characteristic of the incoming Neolithic culture that exhibits a more versatile weapons. Along
with this, domestication of animals started. Staring with dogs, man began to domesticate other
animals like sheep, goat, pig and cow. This gave them constant supply of meat, milk, hides
and wool and this made the life less uncertain. But still people had to roam about to find water
and food at least for their pet animals. Cultivation soon followed the domestication and that
was the end of the nomad life of man. Cultivation started around some fertile river banks of
Near East, such as the river Tigris, Euphretis, Nile and the Indus. New range of tools was
prepared in connection with the cultivations. This was soon followed by potteries (7000 B.C.)
and preparation of utensils by burnt clay possibly to store the crops and also for cooking the
food-materials. This was followed by spinning and weaving and producing cloth and other
garments. Use of metal is another enormous advance in the way of civilization which started
around 4500 B.C. The first metal which was used was copper recovered by smelting of
chalcopyrite. Copper age continued for another 1000 years and about 3500 B.C. Bronze age
started when bronze was prepared by mixing copper and tin. This continued for another 1000
years. Sometime _lat~r perhaps by about 2000 B.C. man discovered the art of making ir~n 'and
that was the begmnmg of our modem civilization. · '
Chapter 27
RECORD OF VERTEBRATE FOSSILS OF INDIA
27.1 l\tAJOR OCCURRENCES
In India, there is no report of any significant vertebrate fossil before Permo-Carboniferous. One
possible reason behind this is that in India, true continental rock deposits are unknown prior to Upper
Palaeozoic. Bulk of the vertebrate fossils which include fishes, amphibias and reptiles are found
preserved within the freshwater/continental Gondwana rocks of India ranging in age from Permo-
Carboniferous to Lower Cretaceous. These rocks are exposed at different parts of Peninsula and
Extrapeninsula. What is more important that these fossil vertebrates are found mostly concentrated
within the Middle Gondwana rock formations rather than in its lower and upper part. The formations
belonging to Middle Gondwana are found well developed in some parts of Madhya Pradesh, Andhra
Pradesh and West Bengal. Denwa and Tiki Formation of Satpura basin, Yarrapalli, Maleri, Kota
and Dehannararri Formations of Godavari basin and Punchet Formation of Ranigunj basin are
especially enriched in the content of vertebrate fossils. Stratigraphically they belong to Upper
Gondwana group. However, a few fish and amphibia fossils are reported from Lower Gondwana
Vi.hi formation (Gangamopteris beds) of Kashmir. In general, fossils of birds and mammals are
unknown from Mesozoic rocks of India, although, there are reports of occurrence of teeth-fossils
of some micromammals from Jurassic rocks of Godavari basin. The major fossils of dinosaurs, the
most dominant reptiles of Mesozoic, are however found mainly within the post-Gondwana Upper
Cretaceous rocks of India, although, a few forms are known to occur within Gondwana rocks of
Godavari basin. Upper Cretaceous Lameta beds of Jabalpur, M.P. and its equivalents in Gujrat are
famous for its dinosaur fossils. The Ariyalur Formation (Upper Cretaceous) in Kaveri basin also
has yielded dinosaurian bones. Fossils of some fishes, amphibias and reptiles are also recorded within
the freshwater-estuarine Deccan intertrappean sediments from different parts of Peninsula which
are mostly of Upper Cretaceous-Lower Eocene age.
There are reports of a very few mammalian fossils from Lower Tertiary rocks of India
because of lack of suitable freshwater or continental deposits of that time. The bulk of the
mammalian fossils in India have come from the continental Siwalk Group of the Lesser
Himalaya of Neogene age (Miocene-Pleistocene). Although, the Siwalks contain all types of
vertebrate fossils but they are actually famous for their mammalian fauna. It is convenient to
list some of these vertebrate fossils of India in tabular forms dividing the whole faunas into
three groups : Gondwana, Post-Gondwana-Pre-Siwalik and Siwalik (Table- I 9, 20, 21 ).
27.l IMPORTANCE OF VERTEBRATE FOSSILS IN GONDWANA STATIGRAPHY
Gondwana rocks are well-known for their content of numerous plant fossils. Most of these
fossils are found from lower (Talchir-Ranigunj Formation) and upper stratigraphic horizons
(Rajmahal-Jabalpur Formation). The middle part of Gondwanu comprising the Triassic portion.
is generally poor in plant content, but contains abundant vertebrate fossils. So in an attempt to
subdivide biostratigraphically the middle part of Gondwana we have to depend upon the
venebrate fossils.
RECORD OF VERTEBRATE FOSSILS OF TNDTA 377
TABLE-19
LIST OF SOME GONDWANA VERTEBRATE FOSSILS OF INDIA
Animal groups Genera/Species Formation/Locality Age
Fish
Ganoid Amblypterus kashmirensis, Vihi (Gangamopteris) Lower Permian
(Palaeoniscoid) A. symmetricus, beds, Kashmir
Gardinerichthyes tewari (Mama) Formation)
Dipnoid Ceratodous (several species) Maleri and Dharmaram Upper Triassic
Formation, Andhra
Pradesh
Chondrichthyes Pleurocanthus pevidiens, Maleri and Upper Triassic
(Sharks) Xenacanthus indicus Dharmaram Formation
Osteichthyes Pholidophorus spp., Kota Formation, Lower Jurassic
(Bony fishes) Lepisosteus, Tetragonolepis, Andhra Pradesh
Depedium,
Indocoelacanthuus robustus
Clupavas neocomiensis, Raghavapuram, Lower
Ichthyopodector Sandstones, Andhra Cretaceous
Pradesh
Saurichthyes sp. Yerrapalli Formation, Middle Triassic
Andhra Pradesh
Amphibia
(Mostly Kashmirosaurus Gangamopteris beds, Late Permian
Labyrinthodonts (= Archaegosaurus) omatus, Kashmir
Actinodon risinensis,
Chelydosaurus
Rhineisuchus, Bijori beds, Late Permian
Gondwanosaurus bijoriensis Madhya Pradesh
Parotosuchus, Puchet Formation, Lower Triassic
Pseudosuchus West Bengal
Brachyops laticeps, MangJi beds, Late Permian
Rhynchosaurus Pranhita-Godavari basin
Parotosaurus rajareddyi, Denwa Formation, M.P., Middle Triassic
Prolacertiformis Yerrapalli Formation, A.P.
Metoposauru.\· maler1ens1s,
MaJeri Formation, Upper Triassic
Kuttycephalus,
Andhra Pradesh
Cosmocerops, Lydekkeria
Gonioglyptus, Glyptognatlms,
Pllchygonia
378 PALAEONTOLOGY
Animal groups Formation/Locality

-
Genera/Species Age
Reptile
(Synapsids Pristerodon mackayi, Kundaram Formation, Late Permian
Dicynodonts) Emydops platiceps, Pranh ita-Godavari
Cristecephalus microrhinus Valley, Andhra Pradesh
lystrosaurus ( several species), Pvnchet Formation, Lower Triassic
Dicynodon West Bengal;
Wadiasaurus, Rechnisaurus Yerrapalli Formation. Middle Triassic
Andhra Pradesh
Diapsids Mesodapedon kuttyi Yerrapalli Formation. A.P. Middle Triassic
(Rhynchosaurids)
Paradapedon hu.xleyi Tiki Formation, M.P. and Upper Triassic
Maleri Formation. A.P
.
(Protosaurs) Malerisaurus robinsonae Maleri Formation, A.P. Upper Triassic
(Thecodohts) Tikisuchus Tiki Formation, M.P. Upper Triassic
Proterosaurus Pu,khet Formation, Lower Triassic
West Beangal
(Phytosaurs) Parasuchus hislopi, Maleri Formation, A.P. Upper Triassic
Branchysuchus maleriensis
(Saurischians) Alwalkeria maleriensis, Maleri Forma.tion, A.P. Upper Triassic
Mesospondylus sp.
Barapasaurus tagorei, Kota Formation, A.P. Lower Jurassic
Kotasaurus vamanapalliensis •
(Pterosaurs) Campylognathus indicus Kota Formation, A.P. Lower Jurassic
Euryapsids Plesiosaurus indicus Umia Formation, Kutch Lower
Cretaceous
Mammal
(Symrnetrodonts) Indotherium pranhitai, Kota Formation, A.P. Lower Jurassic
Kotatherium haldanei,
Trishulotherium kotaensis,
Nakunodon paikasiensis
(fricocooodonts) Gondwanodon tapani Tiki Formation, ·M.P. Upper Triassic
Concentration of plant and vertebrate fossils, at different parts of Gondwana Group also indicate
the pattern of climatic change occurred during Gondwana periods. Lower and upper part of
Gondwana, rich in plant fossils, are generally considered as deposits under a humid climate with
profuse rainfall, while its middle parts with predominant vertebrate fossil may indicate a period
of considerable aridity. Broadly it may be inferred that initial glacial climate was followed first
by a humid climate and then by a periods aridity, after which humid condition reappeared.
RECORD OF VERTEBRATE FOSSILS OF IND/A
) 79
Som~ of t~~ vertebrate foss~ls arc used as age markers of Gondwnna horizons. flor example,
the r:eptile fossil ~ystrosaurus 1s taken as a marker of Lower Triassic horizon of the Gondwuna
~ontments: In Indian Gondw.ana~ this fossil is found to occur within Punchct Formation which
~s thus assi~ned to L~wer Tnass1c. The presence of stahlekriid dicynodont reptile Rech11i.va11rw:
m Yerr~pallta Formation of Godavari basin has confirmed its Middle Triassic age as this reptile-
group 1s not found before this period in any part of the world.
From the occurrence of many of these terrestrial vertebrate fossils in some continents like
lnd.ia, South America, South Africa, Australia and Antarctica, it is confirmed that during the
maJor part of. Upper Palaeozoic-Mesozoic times, these continents were close together forming
the supercontment Gondwanaland near the southern hemisphere.
27.3 IMPORTANCE OF DINOSAUR FOSSILS OF INDIA

Indian dinosaurs are mainly represented by theropods and sauropods belonging to saurischian
group, and also a few omithischians. Most of these are the fossils of armours, skulls, limb bones,
vertebral bones, coprolites, foot prints and a few eggs. Systematics of many fossils are known.
Investigations are going on for identification of many other specimens.
Some of these dinosaur fossils collected from India often show a wide geographic distribution.
Titanosaurus, Amarctosaurus, Laplatosaurus, Stegosaurus are forms which are known to occur
in many other countries outside India. Stegosaurus was a common form of Jurassic-Cretaceous
found almost throughout the world but their existence in Upper Cretaceous in known only from
'India and Madagaskar. Moreover, almost identical genus or even species of Turonian dinosaurs
are found in India, Madagaskar, Brazil, and Patagonia. From this, many workers have suggested
that southern part of South America, Africa and India were close together during Upper Cretaceous
times to form a landmass often designated.by the name "IAmuria" .
Dinosaur fossils are often used by stratigraphers to mark the lower age limit of Deccan Trap
lavas. Lameta beds, with associated Dinosaur fossils are found paraconformably overlain by
Deccan lavas. From this, it is suggested that Deccan lavas eruption did not commence before
the Turonian at least in the central part of India.
India is also famous for occurrence of several egg fossils of dinosaurs. So far two types of
egg fossils are known from India : completely spherical (Megaloofithus) and el?ngat~d
(Elongatoolithus); the former being larger than the other. Megalool,rhwr ( 16cm diam.) 1s
considered as egg of large sauropods, probabl~ of Tltanosau:us. They are known fro.m other
Gondwana countries and also from France, Spam and Romanta of Europe. Elongatoo/11hus (8-
1o cm diam.) eggs are so far known from Dinosaur Park, Kheda district, Gujrat. Possibly they
are eggs of carnivorous theropod dinosaurs.
27.4 SIWALIK MAMMALS . . . .

A list of some mammalian fossils obtained from Siwaltk Group 1s given 111 Thble-21.
A s r, t of evolution and migration of mammalian fauna

• Some. ea ur~s & t es· concerning the evolution and migration of Siwalik mammals may
ome mterestmg ,ea ur . · · I d' the prinl'1tes
be 'd d h s dd n bursting of diversified mammalian popu 1at1on me u mg .• ·
. c~ns1 ere_ e.re.. u ~ . O f a very rapid evolution of different groups of mammal.
e~~::1 c~~~are
w1thm the S1wahk times 1s suggestive f . . robably supported by the
This notable devel?pment and differentia~i~~ of~~~ and fo~ourable physical
abundance of ang10spermous food materrn s, a g
380 PALAEONTOLOGY
TABLE-20
LIST OF SOME POST-GONDWANA-PRE-SIWALIK VERTEBRATE FOSSILS OF INDIA
Animal groups Genera/Species Formation/Locality Age
Fish
Sharks Oxyrhina, Ptycodu.,· Ariyalur Formation, Upper
Tamilnadu Cretaceous
Lamna, Corax, Odontaspis Pondicherry Formation, Upper
Pondicherry Cretaceous
Pycnodus lametae, Pristolepis, Lameta bed, M.P., Upper
Nandus, Myliobatis curvipetalus Deccan Intertraps, M.P., Cretaceous,
Lakpat Formation, Kutch Lower Eocene
Ganoids Sphaerodus, Enchodus Pondicherry Formation, Upper
Pondicherry Cretaceous
Bony fish Lepidosteus indicus, Clupea, Lameta Bed, M.P., Upper
Musferia, Pycnodus lametae Deccan Intertraps, M.P. Cretaceous,
Plaeocene-
Eocene
Amphibia
Anurid Rana pussila Deccan Intertraps, Lower Eocene
Bombay
Reptiles
Dinosaurs
(Saurischian)
Sauropods Titanosaurus indicus, T. colberty Lameta Bed, M.P. Upper
Antarctosaurus septentrionalis Cretaceous
lndosaurus matleyi,
Laplatosaurus madagascarensis
Theropods Coeluroides largus, Lameta Bed, M.P. Upper
Camposuchus so/us, Cretaceous
laevisuchus indicus,
Ornithomimoides (2 sp.),
Dryptosauroides, Jabalpuria,
lndosuchus
(Ornithis- Lametosaurus indicus, Lameta bed, M.P. Upper
chians) Brachypodosaurus gravis, ~ retaceous
Dravidosaurus blanfordi, Trichinopoly and Ariyalur
Stegosaurus (unnamed) Formation, Tumilnadu
Chelonids Hydra.,·pis leithi Deccan Intertraps. Bombay Lower Eocene
- - ~- --
... . .., '. ' .
RECORD OF VERTEBRATE FOSSILS OF INDIA
381
environments. Th~ Si~alik assetnblage is found to be richer in many elements than the present
day fa.una .. The Siwahk ca~ivores are more numerous than the living forms in the same area.
The Siwahk ungulates (penssodactyls and artiodactyls) exceed their living forms in India with
more than 15 genera known from the Siwalik rock. Oxen and buffaloes have been reduced in
number of genera from eight to two at present.
All the Siwalik mammals are not truly of Indian origin. Migration from North America
prob~bly. by way o.f th~ land bridge across Alaska, Siberia and Mongolia made a large
contnbut1on. The S1wahk faunal-assemblage is very similar to the contemporary faunas of
s?u!he~n ~urope, northern Africa, central Asia, China, Burma and Malayasia. From this
s1m1lanty, it appears that at some periods of Siwalk times the land communication between
India and those countries was quite well-marked. The inter-communication between India and
Europe is said to have been restored in Lower Pliocene times . It is believed by some authorities
that on account of the restoration of free land-connection between Europe and Asia in Lower
Pliocene, various forms of mammals (such as the old horse Hipparion), giraffs, primitive
buffaloes, some antelopes, hyenas, and perhaps later true dogs and cats migrated from the one
to the other continent. Most of these migrants probably passed on to Africa. In fact, many of
them are now living in Africa while became exterminated from Europe at Pliocene-Pleistocene.
The primates probably were originated in North America and Africa and they reached India
via intermediate evolutionary centres. The large varieties and number of anthropoid apes in the
Siwalik fauna is a striking feature, which is perhaps not paralleled in other parts of the world.
It is likely that the original centre of radiation of this group was not far from India. They might
have come from the west by way of Egypt, Africa. Numerous specimens of the ape-like
dryopthecines, have been described from the Siwalik fauna. The starting point of some modem
apes like orang-utan, chimpanzee and gorilla probably lies within some of these apes.
The proboscidias originated in Africa, but India became ultimately the centre of adaptive
radiation for the Pliocene forms like Stegolophodon, Stegodon, and the Pleistocene form Elephas.
Elephas is believed to have migrated from India to Europe. Among artoidactyles Hippopotamus
and Sus were believed to have entered India from central Africa where they had their early origin.
Among the giraffids, the Chinji genus Girajfakeryx is considered as one of the oldest form which
was probably ancestral to the large branching-home? Dhok Pat~an forms ~uch .as Hyd~spi~he~ium
and Bramatherium. In India, the giraffids were relatively large m number m Middle S1wahk times
but became rare in Upper Siwalik and were subsequen~ly exte~mina~ed. Living members of this
family are restricted to Africa where it seems to have migrated m Pleistocene.
The Camelids were originated in North America from where t~ey migrated first t~ ce~tr~
As1a and then I nd'1a, p robably at the beginning
· . of Pleistocene.
. h' ..Bov1ds
· seem to have their ongm
in Central Asia from there they migrated mto India by C mJ1 times. . . .
Regard .mg th e horse, thei·r early evolution is traceable
.
in North Amenca where H1pparton
· d I d' · th h
· · · h )
(a pnm1t1ve orse , evo lved in Pliocene from Merych,ppus, soon migrate to n ta . roug
. . . N America and are believed to have migrated
Europe. Equus also evolved m Upper Pliocene m ·
into India by the end of Pliocene where it.shows further development.
s· l'k mammals
B. Causes of disappearance of •~a 1 . malian faunas are in many respects much richer
It has been pointed ouf that the Siwahk .mam bl of this region. The disappearance of
and more diverse than the living ~am~ahan asset~ itr:deed very striking and calls for some
the large bulk of the spectacular S1wa.tik ~amma . · 1 the causes of their disappearance the
explanations. While it is difficult to identify precise Y
following factors might be responsible.
382 PALAEONTOLOGY
The most commonly suggested reason for the disappearance of ~he Siw~lik vertebrates is
the intense cbld climate of the Pleistocene ice age. Lowering of temperature might be apparently
a very potent cause for extinction. The last glacial period witnessed almost a wholesale exti~ction
of the fauna, either as a direct or indirect result of the glaciation. The occurrence of the glaciation
has been well established by geological investigation in the Himalaya, and evidences are also
present to indicate the incidence of a very cold climate over the Indian Peninsula at the time.
The highly specialized nature and morphological complexity attained by the Siwalik mammals
in course of their evolution were also perhaps of some consequence in their decline. It is known
that a natural result of extreme biological specialization could be fatal, particularly in an area,
influenced by sharp changes in temperature or other ecological cum environmental conditions.
The simpler organisms could have survived such changes either by slow adaptation or migration.
The intermittent upheaval of the Himalayan mountain probably created a devastating effect
on the Siwalik mammals. The tectonic disturbance that swept the Siwalik country during the
Pleistocene phase of the Himalayan orogeny probably destroyed and drove away large hordes
of mammalian life.
C. Climatic conditions
A more or less warm and moist subtropical climate has been envisaged for the greater part
of the Siwalik history. This is borne out by the prevalence of a rich, largely forest-dwelling
mammalian fauna, fossil woods and tree trunks within Siwaliks sediments. Lithological attributes,
particularly those of the earlier Siwalik sediments also indicate increased rainfall and incidence
of seasonal changes. In the later Siwalik times the humidity was probably reduced and the
climate was less wet at certain stages (for instance in the Dhok Pathan time). A distinct cold
icy climate with a devastating effect on the fauna swept the area in Pleistocene. From some
levels of the Upper Siwa1ik have been described fine silty loess-like deposits which may
represent wind deposition in a relatively dry climate.
Surrounding habitats : Ecological consideration of the Siwalik mammals may throw light on
the geographic conditions existing in the Siwalik hill and valleys at that time. The Siwalik fauna
comprises forest-living, aquatic to semiaquatic, savannas or steppe-type of organisms. The
trilophodonts and dinotheres represent the primitive elephants whose teeth were adapted to eat
succulent trees and plants and harsher vegetation. The Siwalik Hipparion, from its relatively broader
hoofs and well developed lateral digits, appears to represent a savanna·or swamp type of organism.
Equus was adapted to harder ground and harsher vegetation. Among the bovids present, the antelopes
suggest prairies and steppes while the goats and oxen indicate a less arid habitats. The presence of
some rare organisms indicates a spell of scanty rainfall and desert conditions. The water-deer or
tragulids were river-bank dwellers and the Hippopotamus is suggestive of aquatic (or river-bottom)
habitat. The pigs and canids were mostly forest-dwellers but s_o me of the pigs (from the Chinji to
Dhok Pathan levels) were probably adapted to relatively more arid conditions, Gira.ffakeryx was
probably a forest-dweller, but the giraffas prefer open grassland with scattered trees. Broadly, it
has been envisaged that the Siwalik country was characterized during most of this period by the
belts of rich forests and open grassplains with streams flowing across. Considering the congenial
environmental and climatic conditions during the Siwalik sedimentation and t'ie remarkable find
of fossil apes from the Siwaliks of Haritalyangarh, it appears likely that remains of early man may
be present in the Siwalik deposits of India. No hominid bones have, however, been obtained so far
from them except some stone-implements and human artifacts which are also reported _from
Quantemary deposits of some parts of India. .
l TABLE-21 ~
g
~
a LIST OF SOME SIWALIK MAMMALS OF INDIA
I
8 Mammalian Groups Siwalik Formations and Kamlial Fm. Chinji Fm. Nagri Fm. Tatrot Fm. Pinjor Fm. Boulder i;:,
Dhok
i
"'C orders (Lower (Upper (Sarmatian) Pathan Fm. (Astian) (Villafranchian) conglome- C)
~:
"'?]
~
Tortonian) Tortonian) (Pontian) rate Fm.
'° ~
~
p
~
tt,
Primate Prosemii /,zdraloris ,/
s:
Cercopi- *Macacus ,/ ~
thecids
~
~
*Papio
*Senmopithecus
,/
,/
" ,/
C)
"'?]
Pongids Dryopithecus ,/ ,/
Gigantopithecus ,/ ,/ ~
:i:
Bramapithecus ,/ ,/
Sivapithecus ,/ ,/ ,/
(Ramapithecus)
*Simia ,/
*Pongo ,/
Camivora Creodortaa Dissopsalis ,/
Canids Amphicyon ,/ ,/ ,/
Vishnucyon ,/ ,/
Sivalictis ,/
/n.dractos ,/
*Canis ,/ ,/
*Mustella ,I ,/
~
*Ursus ,/ oc
~
'..>)
00
Mammalian Groups Siwalik Formations and Kamlial Fm. Chinji Fm. Nagri Fm. Dhok Tatrot Fm. Pinjor Fm. Boulder ~
orders corresponding age in (Lower (Upper (Sarmatian) Pathan Fm. (Astian) (Villafranchian) conglome-
~
Tortonian) Tortonian) (Pontian) rate Fm.
es
Camivora *Mellursus ./
Felids Vish11ufelis ./
Mel/ivorodo11 ./ ~
Sansa11osmilus
Smilodon
./
./ ./
i
*Felis ./ ./ •
*Panthera ./ ./
*Crocuta ./ ./
*Lutra ./ ./
Probo- Dino- Di11otherium ./ ./ ./

sddea therids ;
Trilopho- Trilophodon ./ ./ ./
.
1
dontids
Serridentinus ./
Tetralophodon ./ ./
Synchonolophus ./ ./ ./
Cherolophodon ./ 1
Elephantids Stegolophodon ./ ./ ./ ./
Stegodon
~
./ ./ ~
Archidiscodon ./ ~
H_ypselephas ./ ~
~
c--,
r
* Elephas ./ ./ 0
C')
~
'.
, Mammalian/ Groups Siwalik Formations and Kamlial Fm. Chinji Fm. Nagri Fm. Dhok Tatrot Fm. Pinjor Fm. Boulder
:::i::,
t'?)
:::s::· 8
orders corresponding age in (Lower (Upper (Sarmatian) Pathan Fm. (Astian) (Villafranchian) conglome-
Tortonian) Tortonian) :::i::,
(Pontian) rate Fm. t:::,
s 0
°'!"]
~
Proboscidea Elephantid *Loxodonta ./
~
"'-4
Perisso- Equids Hippario11 ./ ./ ./ ./ ./ rtj
~
dactyla
*Equus ./ ./ ~
Rinocera- Gaindatherium ./ ./ .,,~
tids 0
c-;
Aceratherium ./ ./ ./ ./ ~
Baluchitherium ./ ./ ./
0
*Coelodonta
*Rhinoceras ./ ./
./
./ ./
-
°'!"]
<'.
\)
i::
Artioda Anthraco- Heminuy x ./ ./
ctyla therids
Hyenaelursus ./
Telmatodon ./
Mericopotamus ./
Suids Listriodon ./ ./ ./
Conohyus ./ ./
Propotamochaerus ./ .;/
Dico1Jpbochaerus ./ ./
Hippolzyus ./ ./ ./ ./
*Sus ./ ./ ./
Camel ids *Came/us ./
Tragulids Dorcabw,e ./ ./ ./
w
Dorcatlterium ./ ./ ./ oc
VI
Boulder w
Siwalik Formations and Kamlial Fm. Chinji Fm. Nagri Fm. Dhok Tatrot Fm. Pinjor Fm. 00
.Mammalian Groups conglome- °'
corresponding age in (Lower (Upper (Sarmatian) Pathan Fm. (Astian) (Villafranchian)
r::s::
orders
Tortonian) Tortonlan) (Pontian) rate Fm.
es
Tragulid *Tragulus ,/ ,/ ,/
*Cervus ,/ ./
Hippo- *Hippopotamus ,/ ./
potamids
Giraffids Giraffakeryx ,/ ,/
Vislmutherium ,/
Hydaspitherium ,/
Bramatherium ,/
Sivatherium ./
*Gira.ffa ,/ ./
Bovids Perimia ,/
Tragoceras ./
Proleptobos ,/
Proamplzibos ,/
*leptobos ./ ,/
*Bos ,/ ./
*Bison ./ ./
*Capra ./
*Buba/us ./ i
*Surviving genera ~
t"I']
Besides the mammalian fossils mentioned in the table, Siwalik rocks also contain many other vertebrate fossils including fishes, amphibias,
and reptiles. Some imponant fish fossils are Rita, Macrona, Ophiocepha/us, Charcarias, etc. Mio-Pliocene subcrop rocks of Baripada, Orissa ~
have also yielded skeletons of some fish fossils like Scolicodo11 (modem shark), Priotlon. C/rarcariolamna. etc. From fluvio-glacial Karewa c3
beds of Kashmir, fossils of mammals such as Elep/ras, Bos namadicus, Equus 11a11wdicus, along with some s tone artifacts used by the early
man, have been collected. a
8
"'"<
PART - IV
PLANT FOSSILS
Chapter 28
INTRODUCTION TO PALAEOBOTANY
28.1 LIMITATIONS IN USING PLANT FOSSILS

Palaeobotany is the branch of palaeontology dealing with fossil plants. Study of the present
organic w~rld reveals that number of plants exceeds many times to that of animals. It is also
evident from the fossil record that plants appeared much before the animals. In spite of these
facts, fossil-plants are scarcer than fossils of animals. Not only so, our knowledge about the
morphology, systematics, evolution and phylogenetic trends of different ancient plant groups,
remains incomplete in many cases. The reasons behind this are as follows :
(i) Most of the plant fossils in our fos~il-record are terrestrial types and it is known that
preservation of fossils of terrestrial organisms is rather an uncommon event as many
such plants became decomposed and destroyed after their death for want of favourable
conditions.
(ii) Except stem, root and seed, other parts of plants (leaf, flower and fruit) are mostly soft
and have less chance of preservation. Fossils of algae, fungi, mosses are extremely rare
for their lack of suitable hard parts.
(iii) A large plant after its death generally becomes disarticulated; leaves, seeds and flowers
are detached from stem and the stem . itself broken into pieces. All tliese parts are
preserved separately and found as fossils of different form gl!nera and species. From
these fragmentary remains it becomes extremely difficult to reconstruct the structure of
a complete plant.
(iv) It is found that different parts of a plant have not evolved at the same rate and also
there are evidences of parallel evolutfon among the unrelated groups of plants. For
example, similarity between the fructification of a cycad-group to the flower of an
angiosperm does not indicate that they are genetically related. Again, there is a lot of
difference between Palaeozoic lepidodendrons and sigillarians with the modern small
lycopods, but they are all included within the same Lycopod group.
In general, the records of plant and animal fossils indicates one interesting point, that is
evolution of animals and plants are quite related with one another. But major changes of plant
kingdom preceded the major changes in animal world. For example, appearance of terrestrial
plants (vascular plants) took place in Silurian while land animals appeared in Silurian-Devonian
period. Again large gymnosperms appeared in Carboniferous-Permian which was fqllowed by
the appearance of large-sized reptiles in Mesozoic. The appearance of angiosperms in Cretaceous
was followed the abundance of birds and mammals Cenozoic. The phenomenon may be
explained by ecological cum environmental dependence among these two groups of organism.
Birds cannot live without fruit-bearing plants and existence of dinosaurs was impossible without
dense forest of gymnnosperms in Mesozoic times. ·
389
PA LAEONT<>LO<iY
:wo
----Flower
M--- Leaf Apex

11.r..--- Mid-Vein
,,.~~ Margin
Of Leaf
BranchStcm----a1111i._,
Fruit------.___............ PARTS OF A ·LEAF LAMINA
lnternode-------""""fl
Node----------l~Z
(a) A COMPLETE PLANT AND ITS DIFFERENT PARTS
Definite (Cymosc)
Indefinite {RacemoK) Dichocomous
La&cral
(b) BRANCHING
,. FIG. 28 • I : GENERAL MORPHOLOGY OF PLANTS
INTRODUCTION TO PALAEOBOTANY 391
In spite of lot of incoryipleteness of record of plant fossils, study of palaeobotany may be

useful in biostratigraphic subdivision of rocks. Plant fossils are also used in local correlation
of rock sequence but less commonly they can be used in intercontinental correlation due to
their restricted geographic occurrences, mostly controlled by local environments and climates.
These restrictions have made plant-types of one climatic zone quite different from that of other.
Usually a plant bears some morphological imprints indicating the type of environment in which
it is inhabiting. Comparing such features within fossils, plants are frequently used in the
interpretation of past environment, climate, and other geographic elements and in this respect
they are even more important than animal fossils. Again, a botanist can get a lot of information
about the phylogeny of different plant groups now found in the earth from the study of plant
fossils . For example, the gymnosperm group ginkgoales is at present, represented by a single
species Ginkgo biloba, but in Mesozoic there were at least IO to 12 genera belonging to this
group. Pteridophytes were much numerous than those found today. There are many groups of
pteridophytes and gymnosperms which were totally extinct today such as sphenophyllales,
cycadofilicales, cordaitales and so on.
The general evolutionary pattern of plants still remains unclear. It is difficult to say that
different group of plants appeared one after another like bryophytes were evolved from
thallophytes or gymnosperms from pteridophytes. It may be possible that some earliest and
generalized thallophytes might have given rise_to different groups of higher plants.
28.2 OUTLINE OF MORPHOLOGY OF PLANTS

Different parts of a plant are found as fossils : For ident~fication_and description of these
fossils, it is essential for a palaeobotanist to have a considerable knowledge about the
morphology of a plant. An angiosperms may be called as a complete plant as it has all the
different organs which a plant could have possessed. Our description has thus focussed on those
parts of an angiosperm commonly found as fossils.
A complete plant (Fig. 28-1 a) has the following morphological parts : roo.t, stem, leaves
and fructifications (fruit and seed). The root,. stem and leaf perform the different vegetative
functions of a plant such as attachment to soil (by root), conduction of water and food (by
root and stem) and preparation of food, respiration and transpiration (by leaves); while flower,
fruits and seeds are relateded with the reproductive function of a plant. A brief morphology of
each of these parts mentioned above is discussed below.
A. Root system
Root system normally lies below the soil consisting usually of a main root (prop root) and
some branches from it. The terminal end of the root is protected -by a cap-like structure called
· root cap and above the root cap ·o ccur a cluster of very fine delicate hairs called roqt hairs.
The former helps in penetration of soil and the latter in absorption of water from soil through
the process osmosis. Within the fossil world root fossils are comparatively rare.
B. Shoot system
Shoot system is normally aerial and consisting of the main stem, its branches and associated
leaves, flowers and fruits. Commonly, a leaf bearing branch. is called vegetative shoot and a
branch with flowers and fruits is called reproductive shQot. Tissues (xylem) within the stem
Palae(Gco)WP-50
-
392
• I . -. . . .. - . -~ ~ J - -
-
PALAEONTOLOGY
-.....
..... ..._
· h d' , lved minerals (absorbed by roots) to leaves where food
and its branches carry water wit 1sso d . . .
.1s prepared m
· the presence o f sun r1g.ht and chlorophyll. The prepared foo 1s agam earned by
stem through phloem tissue to the different parts of the plant.
Stem · The outer surface of stem may be smooth, striated, rugged.' hairy ~r spinose. _Often
a. dead outer
· .
tissues ,orm a hard woody layer, covering and
&
. protecting the mner soft tissues
called bark. Leaves/branches develop from some pomts of the stem called nodes. The
· be tween t wo success1·ve nodes is called intenwde. One or more leaves and branch.
portion . .
stems can develop from one node and the arrangement of leaves ~n. nodes a 1so mamtams
some distinct patterns called phyllotaxy. Woody plants often exh1b1t scar-mark left by _a
leaf-base after its fall or detachment from stem called leaf scar ( often found on a fossil
stem).
A stem may be soft, weak, slender or juicy or it may be stout, hard or ~o~y. Pl~nts with
mainly soft juicy stems are called herbs. They are usually s~all~r m d1mens1ons a~d
according to their total life time they may be annual (paddy), bienmal (beet) or pere~nial
(bananas). Shurbs are medium-sized plants, with numerous narrow woody stems but. w1th?ut
a main stem. Mostly they are perennial such as china rose, rose etc. Large plants with thick
woody stems are called trees which are mostly perennials e .g. mango.
Besides normal stems, plant may develop stems in abnormal position for performing
several other functions which ate called modified stems. Underground stem may develop
for storing food and also for reproduction such as rhyzome (e.g. ginger), tuber (potato)
etc. They often exhibit nodes and internodes like a stem, Aerial parts of stem may also
be modified variously to perform several other functions such as tendrils of climbers,
thorns of many other plants. ..
As regards the branching pattern (Fig. 28-1 b) of the stem there are two types : lateral
branching is a type where a branch stem emerges laterally from the side of the main
stem. This may be again of two types : racemose when main stem grows indefinitely
and cymose where a terminal bud stops the growth of main stem and a lateral branch
emerge below it from one side only. In dichotomous branching the terminal bud of the
main stem bifurcates into two lateral branches like a fork and this process . continues for
each branch repeatedly. ·
The inner tissue structure of the stem of a plant is of great importance as regards the
placement of a plant in the classification.
Tissue structure of vascular and non-vascular plant is quite different as the latter is devoid
of any vascular bundles. Monocot and dicot plants can also be separated by inner tissue
structures of a stem. Fortunately, a large number of stem and wood fossils are found in
pe.trified conditi_o~ (hard and ~ineralized), thin. sections of which may be studied under
microscope. So 1t 1s also essential to have a knowledge about the tissue pattern of common
s_tem. In general, ~II. vascular ~lants (pterosids) exhibit a common plan as regards their
tissue-structure w1thm stem (Fig. 28-2a). Most of them exhibit an outermost cell layer
called epidermis followed by cortex cells forming ground mass of the stem. Within cortex I
occur t~e vascular bundles or conducting tissues called steles and often a central piJh.
Cortex 1s ~eparated from the portion bearing vascular bundles by a layer of cells forming
endodermis. The pattern of steles in pteridophytes, as regards their position within stem
I
Epidermis - - - - - - . - - - - - - - - - - - - - - - ,
Corle>.---
• Xylc111
"Phloi:111
• Actinostelc Plcctostc:lc
Haplostclc
l'reridophytes (Protostelc)
~ - - - - - Epidcrmi~
Cortex--....
Afonocot
Dicot Gynmosperm
Angiosperm Siphnnostdc
(a) DIAGRAMATIC SECTIONS THROUGH STEMS OF VASCULAR PLANTS
Cortex
Endodermis ------1!:1":::n
Schlcrcnchyma, _ _ _ _~~~
Layers a11,-._ _ _ Schlcrcnchyma
Phloem Tissue
Cambium-----c:111z.
Xylem
Monocot Stem
Pith Ray
Dicot Stem
(b) ENLARGED VIEW OF T .S OF MONOCOT AND DICOT STEM
FIG. 28 - 2 : INTERNAL MORPHOLOGY OF STEM
l'ALAEONTOtOc;y
394
Orhicular
Sputulalc
· Elliptical
Subulatc Ovate
Aciculv Linear Lanccolatc Oblong
Lunate
Kidncy-Shapcd Falcatc Obovatc
Chordate Sagittatc
(a) SHAPE OF LEAF LAMINA
(j Acute Acuminatc
0Rounded
Cuspid ate
Emcrginatc Truncate
(b) LEAF APEX
Enlire Sinuaac Dcntatc Crcnatc
Lobed
(c) LEAF MARGIN
FIG. 28 - 3 : MORPHOLOGY OF LEI\F
!NTRODVCTION TO PALAEOBOTANY
395
and alignment of xylem and phloem tissues ma be .
siphonoste/e. Protoste/e is the mo t . Y of two types · protostele and
d h t' s simp 1e type where central pith is absent and xylem
OI
an P em issues occupy the central part. It is again of three types : in liaplostele xylem
oicur~ at centre andd pholem outside the xylem; in actinostele, the stele beco~es star
s ape at ce_ntre ma e up of xylem tissues whereas phloem occurs in between the arms
of the sta~; m plectostele, xylem and phloem become plate-like alternately arranged at
centre. (Fig. 28-2a)
Sip_honostele ~ound in gymnosperms and angiosperms shows a central pith surrounding

which occu~ rmgs of xylem and phloem tissues. In ectophloic, phloem occurs outside
xylem and m endophloic it is reverse.
In higher plants like gymnosperm and dicot angiosperms the vascular bundles are free
but collateral within cortex.
A cross section of a stem of a common dicot plant (Fig. 28-2b) shows the following part:
an outer epidermis (made up of thic~_sclerenc.hy.ma tissues), followed by cortex (made
up of thin parenchyma tissues) and an inner endodermis. Below this endodermal layer
occur vascular bundles in a row separated from one another by cellular medullary rays. At
central part of stem occurs the pith (with thin parenchyma ceJJs). A vascular bundle is
composed of outermost layers of large sclerenchymc!_tissues successively followed inward
by phloem layers, cambium cell-layers and xylem tissues (composed of tracheid, trachea,
wood fibre and wood parenchyma). However, in monocot stem (Fig. 28-2b) the vascular
bundles are found scatteredly distributed within the groundmass of parenchyma tissues,
each encircled by an ~idermal layer and a few layers of schl_erench ma tissues. The
tissue
pattern of a vascular bundle is also different which is maffify composed oNarge number of
schelerenchyma tissues surrounding an outer..r>hloem ..zone and the inner xylem system.
b. Leaves (Fig. 28-3) : Different forms of. leaves are found within plants such as foliage
leaf, scale leaf, brae leaf and floral leaf. Scale leaves are generally ~ssociated with some
modified stems, while bract leaves and floral leaves are parts of flower. Infact, the petals
of flowers are considered modified leaves (floral leaves) and sepals of flowers are one
kind of bract leaves. However, common leaf of a plant is the foliage leaf which has
normally the following parts : leaf base, petiole and leaf lamina (Fig. 28-la). Leaf base
is the point of attachment of a leaf with a node which may be a depressed (concave) or
ek•vated zone called leaf scar, position of which is often seen when a leaf is detached
from stem. In case of common gtass or sugarcane, a sheathing leaf base is found
encircling the entire stem. A leaf may be attached with stem by means of a stalk called
e t~/e when the leaf is called p_etiolate. There are many leaves which are without any
uc structure, called sessile, where leaf is attached to stem directly by its basal part.
Leaf lamina, the main part of the leaf, usually becomes green coloured due to presence
of £l)lorophyll pigment within the tissues. The shape of a leaf lamina (Fig. 28-3a) is quite
variable such as linear, lanceolate, elliptically oval, spatulate, sickle-shaped or falcate,
rectangular, cordate, orbicular, sagittate, reniform etc. Similarly the apex of a leaf lamina
(Fig. 28-3b) shows a wide variation such as acute, acutely rounded, acuminate, rounded,
truncated, emerginate, etc. There is also variation of the nature of the margin of leaf (Fig.
28-3c) such as entire, wavy, dentate or s. rreted, indentate etc. The most important part
of a leaf is the venation which is ma· :I , of t wo type:; (Fig. 28-4) : reticulate and parallel.
PALAEONTOLOGY
396
Multicostate-Divergenl
Multicos\atc Convergent
Unicostate
(a) REUCULATE
Multicostatc-Divergent
Unicostate Mu\ticostatc-Convergcnt
(b) PARALLEL
FIG. 28 - 4 : TYPES OF VENATION lN LEAF
_,-
••
397
Pinnate Palmate
Simple ·L eaf
Compound Leaf
(a) TYPES OF LEAF
Node----~
Stem---~~~..a
Primary Rachis
Uni pinnate Secondary Rachis
lmparpinnatc Paripinnatc
(b) TYPES OF COMPOUND LEAF
Ahernalc Opposite Whorl
(c) PHYLLOTAXY OF LEAF
FIG. 28- 5: TYPE OF LEAF AND PHYLLOTAXY
~
\C
Stigma} oc
Style
~=====~==:--==:--=::::__:::__:::__::::__:::__::::__:::--=::~ovary Gvnaccium
Anther } Sramcn(6J
A nd roc c 1u 111
Filament
~ ~ - - - - - - - - - - Petal (5'} Corolla
~~·-··, ~ Calyx (5)
' - - - - - - - - - - - - - - - - - - - - - Thal:111 11"

" - - - - - - - - - - - - - - - - - - - - - - - - - - l'..: tiole
(a) AN IDEAL COMPLETE FLOWER (In Longitudinal Section)
Petiole
xocarp ~ ~ Exoc""' } Pericarp
Endocarp
Dry Mesocarp
i:',f .~\ l ~1)
11'4{ }~\ \ \\'
p
1
, ~;.
n
M~oc..,,
.,, ·, :.
Endocarp
Seel.I
·-- '\ \\ \ /,/11 ~I'. ... .~.'- , ....... JJ Seed
Complete Fruit Fruit Cul Midway

~
Complete fruit Fruit Cut Midway ~
h:bhy Fruil ( Ma11 ~11, t'I')
0
~
l>ry Fruit (Nul)
(h) SIMPLE FRUIT ~
5
Cl
FIG. 28 - 6: FLOWER, FRUIT AND SEED "'<
'
399
In reticulate venation one or more veins bifurcate repeat di f · h
. . e y ormmg a mes structure.
Whe~ su~h a leaf po~sesses a d1stmct mid-vein (non-bifurcating) from apex to base, the
venation 1s called -re!JsJJlate..u.nicostate
. . type · In other cas·e , more th an one prmc1pa
· · l vems
·
m.ay be .present (reticulate ~ulb~o~_tate) which may be convergent or divergent types.
Like retic~late ty~e, par~llel venation may be parallel unicostate (a mid-vein and parallel
lateral vems on either s1d.e of it) and parallel multicostate, which may be again of two
types called f!arallel multicostate convergent (major subparallel veins emerging from base
and converging at the apex) and parallel multicostate divergent (major veins emerging
from base and diverging to the apex).
A leaf. is said simple when only one leaf l.amina. is present or it may be compound when
a leaf 1s composed of more than one lamina (Fig. 28-Sa, b). The individual of a compound
leaf is called leaflet. Leaflets are usually free from one another, may be petiolates or
attached by their bases to an axis calls rachis. A compound leaf is called.1wmate when
leaflets are arranged on either side of the r<!chi t_()ike a feather) oppositely or slightly
alternately. In this case, leaflets may be found in ev~n number or in pair when the leaf '
is called paripinnate or the rachis may be terminated by a single leaflet making the
number o leaflets o d when it is called im ari innate. A compound leaf where leaflets
are found at the terminal point of the rachis, like the fingers in the hand, is called palmate
compound leaf.
Arrangement of leaf (phyllotaxy) (Fig. 28-Sc) on the stem may be alternate on either
side of stem; opposite where two leaves emerge from one node in opposite direction and
whorled when more than two leaves emerge from one node.
c. Flower and fruit (Fig . .28-6a) : Different parts of a flower are as follows.
(i) Thalamus : Swollen part of flower base/flower axis.
(ii) Sepals : Several green coloured leaf-like bodies at the base of a flower, above the
petiole (when the flower is petiolate), the number of which may range from three to
numerous.
(iii) Petals : This is the second whorl of flower composed of several, usually bright
coloured laminae, which may be similar or dissimilar looking. Number, shape and
size of petals are quite variable.
(iv) Androecium : This is the third whorl of the flower containing several male
reproductive organs, each called stamen that yields numerous pollens.
(v) Gynaecium : This is the fourth whorl of flower bearing one or more female
reproductive organs, each usually shows three parts such as a basal ovary, a style at
middle and a stigma at top that receives the pollens.
A flower may contain all these five different parts when it is called a complete flower (such
as china rose). There are some flowers which bear either male or female reproductive organ.
Bracts are special leaves from the axis of which a solitary or a cluster of flowers arise. Bracts
may vary in number, size, shape, colour and mode of arrangement. Their primary function is
to protect a flower bud at its young stage.
Fruits (Fig. 28-6b) may be of two types : dry and fleshy. A fruit usually has a petiole or
stalk, and an outer exocarp outside the fruit and an inner endocarp outside the seed within the
fruit. In between this two, there is usually a thick/thin mesocarp which may be dry or fleshy.
Palae(Geo)WP-51
lwpter 29
MAJ R SUBDIVISIONS AND

GEOLOGI AL HISTOI{Y OF PLANTS
29.t MAJOR SUBDI\ ISIONS OF PLANTS
Plant grou may b~ com cni ·ntly subdivid ·d on th· basis of pr ·s --nee or absence of vascular
tissu ·s. Plants whi ·h hnve w"II d"v •lop ·d syst ·ms of cannls and lissu ·s for carrying water and
food are called ltasc11lar p/a11ts and thos" which luck them ure called mJ11-vasc11lar plants. The
form r group is often culled trach.eopliytes as th ·y b--ur food and water conducting canals,
tracl,aea. Obviously their origin is intin1ately associuted with the event of invention of land
by plants that probably took place in Ordo-Silurian times. Non-vascular plants includes such
Im er group as thallopliytes and bryop/iytes. The fornu:r includes various algae and fungi while
the latter are represented by hepaticeae, livewort and various mosses. As these plants are mostly
\\eak and soft-bodies and their fossils are extremely rare (except a few algae). The broad
subdivision of vascular plants are as follows-
A. Pteridophyta : spore-bear\ng plants.
a) Lycopodiales e.g. Lycopodium.
(b) Equisetales e.g. Equisetum.
(c) Filicales e.g. ferns.
B. Spermatophyta : seed bearing plants.
(a) Gymnospermae
naked-seeded plant without fruit. e.g. cycads. conifers.
(b) Angiospermae seed covered by fruit. e.g. pea, puddy. ·
(i) Monocotyledonae one seed-leaf. e.g. paddy.
(ii) Dicotyledonae two seed-leaf. e.g. pea .
. . Subdivis_ions of_n~n-v~s~ular pl~nts are mostly based on living forms. Vascular plants, in
spite of therr te~restnul habitats, left a large number of fossils within sediments ranging from
Upper Palaeozoi~ to Quate1:nary. Study of these fossils of vascular plants has revealed their
great morphologic complexity and diversity which are difficult to compare with their living
counterparts
. . at the present
. · . somc pa Iaeon to Iogrcn
earth · There ur·e ,·,lso · I 1·mutations
· · to put these
fossils into the classrcal botani ,, I d. · .· · t· · I ·
. ca rvrsrons o vascu ur plants. Fossils are mostly fragmented
and reprod~ctrve organs including flower, fruit or seed are mostly lost. There are muny fossil
grl~~-~fis whrch have no comparable parts in the recent. For this renson, pulaeobotunists huw
c ass, red vascular jJlant. . t b · t· . .
s on t 1e .1s1s o such teuturcs whrch ure usually visible in fossils.
The basis of subdivision of fossil vascular plants muy he selt.lctcd as follows :
(a) position of reproductive organs like sport!/tlower/seed.
(b) anatomy of vascular bundles.
(c) nature of leaf und stem und their relution.
",. 400
MAJOR SUBDIVISIONS AND GEOLOGICAL HISTORY OF PLANTS 401
Based on these, four groups of fossil vascular plants have been usually recognised viz.
Psilopsida, Lycopsida, Sphenopsida and Pteropsida.
Fossil plants are named according to the rules that govern the naming of living plants, but
with one difference. Plant fossils are mostly found in fragmented condition and it is impossible,
in most of the cases, to reconstruct a complete plant from their fragmented fossils. Thus, in
case of fossil plants usually segments of a plant like leaf, stem, seed, flower etc. are given
generic and specific status. They are called form genera or form species, which are obviously
artificial as contrasted with natural genera and species of botany. It is quite possible in case of
plant fossils that different generic names have been given for different parts of the same plant.
29.2 ALGAL STRUCTURES: STROMATOLITES

Stromatolites are stratiform, columnar or nodular structures found generally within carbonate
rocks (but may be present in other sedimentary rocks) resulting in from the combined effect of
life activity in lower group of plants and sedimentation. Sediments are bounded by the activities
of successive mats of blue-green algae (cyanophytes). Observation has shown that during the
periods of non-deposition, a thin algal mat is formed above the level of already deposited
sediments. Then, after deposition of a fresh carbonate sediments, algae permeate them and bind
them together and at the same time the algal structures are retained within the sediment.
Sediments thus exhibit variable types of forms and internal laminae depending upon the types
of algae and their activities. Stromatolites in true sense, cannot be regarded as true fossils. They
are activities of organisms preserved in rocks and can be treated as indirect fossils. Observing
the nature of .wide variations of these biogenic structures in rocks right from Precambrian (where
other organic fossils are rare), and the restriction of certain types of structures to some definite
geological age, palaeontologists are, at present, using stromatolites in the classification and
correlation of rocks especially of Precambrian age.
Stromatolites are classified morphologically into 4 distinct groups (Fig. 29-1) viz.
conophytonida (non-branching, discrete and cylindrical columns), kussiellida (passively
branching, cylindrical columns), tungussida (actively branching columns, divergent and widening
upward) and gymnosolenida (branching, pseudocylindrical columns, smooth-walled).
Stromatolites occur in rocks extending from Precambrian to Recent and are found growing
today as well. They are more predominant in rocks older than Ordovician and are most common
in Upper Precambrian (Proterozoic). Their oldest traces are found in Archaean rocks older than
3300 m.y. Use of stromatolites in stratigraphic correlation and zonation of ancient sediments
especially of unfossiliferous Late Precambrian sediments have been established through the
intensive research of some Russian scientists. They divided the Proterozoic rocks into Pre-
Riphean (2700-1650 m.y.). Riphean (Lower : 1650-1350 m.y., Middle : 1350-950 m.y.,
Upper : 950-670 m.y.) and Vendian (670-570 m.y.) on the basis of stromatolites present in
them supplemented by radiometric age data.
29.3 PSILOPSIDS
Psilopsids are probably the oldest and simplest forms of vascular plants. The li:ing forms
are called Psilotales and the extinct group Psilophytales. A few characteristics of psilophytales
are as follows :
) .i:.
0
N
~ - a .a. ,¢1_!.ic:;~ ~
~
~=~=--
....-::;...~~-
~~~
~~~:':~ --
'~::..~~~
~:-:..
~.,-~
.....
.
-::,Y
~
--
~ - : Cl",r•
-::-_.... '
~
Conoph)100
-:;::::
Kussii.:lla Tungussida
Gymnosolenida
(a) DIFFERENT FORMS
~..-~a ~,.~
~-~:::~·-
~-:~-~·
~;1~t~
- -
~I\ - ;;.-~}
'- · '\
~~I
~ :::!!! .::..
~~§
~:::::=
~~ $
~~~~~
~--~
~ ::.:_~
~)
~
~
/.:
{~
~
,-==
~
-"
~ ~ ~
'% -- ~ ~
~'
~ ~
~
-----
~'""
Discrete
Parallel Slightly Divergent
Strongly Divergent
(b) NATURE OF BRANCHING
Flat ~
~
Slightly Convex
~
Strongly Convex
ASConical Crenulated
(c) NATURE OF LAMELLAE ~

s:
~
21
~
FIG. 29 - l : STROMATOLITES 0
C')
"'<:
MAJVN SI///)/\ /,\' /()NS \Nil(,' /.'()/ ,{) ( ;/ (';\/ , IIIS'/'ONY OF /'LANrs 403
n) A sl n1i)h' b,,d w 111 low lh' µ;r•,• nf oq.\11n-diff ·re11tiation; stem simple and sparsely
brnn ·h\'d di ·h\1t111)111u ,' I ; som ·tin,,· s showing rhiwids but no root.
(h) St III u s 11 11II 111\.. •d ,vlth11111 h·uf nr with f ·w spines; stornatas on stern su~face; .vascular
systt•lll (wll\ 11 pre s ·111) r~ntrnl, shnwinp; 11 circular structure in cross section with outer
phloem ,•11d rl'li11p. h'm . .
le Sp\W\' tw11rl11µ orp,1111s 111· • t ·1·1ni1111I, •ith "r on main shoot .or on its branches representing
just 1111 •11l111'8t111r nt uf st ·rn-tip with a singl · spore cavity (homosphorous).
Th. first r~plH't of psiluphyt11I s is kn wn from Middle Silurian and they continued upto
Upp r D •nuvia11. Th •y 111· 1rmst common in Low ·r to Mkldlc Devonia~1. Among. many forms
J •s ·rib ·d frum lh, Midi.II \ l) ·voninn Rhyni · hert of Scotland, Rhynw and Pstlophyton are
the 11\QSI ~UI\IIIIUII.
Psilvpli 1w11 11 :i . 9. ~a(i)1 is n well known form nearly one metre high: T~ere are ~uba~rial
rhyzo111 ·s supporting th• nurrnw cylindri al aerial shoots. Rhyzornes bear ha1r-ltke rhyzmds atd~d
in absorptiun of wnt "r from soil. Dium ·tcr of rhizome or stem rarely exceeds I cm. T~e aer~al
shoot. is di ·hutmuuusly hrnnch·ll for more than one time. Branches are mostly endmg with
bifurcut -·d tips, som • of whi ·h ly•com~ swollcncd bearing sporangia. Bifurcated tips of ve~etat~ve
brnnd1 ·s ar • oft •n cir inut •\y coiled. Covering most of the aerial parts are short spme-ltke
struct111' •s oft ·n rcse1nbling ll~nr. The fossils of psilophytons are known to occur in Devonian-
arbonif ~rous rocks of many countries such as Canada, Belgium, Scotland, France, America,
Norwuy und also lndiu.
29.4 LY OPSIDS
This group is charnctcriscd by its simple foliage, small leaves spirally arranged, densely
covering the entire stem. Leaves are small, decreasing in size towards upper part. Stem bears
subaerial rhizomes and thick uerial shoots, dichotomously branching. There are cluster of fertile
leaves called sporophylls which on their undcrsurface bear sporangia (cones).
First report of lycopsid 8araRwmiatlri11 longijcJ/ia is known from Upper Silurian rocks of
Australia. They developed rapidly during Devonian and Early Carboniferous and reached their
climax. in Middle-Upper Carboniferous. They suffered a sharp decline during Permian and
appeared in u gr •atly di1ninished number in Mesozoic.
At present, there ure only four living genera of lycopsids which are Lycopodium. Selaginella.
Phyllvglossum and lsoetes. All of them except Phylloglossum are known from fossil record.
The two Upper Carboniferous-Permian forms of Europe and North Ameria, Lepidodendron
and Sigillaria are the most familiar. Lepitlodendron was one of the largest form of lycopsids
of Palueozoic (Fig. 29-2b). It grew over 35 metre with the main trunk of L meter across. The
t~ickcning wus due. to: secondary growth, producing woody materials composed of spongy
tissues. At the top of this tall plunt was n crnwn of branches mostly leafy, some of which were
fertile bearing cones. In fossil stem-surface spiral or rhombic leaf scars were present. At its
foot was an usual dichotomously branching root system with several roots each bearing slender,
spirally arranged rootlets. In fossil, they appear ns depressed pits on the surface. Lepidodendron
is a form which has been reconstructed from three fossils originally described as shoot
L.epiclopl,yl/um, root Stigmmiia und cone Lepidustrobus.
-· Scanned by CamScanner
PALAEONTOLOGY
.t()4
Sporangium
Sterile Stem ----,,1\
Fertile Stem ----~"lTL
RhizojJ - - - - (ii) PSILOTUM

(A Modem Psilopsid)
- - - - Slrobilus (wilh sporangiu)
A Probable Form of
Rhizoids A Carboniferous Lycops1·d Tree
(Height May Be 35 Feet)
Ll"COPODIUM
(A Modem Lycops1,I) lh) LYCOl'SIDS
Leaf~----::;:::~~.,,
Node----
~
r
&---w Strohilus
;1h Spo1w1g;,
.__ _ _ Stem (intcrnodc)

Fertile Branch ----Node
f:QIJIS£1'UM
(A Modem S1lCnop hi)
SPHENOPHrLLUM
hll Sl'U ENOl'Sll) S (A l'cr111i1111 Fossil Sphc1111J\S1d)
i-:Ki. 29-2 : SOM E FOSSI - l'LANTS I\Nl) TH IR LlYINU C<>UNTER-PART.S
.U.-\J R Sl 1BDI\ '/ I ,\JS A D EOLOGICAL HISTORY OF PLANTS 405
29.5 SPHENOPSIDS Fig. _9-_c)
A gene_ral "'h~m1cter of all members of sphenopsids is the presence of a longitudinally striated
stt'm "hich is ,tlso tnmsversdy articulated into nodes and internodes. The nodes bear whorls
of lea'.c;'.S. oupl d with these ch,1mcteristics, most genera possess modified fertile stalks of
spomngiophores ,, hich may be homologous with sporophylls of lycopsids. For the typical
arti ulated hara 't ·r of sh~ms, the forms are also named as Articulaticeae or Arthrophyta. The
onl} survi ing g nus of this group is horse-tail Equisetum, a plant which is few feet in height
with leaYes as , estigiul organs forming low toothed sheaths at each node of the stem.
Some of th important di, isions of sphenopsids are hyeniales, calamitales, sphenophyllales
and equisetales. The earliest sphenopsid is Protohyenia of Lower Devonian age had a rhizome
and u br~nched areal shoot. Pem10-Carboniferous Sphe11ophyl/11m is a shoot-form with fan-shaped
leaves. alymilt's is another common form of Upper Carboniferous of Europe. First appeared
in De onian, sphenosids became dominant in Carboniferous-Permian and were drastically
reduced in the Mesozoic and now surviving with only one genus. Some common fossil forms
are Sphe11opli) I/um , Schiw11e11ra, Phyllotheca, Calamites, Equisetites etc.
29.6 PTEROPSIDS (Fig. 29-3a-c)

It is the largest plant group that includes fems, seed fems, gymnosperms and angiosperms.
True fems are included in one group called Filicales and the other seed bearing forms (rest
three) are included within the group Spermal!)pliyta.
The fems and other advanced groups show the appearance of true leaves on stem. Probably
true leaves were gradual structural modification of stem off-shoots which became flattened and
•
subsequently webbed to form leaves.
A. Filicales (Fig. 29-3a) : Earliest fem Protopteridium has been recorded from Middle Devonian
rocks of Belgium, Scotland, Canada and China. Sometimes called pre-fems, probably, they
gave rise to Penno-Carboniferous seed-ferns and true ferns. These two forms are very much
abundant and similar in many respect and it is very difficult to assign a form genus (leaf
form) whether belonging to a fem or seed fem. Truly speaking, the only distinguishing feature
between them lies in their reproductive structure.
Fems have frond-like leaves growing fi:om a central crown close to surface or raised upon
a trunk of many meter height (tree fems). There are distinct but fibrous root systems. Leaves
are pinnately compound, variable shaped with/without a central midvein and lateral veins
mostly bifurcating. Ferns are mostly heterosporous with spores originating within small
sporangia on the under side of pinnae. Ferns are exceedingly diverse and complex. plants.
Appeared in Devonian they remained insignificant in Palaeozoic but attained climax. at
Mesozoic and are still continuing with reduction of their size and number compared to their
Mesozoic counterparts.
B. Seed-ferns/Pteridosperms/Cycudofilicules : Devonian may be said as the time of great
diversification in the history of plant life. At the beginning. neurly all vascular plants were
homosporous and by the close, heteruspory was common which possibly gave rise to earliest
seed plants, fossils of which are reported from Upper Devonian of North America. Existence
of such a group was first visualized 1900s when Lygi11opteris of filicopsid foliage was found l
~
-
PALAEONTOLOGY
406
Laf----"'T"'1
Stan----=~
Rachis Of 2nd Order
Probable Tree Form of

A Cretaceous Filicales Having
Leaf-Form Gleichenite.~
A Modern Filicalcs (fan)
(a) PTEROPSIDS (fems)
CYCAS (A Modem Gymnospenn)

WILUA.MSONIA SEWARD/ANA.
A Complete Cycad Fossil
(Ratontion After Sahni, 1932)
(b) PTEROPSIDS (Oymnospenns)
FIG. 29-3 : SOME FOSSIL-PLANTS AND THEIR LIVING COUNTER-PARTS
.,
MAJOR SUBDIVISIONS AND GEOLOGICAL HISTORY OF PLANTS 407
in association with seeds. From that time intensive study had proved that much of the fern-
like foliage-fossils of Permo-Carboniferous period are in fact, seed-ferms and now they are
grouped into a new fossil subdivision pteriodospermae or cycadofilicales (having leaf like
filicales and seed like cycads). At present, they are totally extinct. In them, seeds were borne
laterally or apically on leave but never on their underside like those of ferns nor the seeds
cluster into a cone like those of a true gymnosperm. The overall characteristics of a
pteridosperm are (i) relatively slender stem, often creeping, subaerial type, (ii) large froncJ.
like or simple leaf, (iii) seeds borne mostly single on apex or on lateral side of modified
leaves, (iv) occurrence of secondary wood and phloem, (v) pollen producing organs
exannulate and difficult to distinguish from fructification of ferns.
Several groups (nearly seven) of peridosperms have been so far distinguished of which three
were restricted in Palaeozoic; some restricted to Mesozoic and a few ranges from Palaeozoic
to Mesozoic. Taxonomically, they are classified along with other pterospsids within the
naked-seeded gymnospermae but in many respect they are different from other common
members of the group yet showing close affinities with cycads and ginkgos. The living genus
Cycas may be readily distinguished from a pteriodosperms by its distinctly organised
strobilus (cones of seeds) but its megasporophyll bears considerable resemblance to the
foliage leaves of seed-fem. Thus derivation of cycadaceous ovulate strobilus from seed-
bearing fronds of the pteridosperm may not be ruled out. On the other hand, a genetic
_connection between the pteridosperms and the psilophytales has been suggested by many
authors for the fact that the oldest known seeds of the pteridosperm were borne singly or
in pairs at tips of bifurcated stalks in a manner strongly reminiscent of the terminal
sporangium of the psilophytales. The main objection in accepting cryptogamic ancestor of
pteridosperm is the lack of any record of existence of heterosporous ferns which could have
given rise to seed-habit except the Palaeozoic fern Archaeopteris latifolia which could have
been genetically related to seed-ferns.
Some common Palaeozoic genera of pteridosperms are Lyginopteris, Lygenostoma (oldest
known Devonian form of North America), Calymmatotheca, Crossotheca, Neuropteris.
Glossopteris, Gangamopteris, Vertebraria, Thinnfeldia, Calypteris, Caytonia etc.
C. Gymnospermae (Fig. 29-3b) : True gymnosperms within pteropsids are grouped into
Cycadales, Bennettitales, Ginkgoales, Cordaitales and Coniferales.
(i) Cycadophytes : Members of cycadales and bennettitales are very similar in appearance
to one another and are sometimes grouped into one division 'Cycadophytes'. It has ·been
reported by some authors that they exhibit minute difference in their internal structures
which are not accepted by others who preferred to put them within one group.
Cycads appeared first in Triassic but attained their climax in Upper Jurassic and Lower
Cretaceous times. They showed abrupt reduction after Cretaceous and at present only nine
genera of this group are existing. Mesozoic is sometimes called 'age of cycads'. A I
generalized cycad has following morphologic characteristics. A trunk, may be branched, I

shows nearly 20m heights bearing rhombic, depressed spirally arranged leaf-bases at the ;
basal part and at the top a crown of frond-like leaves, each with two rows of pinnae ~
arranged on either side of a primary rachis. The frond, may be a meter long and being
Palae(Gco)WP-52
-
..:.-
c'it;
(a) A MODERN CONIFER (pine) (b) GINJEGO LJBOBA (a modem ginkgoalcs)

~
~
~
!'?']
FIG. 29 - 4 : LIVING CONIFER AND GINKGO ~

d
8
C)
~
MAJOR SUBDIVISIONS AND GEOLOGICAL HISTORY OF PLANT:~· 409
tough, is commonly. found a.s fossil. Cone of a cycad is borne singly at the centre of the
crown and the plant 1s exclusively unisexual. The cycad inflorescence hac; a striking flower-
like structure that is similar in several respect to the flower produced by modern
angiosperm Magnolia . The spirally arranged bracts beneath the stamen-ring of a cycad
cone may ~ compared to the calyx and corolla of Magnolia. The presence of large number
of stamens m the staminate ring finds a counterpart in the numerous stamens of Magnolia.
There is also similarity between ovaliferous receptacle of Magnolia bearing a cluster of
separate fruits to the cluster of seeds within cycad-cone. The floral similarities are also
accompanied by similarity in vegetative structure. Ve ssels and tracheids bearing
scalariform pits, a feature of Magnolia is present in all cycads. These resemblances may
be considered nothing more then analogies brought about by parallel evolution during a
long course of time. Some authors, however, intend to believe that this group (cycad) did
give rise to some of the modem flowering plants. If this assumption is true, it would partly
solve the old and perflexing problems of the origin of angiosperm.
Some common cycad fossils, are Ptilophyllum, Pterophyllum, Otozamites. Dictyozamites,
Taenoipteris, Williamsonia, Bucklandia, Cycadeoidea, Ctennis, Psuedoctenis etc. The most
completely known cycad fossil is Williamsonia sewardiana reported by Sahni ( 1932) from
Lower Cretaceous intertrappean sediments of Rajmahal Hills, India. The reconstruction
of the plant (Fig. 29-3b) shows a tree with a sturdy columnar trunk, not more than 2m
tall, of Bucklandia bearing at top a crown of Ptilophyllum leaves resembling the small
living Cycas. Stem exhibits rhombic leaf-scars spirally arranged forming a thick layer
around the outer surface of trunk and lateral branches are projecting through it, some at
places bearing flowers or fructifications (Williamsonia). The side shoots are constricted
at base suggesting that they broke away and served as propagative shoots. Major part of
these side-branches is like main stem but towards the tip they are inflated forming the
cone surrounded by bracts. The cone or flower is uniaxial consisting of a central receptable
covering long stalked ovules and club-shaped interseminal scales. Internally. in thin
section, the stem shows a pith surrounded by a thin ring of xylem tissues which are
compact with scalariform tracheids (suggesting it a cycad).
(ii) Cordaitales : They are a group of Palaeozoic arborescent gymosperms which together
with the seed-ferns constitute the bulk of seed plants of the Permo-Carboniferous coal
forest. Although, these two groups attained the climax nearly at the same time, they
differ markedly in their habit and external appearance. Cordaitales appeared first in
Devonian, attained their climax in Carboniferous-Permian and got extinct at the end of
Palaeozoic. Although, their origin is obscure, their oldest known members reveal such
an advanced state of development that it is certain that they were preceded by a long
evolutionary stages which is entirely unknown. Some characteristic features of this group
are as follows : A long tree with a trunk over 30m tall, showed a dense crown of
branches bearing broad simple leaves showing several parallel veins. Stem, often showing
annual rings, had very well developed secondary wood. vascular bundle and pith. Stems
are often found as large petrified wood-fossils. Inflorescences were borne on the stem
among the leaves whose main axes were slender, bearing stiff. taper-pointed bracts and
within the axis of each bract was a short bud-like strobilus.
410 PALAEONTOLOGY
Some common forms of Permo-Carboniferous cordaitales are Callixylon (stem),

Cordaites or Noeggerathiopsis (leat), Dadoxy/011 (stem), Cordaicarpus (seed), Artisia
(pith cast), MesoJ.y /011 (stem), Cordaianthus (seed) etc.
(iii) Coniferales : Conifers are woody, naked-seeded plants often growing to a large size (Fig.
29-4a). In possession of such vegetative features as the entire leaves and extensive
secondary wood, many of them show a decided resemblance to cordaitales although,
there are differences in their reproductive organs and primary wood structures.
Firs.t appearance of conifers is noted in Late Carboniferous but its course of evolution
in Palaeozoic is not well understood. After Palaeozoic, they rapidly evolved during
Triassic and Jurassic when they attained their climax along . with the cycads. Some of
the modem families probably became seggregated af that time but it .is ~ot possible to
trace them upto generic level which are identifiable only after Tertiary. Decline of
conifers started from Late Mesozoic as· they failed to compete with the incoming
angiosperms. At present, such forms like pines, larches, spruces, junipers and firs seem
well established only within a cold and high altitude forest.
Modern confiers are tall trees and each individual bears male and female reproductive
structures (bisexual). The pollens are often characteristically winged with air sacs suitable
for dispersal through air. The leaves are typically needle-like, spirally arranged on stem,
showing parallel veins. Longest (!f all trees (fossil or living) belonging to this group,
Sequoia, the giant redwood of California, sometimes becomes more than. l OOm tall.
Seeds of the conifer are woody, often winged, arranged on cone in a spirally manner.
Different part of conifer may be found as fossils which include stem and leaf, often
found together (shoot), cones! infloresence, seeds and pollen. Two important conifer
fossils of Palaeozoic are labachia and Bµ,riadia. Among the Mesozoic fossils,
Elatocladus. Araucarites, lndostrobus, Taklio~trobus, Brachyphyl/um. Taxas,
Araucarioxylon, CherioleP_sis, ~eplzalotaxas, etc. are worthmentioning. Some Tertiary
forms are Pynus, Keteleena, P1tyostrobus, Pityol~psis etc.
(iv) Ginkgoales : Gin_kgoales are an ancient group of plants which reached their climax of
dev~Jopment durmg Jurassic-~retaceous times. They probably developed from some
ptendo-spermous ancestry durmg later part of Palaeozoic. Points in common with the
latter are revealed by the structure of the seed, the swimming sperms and double forked
leaflets. However, at present the whole groups is represented I b · ·
c m· k~o b'o'l b (F" 2 ' on Y Y a smgle species
a 1g. 9-4b). The features, like the production of mobile . d h
tenmnally borne d . . .. sperms an t e
. see s, constitute a set of prumt1ve characters that have . d ti h' .
species the appellation of a '/i.,,ing fossil'. . earne or t is
?ur kn~wledge on extinct ginkgoales is founded almost exclu · .
impressions. Fructificution except for isolated . d s1ve 1y upon the,r leaf
include, a simple or compound type wi.th a Jo see s . ar,e rarely found. Leaf characters
· several spatulate foliage each of which is ng
up petio e often striate d ' bearmg
ti k d I '
· at t h e
divergent, each bifurcating towards apical pan ofr he o~Jy or deeply~ Veins are also
o t e Jeat.
Gi~k~oalc~ extends us far into the geologic pasr as L· .
Rh1p1dops,s are the forms of Early Permian .ar~ Pe~m1an. Psygmophyllum and
1
paced wnhm gmkgoales. Another form is
MAJOR SUBDIVISIONS AND GEOLOGICAL HISTORY OF PLANTS
411
also
f tentatively
d included within this group whi'ch ·ts a·tn kgop hy LL um of upper Devonian
·
1 1
o re an. · The name ~inkgoites is used by many authors to include all Ginkgo-like
leaf-fossils of Mesozoic, although some like to separate it from the genus Ginkgo. A
few other genera of Ginkgo-like leaves are Baiera (Jurassic), Phaenicopsis, Windwardia
(Lower Cretaceous), Hartzia, Sphenobaiera etc.
D. ~n~iosper.rnae : The time of first appearance of angiosperms is somewhat controversial.
Thetr fossils are recorded in Cretaceous or somewhat earlier but their abundance in Late
Creataceous is difficult to explain. Probably, they have had a longer and more extensive pre-
Cretaceous history than so far been revealed by their fossils. The scarcity of their fossils in
Early Mesozoic is possibly due to their predominantly upland habits where their remains
were not readily buried and preserved. It is also true for all plant groups that they are seldom
represented in fossil-record until they become well-established. Our paucity of knowledge
about pre-Cretaceous angiosperms indicates that they were not only subordinate to
gymnosperms as concern their numbers but there was also some ecological barriers for their
preservations.
Evolutionists are also at a loss to explain their origin. It is hypothetically assumed that the
angiosperm-line took shape at some unknown time during the Mesozoic Era. All the seed-
bearing plant groups, even the ferns have, at times, been proposed by various workers as
the possible ancestor of these flowering plants. Considerable attention has been centred on
the cycadoid fructification. and its resemblance to the flower of the angiosperm Magnolia
(mentioned earlier). If one accepts this relation, the question of recognizing the intermediate
stages would remain.
The oldest plant, known to literature, that bears any undisputable resemblance to an
angiosperm is Furcula granulifera discovered from Rhaetic (Late Triassic) rocks, eastern
Greenland. An example of possible Jurassic-Lower Cretaceous angiosperm is a piece of
secondary wood, Homoxylon rajmahalensis from Rajmahal hills of India.
Some of the oldest and undoubted angiosperm fossils have come from Lower Cretaceous
Kome beds, western Greenland. One form is Populus primaeva which is taken as the oldest
leaf to be referred to a modern ·angiosperm genus. Remains of monocots are always fewer
than those of the dicot plants in Cenozoic times. This is because many monocots inhabit a
drier environment where there is less chance of fossil-preservation and also because mostly
they are low-growing forms and their detached foliage is unlikely to be carried by wind to
the nearby preservational sites.
In general, monocot plants have linear or palmate leaves with parallel veins and their flowers
usually exhibit a triradial symmetry. Vascular bundles within their stem are found scattered
throughout cortex. Dicots on the other hand exhibit more varied forms with leaves more
commonly showing reticulate venation and flowers showing multiradial symmetry.
Most of the angiosperms are represented by large wood and leaf fossils but less fossils of
flowers, fruits or seeds. Petrified palm trunks are usually assigned to one stem genus
Pa/moxylon. Large leaves of sabal (Sabalites) are also found from Eocene rocks ?f New
Mexico and India. Nipa is a palm fruit of Eocene age found in London Clay Formation and
also in Deccan intertrappeans.
·412
PALAEONTOLOGY
l
Dicot plants, preserved in great number, include the arborescent forms such as oak .
. ' ' 'Wt 11 OS
poplars, sycamons, birches, allders, elms, hackberries, mapples, lindens, dogwoods etc. Man;
of the larger shrubs are also found as fossils, but low-growing herbs are poorly represented.
As a rule, however, the majority of the specimens found as fossils within different continental
deposits were derived from near-by source. Very often a single fossil flora of a locality
may represent the presence of several type of local habitats surrounding the site of deposition
at that time. A few familiar dicot fossils of Cenozoic are : Nymphaea, Rosa, Salix, Arbutus
Lithocarpus, Sorbus, Magnolia, Acer etc. '
As a whole, study of Cenozoic angiosperm fossils may give clue about climatic changes,
distribution of geographic elements, sign of orogenic movements, glacial advance and retreat
and ecology or habitats of the plants of that time.
29.7 GEOLOGICAL HISTORY OF LAND FLORAS

Record of land flora within Precambrian-Early Palaeozoic remains a matter of speculation.
However, evidence of presence of animal burrows within the Late Ordovician terrestrial
palaeosoil may indicate existence of a plant-cover sustaining the soil fauna. Probably, these
plants belonged to algae and lichens. The terms palaeophyte, mesophyte and cenophyte are
considered convenient application to the terrestrial flora of the world broadly corresponding
to the geological periods Palaeozoic, Mesozoic and Cenozoic respectively. However, appearance
of members of mesophyte and cenophyte took place ~t Late Permian and Late Cretaceous times
respectively. In general, palaeophytes exhibit dominance of pteriodophytes, mesophytes,
dominance of gymnosperms and cenophytes, dominance of angiosperms (Meyen, 1987).
A. Palaeozoic flora
The important floral groups within the palaeophytes so far recognised since Silurian may
be listed as follows :
Floral groups Important members Occurrences Age
Cooksonia flora Cooksonia, Parka, Czechoslovakia, Lower-Middle,

(pre-pteridophytes) Prototaxites England, Australia. Silurian.
UJSterophyllum flora UJsterophyllum, Same as above and Upper Silurian to

(pre-pteridophytes) Sporogonites Kazakhastan. Lower Devonian -
Psilophyton flora Psilophyton, Siberia, West Europe, Lower Devonian
(earliest pteridaphytes) Drepanophycus, North and South
Protolepidodendron America, China,
Australia. -
Hyenia flora Hyenia, Siberia, Europe, Middle Devonian
Protopteridium, N. America, N. Africa.
Ca/amoph yt011
Middle to Upper
-
Svalbardia flora Svalbardia, Siberia, Europe,
lycopodite.'i, N. America, N. Africa. Devonian
lepidodendrops-is --
4D
Floral groups Important members Occurrences Age

Archaeopteris flora Archaeopteris, Same as above Late Devonian
Callixylon,
Palaeoxylon
Rhacopteris flora Rhacopte ris, Western Europe, Late Devonian to
Sphenopteris, North America, SE Lower Carboniferous
Sphe11opteridium, Asia (India).
Cyclostigma
Eur-American flora Lepidodendro11, Europe, North Permo-Carboniferous
Callipteris, America, North
Lepidostrobus, Africa, Caucasus,
Pecopteris, Asia Minor.
Trigonocarpus,
Neuropteris
Cathysian flora Tringia, Northern China, Permo-Carbon iferous
Gigantopteris, North Korea, Indo-
Lobaton11ularia China, West-North
America.
Angara flora Callipteris, Asia, East of Ural Permo-Carbon iferous
Psygmophyllum, mountain, North
Rhipidopsis, Mongolia and Korea.
Angaroplois
Glossopteris flora Glossopteris, South America, South Penno-Carbon iferous
Gangamopteris, Africa, Australia,
Schizoneura, India, New Zeland,
Vertebraria. Antarctica. I
B. Mesozoic flora
Floras with Mesozoic affinity, however appeared in Upper Permian. In fact, the boundary
between palaeophyte and mesophyte is a transitional one involving the period Upper Permian-
Lower Triassic. Dicroidium flora of Euresia, Voltzia flora of North America and Europe and
Bunter flora of Europe are some floral groups of this transitional zone. In general, Triassic floras
are geographically differentiated into four provinces such as Siberian, European, Middle Asian
and East Asian. Siberian flora is dominated by equisetales, ferns and gymnosperms (peltasperms
and ginkgoales). The richest European flora are found in Middle-Upper Triassic of Western Europe
often designated as Keuper flora that contains major groups of gymnosperms such as :
bennettitales, caytoniales, ginkgoales, coniferales and cycadales along with several groups of forns
and equisetales.
Jurassic and Lower Cretaceous are usually referred to as the periods of gymnosperm
supremacy. The dominant plant groups represented are cycads, conifers, ginkgos but fr 1~ 1s are
not much inferior to them. Siberian Province became much wider covering northeast Europe
/
. I , ~ .
414
PALAEONTOLocy
where prevailed a warm and humid climate. Two diagnostic members are ferns (Cladophlebis
Coniopteris) and gymnosperms (Schizolepis, Pityophyllum and Podozamites). ·
In Jurassic, Equatorial Province covered western Europe, Caucasus, Middle Asia, China and
Southeast Asia. The dominant floral members are gymnosperms (Pterophyllum , Ptilophyllum.
Otozamites, Dictyozamites, Zamites, Williamsonia) and ferns (Phlebopteris Marattiopsis and
Gleichenites).
The Upper Jurassic-Lower Cretaceous floras show greater uniformity. These floras are of
particular interest for the gradual appearance of angiosperm elements in them. Four major
geographic provinces are recognised here based on plant fossils, viz. Canadian-Siberian,
European-Sinian, Equatorial (north Gondwanaland) and Notal (south Gondwanaland). Siberian
Province extended further towards north compared with its Triassic counterpart, where climate
was humid and favourable of coal accumulation . The chief floral members of this province are
ginkgoales like Baiera, Ginkgo; coinifers like Pityophyllum and cycads like Nilssonia. Ctennis
and Pterophyllun. European-Sinian Province included Europe, Middle Asia and East Asia. Here
prevailed an arid to semiarid condition indicated by a sharp decrease of ferns and equisetales.
Some important fossil members are : Pterophyllum, Ptilophyllum, Otozamites, Pachypteris and
Pagiophyllum. Equatorial Province included areas like nothern part of South America and North
Africa. Plants fossils are known only from a few localities. The Notal area included southern
part Gondwanaland including South America, India, Australia, Southern Africa and Antarctica
where developed typical members of Upper Gondwana Ptilophyllum flora.
C. Cenozoic flora
Change from mesophyte to cenophyte was also very gradual and transitional and the elements
of flora with true cenophytic affinity (angiosperms) appeared in Upper Cretaceous much below
the Mesozoic-Cenozoic boundary just as mesophytic elements appeared in Upper Permian . .From
this, it is often postulated that evolution of floral groups proceeded in advance of the faunal
evolution. The general habit of the flora gradually changed from the Upper Cretaceous owing
to gradual dominance of angiospermous elements with consequent of fall of gymnosperms and
ferns. Upper Cretaceous angiosperms differed markedly from those of Cenozoic. Change from
older to younger extant groups took place during Palaeogene. Thus transitional period from
mesophyte to cenophyte was considerably long.
The modem pattern of plant distribution Was caused by redistribution of continents and oceans
and cons~uent change of climatic belts of the world during the Cretaceous-Palaeogene periods.
Gradual d1sapP<:arance o~ the Tethys, birth of the Atlantic ocean, separation and drifting of
gondw~na continents with creation of seas between them, rise of the Alpine-Himalayan
mountains were some of the major events of this time.
In the cenophyte.
of the present day th d'f~ · · · ·
, e • 1erent1at1on of the floras are of maxmm express10n
that has. resulte~ m the prese.nt picture of the modem plant provinces of the world which include :
Holarcttc, Tropical, Australian and Antarctic zone Holarctic p · · • d Boreal
zone (arctic belt towards north) d T h · rovince ma111 1Y me 1u es
sout h) . an iet yan zone (temperate and subtropical zones towards
The most characteristic feature of B . .

of broad leaves that can b tt 'b d orea1 zo.ne at the beginning of Cenozoic was the prese~ce
e a n ute to paucity of sufficient light at high latitude. The maJor
" $ , a 15iis
, __;, __ :_ -- _ .... _.. ,. - -- --

415
forest elements were Ginkgo, Glyptostrobus, Magnolia, Osmunda . Thuja, Betula, Quercus.
P~pulus, Acer etc ..Towards the end of Miocene with further cooling off and differentiation of
chmate, many e~rhe.r Palaeogene members disappeared from Boreal zone giving way to a new
type of flora which included .cold-resistant conifers and small-leaved arborescent angiosperms.
The common members of this flora are Picea, Pinus, Abies, Larix, Salix, Tilia, Rosa etc.
. . Tethyan area supported at the beginning of Cenozoic a subtropical floral belt. Geographically,
~t mcl~ded US~, Europe, Middle Asia, China and Japan. Presence of evergreen trees and shrubs
mcludmg tree-hke fems, arborescent angiosperms like Parathesis, Porsea, Sapidus, Ficus, Sabal,
Clethra etc. were the characteristic of this zone.
Tropical Province during Cenozoic involved African, Indo-Malesan and Neotropic zone. At
the beginning of Palaeogene, most of Africa was covered by tropical rain forest, members of
which belonged to families like Arecaceae, Ebenaceae, Fabaceae, Rutaceae etc. With further
climatic differentiation during Oligocene-Miocene, this tropical forest was gradually replaced
by open woodlands of the savanna forest with such members like Fabaceae, Annonaceae,
Combretaceae, Ebenaceae, Euphorbiaceae etc. With further aridization at the end of Pliocene,
xerophytic and halophytic plants began to invade the area with such members like Chenopo-
diaceae and Tamariceae.
During the entire Cenozoic, Indian peninsula, Indochina and Malaysia constituted the Indo-
Malesian Provil)ce. Palaeogene flora of Deccan intertrappean beds in India are represented by
conifers (Auracariaceae and Podocarpaceae). Palms were remarkably diverse with more than
30 genera and some of the wood fossils were allied to Cocos, Nypa and Viracarpon. Oligocene
wood fossils, mostly belonged to tropical families of the Old World such as Tiliaceae, Guttiferae,
Sapindaceae, Anacardiaceae, Fabaceae, Combretaceae, Euphorbiaceae and the most important
family Dipterocarpaceae. The latter spread out from Malaysia to the East and West.
Dipterocarpus appeared in India at Miocene. Formation of temperate and subtropical forest in
the Himalayan belt began to large extent from Oligocene and mainly after the main Himalayan
uplift at Middle Miocene. Gradual fall of temperature resulted in the formation of a mountain
forest with dominant conifers. In the other parts, the flora remained mainly tropical type upto
Neogene, after which various degree of aridization occurred. In the Kutch valley, the Palaeogene
rain forest were gradually replaced by the Neogene deciduous forest.
The Neotropic Province included most of the So~th and Central America .. It was re~resented
by rain forest in the Palaeogene, dominated by ang1~sperms tha~ have c_ontinued to hve today
in the tropical f th New and Old World. The nch Neotroptc flora includes gymnosperms
zone o e f .. .k M . p
like Ginkgo, L"b d
L oce rus,
p odocedrus and angiosperm. amt 11es 1I e yncaceae, roteaceae,
T·1 ·
Polygonaceae, N ye tagmaceae,
· M'imosaceae, Caesalpinaceae,. Fabaceae, Rutaceae,
. 1 1aceae,
Cl· ·
Euphorb· M 1 · h' Meliaceae Myrtaceae, Ascleptadaceae, Rub1aceae etc. 1mat1c
1aceae, a pig 1aceae, fN · · u l'ft
changes during later Cenozoic were also reflected in many parts o eo~rop1c province. ~ 1
of the Andes results further differentiation of vegeta~ion of S0~1th America ~here the t~~:~cal
forest was partly replaced at first by savannas and finally by steppe and desert vegeta ·
The Notal Province included the entire Antarctica, Australia, New Zealand and southern ,~ost . .cf
h· vince was more homogeneous comp,ired ·
Part of South America. The Palaeogene flora of t ts pro . . . Podocaq,on ,~ •
to the present day flora. This included dominant conifers (Dacru/111111 , Phy lloclac.1uJ. :- ···
....---i·.; ~
Palae(Geo)WP-53
________..._.....- ...-~,r--~.?'!.
.. :~~- -~,1~'·"/ ,
416 PALAEONTOLOGY
and angiosperms (Myrtaceae, Araliaceae Winteraceae etc.) At the end of Palaeogene, Australia
became isolated from Antarctica and moved towards North causing transformation of floras into
several independent groups.
During the first half of Palaeogene, Antarctica was covered by conifers and small-leaved
forest with few angiosperm elements. By Middle Miocene when stable ice-sheets were formed
the vegetation become scanty, composed of few shrubs and herbaceous plants.
New Zealand is the only region of the world where ancient Palaeogene floral types are
preserved to date having been subjected to least alteration through time indicating less shifting
of this portion of Gondwanaland and prevalence ~f an unif(?rm climate since Palaeogene.
Palaeogene forest of New Zealand was like that of Antarctica composed of mixed components
(conifers and angiosperms) and such floras are still surviving there.
Chapter 30
RECORD OF PLANT FOSSILS OF INDIA
30.1 BROAD SUBDIVISIONS OF INDIAN FLORAS

Occurrences of fossil plants in India have been reported from very early days from rocks
ranging in age from Precambrian onwards. But many of these reports from ancient rocks
(Precambrian) lack details and in many cases, the identification remains doubtful either due to
their ill-preservation or due to lack of suitable techniques to study them. The first land plant
was reported from Upper Silurian-Devonian beds of the Himalayan basin. Floral materials within
continental rocks become abundant after the Devonian period and the Gondwana flora has
characterised the Indian Permo-Cretaceous rocks. Fossils of angiosperms are found from
Cretaceous onwards. It is convenient to subdivide all these fossil floras of India into four
categories : Precambrian flora, Post-Cambrian-Pre-Gondwana flora, Gondwana flora and Post-
Gondwana flora. However except the Precambrian flora, our discussions will be limited only
within megascopic floral elements.
30.2 PRECAMBRIAN FLORA

Report of fossils within Precambrian rocks of India was not so encouraging before the
successful use of transmission and scanning electron microscopes for detecting ultrastructural
features. At present, there are a lot of information of the occurrence of many such forms like
algae, stromatolites, acritarchs, and also doubtful occurrence of some filamentous calcareous
algae belonging to Dasycladaceae.
Report of the oldest plant fossils has come from Guddadrangayanhalli Formation of
Chitaldrug Schist belt, South India. The assemblage consists of sphaeromorphitae and
netromorphitae of the group acritarch (phytoplankton). The microbiotas which come from the
age group of 2.0-2.3 b.y. are in the form of circular, elliptical, triangular and oval shaped
microscopic bodies. This may be considered as the oldest fossil-record of India (Gouda and
Srinivasa, 1969). Stromatolites like Rugocystis and Navifusa are also known from some other
Dharwar group of rocks.
Evidences of organic life have been reported from Cuddapah Group of Andhra and from
the equivalent rocks of Karnataka (Kaladgi Formation) and Chattishgarh. Fossils occur in the
form of algal stromatolites belonging to such genera as Collenia, Archaeorestis, Cryptozoan,
Hydrophycus, /ndophyton, Gymnosolem, Tungussia etc. The age of this rock is dated as
1.6 b.y. Algal phosphorities are reported from some Precambrian rocks of Rajasthan belonging
to Aravalli Group along with stromatolites like Baicalia, Collenia and Minjaria.
Occurrence of fossil was unknown from Vindhyans for many days although there was a lot
of controversy about some lensoid carbonized structures which were discovered by Jones in
1909. These structures, originally described from Suket Shale of Kaimur Group, were named
as Fenmoria and assigned to primitive brachiopods. Later they were considered as some. non-
organic colloidal substances. Finally. these are assigned to the genus Tasmanite, an acntarch
417
418
PALAEONTOLOGY
(may be a spore of a non-calcareous algae). Other forms withi~ Vind~yans include some
primitive dasycladaceous alga which exhibit some disc-like semi-ovoid or kidney-shaped bodies.
But the most proliferous forms are stromatolites like Collenia, Kussiella, Conophyton, Baicalia.
Mammicus, Columnaris, Jurusania and others which are common in lower part of Vindhyan
Group.
A large number of fossils of acritarchs have been also reported from Lower Vindhyans of
the Son valley and its equivalent Bhima Formation in Karnataka. Upper Vindhyans of Rajasthan
also yield similar fossils. These forms mostly belong to sphaeromorphitae, pteromorphitae,
acanthomorphitae, netromorphitae, herkomorphitae, oomorphitae, and some others with uncertain
affinity.
Recently, algal stromatolites have also been reported from Shali Limestone, Lesser Himalayas
equivalent to Vindhyans (Valdiya, 1969) representing similar forms, often identical species.
30.3 POST-CAl\'IBRIAN-PRE-GONDWANA FLORA (Cambrian to Carboniferous)

In India, first report of fossils of vascular. land plants is known from the Silurian-Devonian
beds of the Spiti Himalaya (Sahni, 1953). Those remains are in the form of impressions and
are fragmentary in nature. The two forms are described, one of unforked axes and the other of
forked axes. They show either smooth or with minute spines on surface. No further details are
available. Sahni concluded that the unforked forms show Hostimella-like aspects while the forked
type has a Psilophyton-like aspects. He identified the latter as P. princeps. But as features. like
xylem, epidermis with cuticle and stomata or sponangia are lacking in these specimens, their
affinity still remains doubtful. The Devonian Aishmugam Formation of Kashmir basin also yields
Psilophyton along with Prolepidodendron.
Ho~ever, fossils of undoubted land. plants were reported as early as 1904 by Hayden.
Matenals were collected from Po Senes (Thabo Stage) of Spiti, associated with marine
formation'. the Fenestella Shales. Specimens were re-examined by Gothan and Sahni ( J937)
who con~idered the plant ~ed as Lower Carboniferous age from the associated marine fauna.
The fossil plants w~re assigned to Rlzacopferis flora reported from many other parts of the
world from. Carboniferous. rocks.
. The details of the flora reco gmse · , d so &-aar are as &-10II ows ·.
Rhacopter,s ovata,
. . R. mequtlatera, R. cf circularis
. · · Spheno Pen wm fiurc,·11arum. Sphenopter1s
t 'd' . ·
sp., Rhodea sp., and a few . doubtful forms
. identified as Asterop hy II anlI Ad' ·
. 1a11tttes. Equva Ient
Feneste II a Sh aIe Formatton of Kashmir also yields Rhac t · . ,
f orms l.k Rh d · . . op ens ovata a1ong with several other
I e o ea, Tnphylloptens, Lep1dosigillaria Ar ·I · ·11 · . .
. . . . · · • c 1aeos1g1 ar,a, Lep1dodendrops1s etc.
One mterestmg point is that most of the pre-Gondwana Carb · .
was broadly cosmopolitan in nature and even have the san . ~mfe1ous floras of the world
hemisphere. But it is difficult to imagine that thes d ~~ genera m both northern and southern
physical barriers and different climates in . . · e f e tcate plants were able to survive the
~ migratmg rom the north t th h I h be n
thus presumed that these plants at that time h d ~ h d o e sout . t as e
in different parts of the world under .· .a rel~c e. more or less the same stage of evolution
a s1m1 1ar c imat.ic zone. .
30.4 GONDWANA FLORA (Permo-Carbo 't
lndia was a member of the hyp th . m erous-Lower Cretaceous)
1
southern pole during Permo-Carb .~ ettca supercontinent Gondwnnaland existing toward the
om erous-Lower Cretaceous times. A typical flora often called
RECORD OF PLANT FOSSILS OF IND/A 419
Gondwana flora developed within the continents of Gondwanaland during this period. Gondwana
flora of India is conventionally divided into two parts viz. Lower Gondwana flora or
Glossopteris flora and Upper Gondwana flora or Ptilophyllum flora. Details of these two floras
have been summerised below.
A. Glossopteris flora/Lower Gondwana flora
Glossopteris flora is the purest of the contemporaneous Permian flora showing least
admixture. It probably originated first in Antarctica which apparently occupied the central part
of Gondwanaland. The earliest members of this flora are Gangamopteris, Glossopteris and
Noeggerathiopsis, which are found within ·the shales above the glaciogene Talcher tillites of
lowermost Permian age. Spores of Glossopteris and other genera are also found within these
shales. The flora attained its climax in Middle and Upper Permian during Barakar and Ranigunj
times after which it declined abruptly with a very few members persisting upto Mesozoic when
it was gradually replaced by a set of new floral group known as Ptilopliyllum flora.
(a) Origin and geographic distribution .
Evidently, Glossopteris flora grew as an indigenous product from the few plants that
were left over at the en.d of the Carboniferous glaciation that affected the entire
Gondwanaland. Glossopteris flora may have accompanied t~e ice-age or may .be slightly
younger than it. Occurrence of dissacate pollens and presence of poorly preserved stems
with growth rings fr~m some glacicll tillites of Talchir Formation may indicate that its
earlier members were co-existing with Late Carboniferous glaciation but Glossopteris
itself and most of the other members of this flora are believed to be post-glacial. The
flora soon reached its climax under a damp and wet temperate climate in marshy
environment. This is evident by its association with huge number of coal seams and
the general absence of growth rings in petrified stem fossils within Lower Gondwana
rocks.
In India, the major occurrences of Glossopteris flora are found localized within the
sedimentary rocks deposited along some major river basins of Peninsular India. These
include, the Son-Damodar basins (covering parts of Madhya Pradesh, Bihar and West
Bengal}, the Pranhita-Godavari Basins of Andhra Pradesh and Maharastra and the
Mahanadi basin of Orissa. Besides these, several isolated Lower Gondwana rocks
reported from some localities such as Sikkim, Kashmir, Rajasthan and East Coast also'
yield Glossopteris flora.
(b) Botanical affinity .
The floral elements mostly include peteridosperms. However, pteridophytes like
lycopsids, sphenopsids and filicales are also found but in less abundance. There are also
a few elements of true gymnosperm besides cordaitales group, which was not only
dominant but also restricted within this flora. Most of the fossils are impressions.
Carbonized and petrified fossils are also found. As usual, these fossil plants are mostly
represented by a large number of form genera and species and example of a complete
plant-fossil is virtually absent. But there is every possibility that. different parts of the
same plant have been recognised as different genera. However, close association of the
leaf fossil Glossopteris, stem-fossil Vertebraria, fructication Dictyopteridium and
dissacate pollen within rocks may indicate that those may be the parts of the same plant.
,
420 PALAEONTOLOGY
The diverse plant-parts preserved, include leaf (both simple and compound), stem,
vegetative shoot, fructification and spore and pollen. Although, ~ost of. the members
are indigenous, a very few forms like the genus Psygmophyllum 1s considered to be a
migrant from Angaraland as it was a chief member of the contemporaneous Angara
flora. As this fossil is found only in Kashmir basin within the Permian Gangamoptris
Bed, it may be concluded that Kashmir-Pamir area might have formed a land bridge
between the Gondwanaland and the Angaraland at that time.
(c) Fossil components, -stratigraphic distribution and age of the Dora
Glossopteis floral members are found in all the rock formations of Lower Gondwana
sequence of the type area (Damodar valley), such as Talchir, Karharbari, Barakar, Barren
Measure and Ranigunj Formation and their equivalent formations in other gondwana
basins. Some important form genera and their stratigraphic distributions in India are
shown in Table-22.
A long standing idea about the stratigraphic age of Lower Gondwana flora is that it is
ranging from Upper Carboniferous to Lower Triassic. In other words, it is beginning
with initiation of Talchir and ending with Lower Panchet. However, in most of the field
sections, the first appearance of Glossopteris flora is noted in rocks immediately
overlying the glaciogene tillites occurring at the base of the Talchir Formation. Again
the main bulk of the flora became extinct at the base of Panchet Formation. So it is
essential to demarcate the age of the tillites and the base of Panchet to find out the
total range of the Glossopteris flora. In this regard, the following points should be noted.
(i) The glaci~gene tillites. at basal Talchir are correlated lithologically with similar types
of format1on.s found m other p.arts of southern continents, such as, Dyka Tillites
of Sout~ Afnca. and Pagoda Tilhtes of Antarctica which are mostly Middle to Upper
Carbonaferous m age. ·
(ii) Fortu~ately, .in Indian peninsula, this glaciogene tillites are found sometimes
associated. with some patches of contemporeneous fossiliferous marine beds. In
DaltongunJ (Jharkhand), Manendragarh (M.P.), and at some places of Sikim and
Aru?achal ~radesh these marine beds contain Lower Sakmarian (basal Permian)
~arm~ f'.(oMssipls) suhch as ~ury~esma, Deltopecten, Conularia and Pleurotomaria. In
mana · · , t e fossils yielded by a r htl
Productus a d S . s ig Y younger marine band are Spirifer.
S It R ~ d treptoryn~hus which are similar to those of 'Productus fauna' of
a ange an are considered as Upper Sakm . (L . .
suggests that I r . aman ower Permian) age. This
.. . ower 1m1t of Glossopteris flora is basal Permian.
(111) Coming to the upper boundar i h be
locations contains Lower . y. t as en found that Panchet Formation at several
is in tum associated ·th T~aassac elements such as the plant fossil Dicroidium which
As most of the wb1 t e well known Lower Triassic reptile fossil lystrosaurus.
mem ers of Glo · 1 · fl f
Ranigunj Formation u d
as the top of Perm· n tr 1
. ssop er,s ora are found extinct at the top o
~mg the Panchet, the upper limit of the flora may taken
totally restricted w11.athn.' nht at ca~e, broadly the Glossopteris flora would become
m t e Permian.
::i.::,
TABLE-22 t'?']
(")
DISTRIBUTION OF LOWER GONDWANA FLORA IN INDIA 0
::i.::,
Younger~ t,
Plant Lower Gondwana Formations
Families 0
Genera Fossil form s "?']
groups Talchir Karharba ri Baraka r Barren Measu re Ranigu nj "'t,
./ ./ ~
Gymosperms Pteriodospermae Glossopteris Leaf ./ ./ ./ <'.
"'"j
./ ./ "?']
Gangamopteris Leaf ./ ./ ./
0
-
v.)
V)
Palaeovittaria Leaf ./
£;
Vertebraria Stem ./ ./ ./ ./ ./ 0
"'?']
Gondwanidium Stem ./ ~
Glossotheca Fructification ./ -
c:,
::i:,..
Dictyopteridium Fructification ./ ./
Cordaitales Noeggerathiopsis Leaf ./ ./ ./ ./ ./
Cordaicarpas Seed ./ ./ ./
Dadoxylon Stem ./ ./
Trigonomyelon Stem ./
Ginkgoales Psygmophyllum Leaf ./

Rhipidopsis Leaf ./ ./
Ginkgophyton Leaf ./
Coniferales Buriadia Shoot ./ ./ ./

Barakaria Stem ./ ./
Cycadales Taeniopteris Leaf ./ ./
Pseudoctenis Leaf ./ ./
.,&,.
-
Iv
\
~
N
(Continuation of TABLE-22) N
Plant Lower Gondwana Formations Younger~

Families Genera Fossil forms
groups
Talchir Karharbari Barakar Barren Measure Ranigunj
Gymospenns Incertae Ottokaria ./ ./
Sumaropsis ./ ./ ./
Seeds of
Callypteridium uncertain ./
affinity
Arberia ./
Cornucarpas ./ ./ ./
Pteriodophytes Lycopodiales Cyclodendron Shoot ./
EqisetaJes Schizoneura Shoot ./ ./ ./ ./
. Phyllotheca Shoot ./ ./
Sphenophyllales Sphenophyllum Shoot ./ ./

(Trizygia)
Raniganjia Leaf ./
Filicales
Leaf ./
Alethopteris
./
Pecopteris Leaf
./ ./ ./ ./
Sphenopteris Leaf
,/
Belemnopte ris Leaf ir-.
),,.
t'?']
0
<:
2J
s
C)
"<
_ ._. - ,..
RECORD OF PLANT FOSSILS OF INDIA 423
Members of Glossopteris flora are widely distributed and mostly reported from other
Gondwana continents of southern hemisphere. Table-23 shows distribution of some important
Indian species of the flora in other parts of the Gondwanaland.
TABLE-23
DISTRIBUTION OF SOME INDIAN PLANT FOSSILS IN GONDWANALAND
South South Folkland

Leading species of India Australia
Africa America Island
Gangamopteris cyclopteriodes ./ ./ ./ ./
G. kashmirensis ./
Glossopteris indica ./ ./ ./ ./
G. browniana ./ ./ ./ ./
G. retifera ./
G. damudica ./ ./
G. ampla ./ ./
G. angustifolia ./ ./
Gondwanidium validum ./ ./ ./ ./
Vertebraria indica ./ ./ ./
Sphenopteris polymorpha ./ ./ ./ ./
Schiwneura i11dica ./
Sphenophyllum speciosum ./
Noeggerathiopsis hislopi ./ ./ ./ ./
Dadoxylan indica ./ ./ ./ ./
B. Upper Gondwana flora

The Upper Gondwana flora of India is rich in cycad-conifer plant-assemblage occurring
within the Mesozoic Gondwana sediments. As the cycad genus Ptilophyllum is the most abundant
member of the flora it is also known as Ptilopliyllum flora. And, as Rajmahal Hills of Jharkhand
is the main site where a large number of floral members are preserved within the freshwater
intertrappean sediments (within Rajmahal lavas), in India, the flora is also sometimes called
Rajmahal flora.
(a) Geographic occurrence
Chief occurrences of Upper Gondwana flora are scattered all over the main Gondwana
province of Peninsular India which are as follows :
(i) Rajmahal hills of Jharkhand, where fossils are found within cherty shales,
intertrapped within the Rajmahal basalts.
Palae(Geo)WP-54 ~
- . - -- .
PALAEONTOLOGY
4 4
"
f. h Pranhita-Godavari Valley.
Kota- hikia\a- angapur assemblage o t e
"' .laba\pur ass •mhlage of the Satpura basin. M.P.
(i" Bansn and Persora assemblage of Rewa, M.P. .
·, . Andhra and Tamtlnadu.
(,) East ·oast oi..: ·urrenccs o f O n ssa,
\Ii) Jaisa\mir ass •mb\age of Rajasthan.
(,iii mia ass "' mb\age of Kutch.
lb) Botanical affinity

Like , er Gondwana flora, most of the members of Upper Gondwan~ are _form genera,
0 · 1 1·k l f t shoot flower fructtficat1on and seed.
represented by segments of p ants 1 e ea , s em, , ,
it is every possible that different parts of the same plant might have bee~ represented
by diffrrent generic names .. So far, an almost complete fossil has bee~ d1sc~vered by
Sahni from the Panchwara pass of Rajmahal hills, representing a plant with Pttlophyllum
\e:l\es, B11cklmulia stem and a flower like Williamsonia. This fossil plant is named as
\\illiamsonia sewardiana (described in the Chapter 29).
The bulk of the Rajmahal flora is represented by gymnosperms mainly cycads, conifers
and ginkgos. Within Pteriodophytes, filicales are the most dominant. The dominant
pteridosperms and sphenopsids of Lower Gondwana flora show an abrupt decline in
number, and the gymnosperm cordaitales is totally absent here. True angiospermous
elements are not so far reported but Homoxylon rajmahalensis, a pterified stem from
Rajmahal intertraps may represent an early member of angiosperm. Petrified patmw~s
are al so known to occur within the Umia Plant bed of Kutch, representing the topmost
G ndwana bed. All such petrified wood-fossils of palms of Cretaceous and Tertiary .rre
n w designated by the form genus Palmoxylon.
c) Constituents of flora and their stratigraphic distribution

The ch~cf con tituent floral members of Upper Gondwana Ptilophyllum flora and their
re ·pectwe occurrences in India are shown in Table-24.
(d) Age of Rajmahal flora
A ge limit of Upper Gondwana flora can be ascertained from the evidence of some marine
f · ·ils associated with them. Stratigraphically. Punchet Formation of Ranigunj Coalfield
ocrnr a t~e basal member of Upper Gondwana white Umia Plant Bed, Uttattur Plant
Bed and H1mmatnagar Sandstones are some of the top horizons of the Upper Gondwana.
Pur~che.t contains Dic:roidium as the only significant floral member of Upper Gondwana
which can. .be .a s.·s·ign~ d t~ L~wer T nass,c
· . · age from its close association with well known
Lower :nass,c r~~t,le toss,\ lystrosaurus (also found in Lower Triassic Lystms""'"s
w,1e
A. ot, ·dSouth
. . Afncu) · So. the . lower
. ·
age r1m1t· o f the flora. is definitely· Lower Triassic.
s r g.u s the upper age hmtl of the fl ora. . th e f o 11 owmg
. pomts
. .
may be worthmentioning.
(i) Umia Plant Bed with Ptil I1 II . ~ .
B •d .. . L up .Y um fiora is closely associated with marine Ukr.1
comaanmg ower C rctnceo , (A · ) .
Austrnlic ras UJld Cl . us ptam ammonate fossils such as ColombicerC1S
· 1 1omcera s.
RECORD OF PLANT FOSSILS OF IND/A 425
TABLE-24
DISTRIBUTION OF UPPER GONDWANA FLORA
~
Plant groups Forms Names of genera with stratigraphic

occurrences
A. Gymnosperms
Cycad ales Leaves Ptilophyllum, Pterophyllum, Otozomites,
Dictyozamites (R, J, K, U, Ut)
Taeniopteris, Nipaniophyllum (R, U, Ut, K)
Stem Bucklandia (R, J, U)
Pentaxylon (R, J, U) '
Flower Wifliamsonia (R, ·u)
Weltrichia (U, R)
Fructification Carnoconites (R, J)
Coniferales Shoot Elatocladus, Podozamites, Pagiophyllum,
Brachyphyllum (R, J, U)
Fructification Araucarites, lndostrobus, Podostrobus,
Takliostrobus (R, J)
Stem Podocarpoxylon (R)
Ginkgoales Leaf Ginkgoites (R, U)
Baiera (B) ..
Pteridosperm Leaf Dicroidium (P)

Thinnfeldia (K)
Pachypteris (R)
Cycadopteris (R, J)
B. Pteridophytes
Lycopodiales Shoot Lycopodites (P)
Equisetales Equisetites (R, U, 0)
Filicales Leaf Marattiopsis, Gleichenites, Cladophlebis,
Hausmannia, (R, J, U, K,), Protocyathea (R)
Matonidium, Weichselia (H)
C. Angiosperms Stem Homoxylon (R)
Palmoxylon (U)
R = Rajmahal Formation, U = Umia Plant Bed, J = Jabalpur Formation; K = Kota Formation, Ut = Uttattur
Plant Bed, B = Bansa Bed, P = Persora Bed, H = Himmatnagar Sandstone, o = Ongole Bed.
I
426 PALAEONTOLOGY
(ii) Rajmahal and Jabalpur flora are very much comparable with Lower Cretaceous flora
of Maryland and Varginia, U.S.A.
(iii) In Jabalpur, Ptilophyllum flora bearing beds are succeeded paraconformably by
Lameta Bed containing Middle to Upper Cretaceous dinosaur fossils like
Titanosaurus, laplatosaurus etc.
(iv) The two xerophytic plants Matonidium indicum and Weichselia reticulata found
in Himmatnagar Sandstone are characteristics of Wea/den flora which suggest a
Lower Cretaceous age of the flora.
Synthesizing all the data mentioned above it can be concluded that Upper Gondwana
flora is ranging between Lower Triassic to Lower Cretaceous.
(e) Distinction from Lower Gondwana Dora
Upper Gondwana flora stratigraphically succeeds the Lower Gondwana flora . Some
features of distinction between them are shown in Table-25.
TABLE-25
A COMPARISON BETWEEN LOWER AND UPPER GONDWANA FLORA
Features Lower Gondwana flora Upper Gondwana flora

1. Composition of flora
(a) Pteriodosperms Most dominant group of the Very few, nearly restricted to
flora Punchet Formation
(b)True Gymnosperms Less frequent; the only dominant Dominant group of the flora
group is cordaitales with abundant cycads, conifers
and ginkgoales
(c) Pteriodophytes Except equisetales, other forms Filicales dominant with also
are rare (especially, the filicales) other members
(d)Angiosperms Absent Possibly present
2. Initial climate Initiated in a cold icy climate; Initiated in a high temperature
attained climax in a wet, humid climate, attained climax in
climate humid condition
3. Evolution grade More primitive type of flora More advanced type of flora
-
4. Age limit Older flora, ranging within a Younger flora, ranging for a
narrower time limit longer time from Triassic to
(only Permian) Lower Cretaceous . ~
30.S POST-GONDWANA FLORA (Upper Cretaceous-Cenozoic)

In India, floral record after Gondwana period may be subdivided into four groups viz. :
((a)) UNpper Crefltaceous Deccan intertrappean flora, (b) Upper Cretaceous-Palaeocene algal flora, .-:J
c cogene ora, (d) Quaternary flora. /
RECORD OF Pl.ANT FOSSILS OF IND/A
427
(a) Upper Cretaceous-Deccan intertrappean flora
Cretaceous fossil floras are mainly located in Peninsular India (except report of fossil spore-
pollen from the Palaeocene-Oligocene rocks of Assam). Again, most of these fossils are known
from some continental sedimentary patches intertrapped within Deccan lavas which were
erupted during this time over a wide area of Peninsula. For this reason, broadly we can refer
these fossils as Deccan intertrappean flora, although, there are also records of few fossils
from Rajasthan and Gujrat lying outside the Deccan province.
Plant bearing intertrappean beds, however, are not uniformly distributed over the Deccan trap
province. Fossils reported from Kateru beds of East Godavari district, Andhra Pradesh, are
mainly of algae such as Holosporella, Neomeris, Acicularia and Terqueme/la.
Bulk of intertrappean fossils have come from different localities of Madhya Pradesh where
fossils are more varied types with remains of pteridophytes, gymnosperms and angiosperms.
Some of the important fossil localities are Mohagaonkalan, (Chhindwara; M.P.), Malurzari
(near Nagpur), Karia Mandie and Sagar (M.P.).
Some important fossil members are as follows :
Gymnospermous wood-fossils with affinity to conifers : such as, Mesoembryoxy/on,
Araucarioxylon : Conifer cones such as, Mohagaostrobus, Pityostrobus, Takliostrobus and
lndostrobus.
Angiosperms are mostly represented by several fossils of root, stem and seed. Some
important fossils are as folJows :
Angiosperm root
Monocot Rhizapalmoxy/011, Acrorhizas.
Dicot Dicotyrhizas., Somzeratiorhizas.

Angiosperm stem
Monocot Palmoxyon (about 22 species), Cyperaceoxylon, Protoxylon.
Dicot Ailanthoxylon, Simarouboxy/011, Sapindoxylon,

Bosellioxylon, Grewioxylon, Rhamnoxylon, Anacardioxylon,
Lecoxylon, Terminalioxylon, Euphorbioxylon, Ebenoxylon,
Vitexoxy/on and a few other forms of family Guttiferae,
Rutaceae, and Datiscaceae.
Fruits/seeds
Monocot Nypa, Palmocarpon, Tricoccites, Vivacarpo11, Graminocarpon.
Dicot Enigymocarpo11, lndocarpa, Harrisocarpon, Sahniocarpon.

Flower Sahniant/Jus, Deccant/Jtts.
Besides, some fossil leaves have been also reported with uncertain affinity such as Smilacites.
Flacourtiatites, Dicotytiphy/lum, Phyllites etc.
428 PALAEONTOLOGY
Majority of petrified stem fossils, found within intertrappeans belonging to palmae are
together assigned to a common artificial form genus Palmoxylon, which might comprise
many natural genera of palms. Many of these stems have a similarity with the recent form
Cocos which perfers a saline marshy land near the sea-shore. This might indicate the
presence of nearby sea-shore in this localities of Madhya Pradesh during the Deccan lava
eruption.
(b) Upper Cretaceous-Palaeocene algal flora

Marine algae fossils have been reported from Upper Cretaceous Bagh Beds of the Narmada
valley and Cretaceo-Palaeocene Niniyur Beds of cauvery basin. Both the beds contain rich
marine fauna which have confirmed their age suggesting Bagh Beds slightly older than Niniyur
Beds. A rich algae fossils have been reported from both Bagh Beds (Chiplonkar, 1944 and
Pal, 1971) and Niniyur Beds (Rao and Pia, 1936). In spite of slight difference in age, there is
remarkable similarity in type of fossil algae of these two localities and the bulk of the materials
represents Dasycladaceae algae. Some important- forms are Dissocladella, lndopolia,
Holosporella, Orioporella, Neomeris, Acicularia, Clypeina. A few codiacean algae of Bagh
Beds are Arabicodium, Boniena and Halimeda. ·
(c) Neogene flora (Mio-Pliocene)

A rich floral fossils of angiosperms has been found within some continental sediments
deposited at different parts of India, mainly from Tipam and Dupitila Formation of Assam
and their equivalents in Tripura, NEFA, Bengal and -Bihar, Middle and Upper Siwaliks of
N.W. India and also from Cuddalore Sandstones (Mio-Pliocene) of Corromondel Coast,
South India. Most of these are represented by petrified fossil woods of angiosperm belonging
to monocots and dicots which have close affinity with the recent forms of that locality.
Table-26 shows a list of some of the important fossil-forms reported from these three
localities (north-east, north-west and south-east India).
From the study of the Table-26 it is clear that many of these genera are common to all
these Neogene deposits of India, especially there is an overall resemblance of flora between
Assam and Cuddalore. From the present distribution of the modem equivalents of these flora,
it is envisaged that India had a broad uniformity in its tropical vegetation during Neogene
times. Dipterocarpaccae, Guttiferae, Anacardiaceae and Combretaceae were dominant
vegetation at that time and some of them had wider distribution in Neogene than that found
in the present day. For example, Dipterocarpus is found at present in evergreen forest of
Kerala, Mysore and Assam but their fossils are found in almost all Neogene deposits of
India indicating a probable existence of luxuriant evergreen rain-forests to deciduous forest
over a large part of India during Neogene.
A few more points whkh may be revealed from the study of overall Tertiary flora of India
are:
(i) Palaeogene flora are mainly found in Peninsula largely of Eocene age while Neogene
flora are dominating in Extrapeninsula mostly of Mio-Pliocene age.
(ii) Although, a large number of families are common to both the groups, there are some
Palaeogene forms which seem to have disappeared by the Neogene times such as
RECORD OF ('LANT FOSSILS OF INDIA 429
TABLE-26
NEOGENE FLORA OF INDIA
Assam Siwalik Cuddalore
Genus Family
(N E India) (NW India) (SE India)
M onocot Palmae ./ ./
Palmoxylon
Dicot
Kayeoxylon Guttiferae ./ ./
Calophylloxylon
Dipterocarpoxylon Di pterocarpaceae ./ ./ ./
Anisopteroxylon Dipterocarpaceae ./ ./ ./
Shoreoxylon Di pterocarpaceae ./ ./
Peltophoroxylon Leguminosae ./ ./
Cassioxylon Leguminosae ./
Cynometroxylon Leguminosae ./
Caesalpinioxylon Leguminosae ./
Accacioxylo11 Leguminosae ./
Prohudioxylo11 Leguminosae ./ ./
Bauhinioxylon Leguminosae ./
Terminalia Combretaceae ./ ./ ./
Mangiferoxylon Anacardiaceae ./ ./
Glutoxylon Anacardiaceae ./ ./
Barringtonioxylon Lecy th idaceae ./ ./
Careyoxylon Lecythidaceae ./ ./
Ebenoxylon Ebenaceae ./
./
Pometioxylon Sapindaceae ./
Sapindoxylon Sapindaceae ./
Alangioxylon Alangiaceae
./
Brindelioxylon Euphorbiaceae
./
Glochidioxylon Eu phorbiaceae
./
Phyllanthinium Euphorbiaceae
./
Siderinium Sapotaceae ./ ./
I
L Parinarioxy/011 Rosaceae ./
PALAEONTOLOGY
430
Araucariaceae, Cannaceae, Cyclanthaceae and Proteaceae. The families which may be
considered typical of Neogene are Dipterocarpaceae. Ebenaceae, Sapota~eae, ~osace?e,
·
Al angiaceae, an d a few o th ers. May
· be .s· ome of them might have existed m earlier
times but at least they are not recorded as fossi Is.
(iii) Leguminosae which was meagerly represented in Palaeogene, becomes abundant since
Neogene times.
(iv) From the .occurrence of Palaeo-Neogene fossils of Kutch and adjoin.ing pa~t~ of
Rajasthan it may be concluded that in contrast to the prevailing xer~phy~1c c~nd1t1~ns
of today, there was much higher rainfall and a moist climate prevailed m this region
at that time. The present aridity of this region is possibly a post-Pliocene development.
(d) Quaternary flora of Kashmir

Quaternary deposits, with reports of large quantity of macro and micro plant-fossils, are
mainly found in Kashmir and adjoining areas of North India. These mega fossils mostly
have their living counterparts in the present vegetation of the area. Fortunately, the
fossiliferous Karewa Beds are more or less contemporeneous with sequences of Pleistocene
glacial and interglacial phases that provides a time scale against which a particular flora
may be dated. Fossils have been collected mainly from three localities of Kashmir such as
Liddermarg, Laredug and Ningal Nata (Puri, 1948). About 70 genera have been identified,
mostly of monocot and dicot angiosperms of aquatic, subaquatic and terrestrial habitats.
Some common forms are Acer, Berbies, lndigofera, Populus, Pi,ius, Picea, Juniperus,
N~lumbium etc. ~he associated lignite beds show several genera of diatoms and pollen.
L1ddermarg flora 1s mostly tropical while Ningal Nata flora is exclusively of temperate nature
containing _willow, p~pler, cherry, walnut, maple, alder, spruce, fir and pine. Their modern
representatives are still found_in the Kashmir valley at high altitude.
In some _respe_ct, the present day vegetation differs from Pleistocene flora of this area The
outstanding differences arise from the absence of oaks and alders a d · f · ·
P resent da fl Th C d n ranty o spruce m
. - Y or~. e e rus-oak-wood and Spruce-deodar-oak wood which flourished here
~ the !ar~y KPle1hsto~ene are no longer present today except on the south of the Pir Panjal.
e oa s in as m1r valley have been replaced by Pi · ·
community today The modern pine fi . arotw, so that we have cedrus-parotia
. - ir community was non exist t d . Pl . Th'
is possibly due to great climatic chan . d - · e~ unng e1stocene. 1s
in Late Pleistocene which at present :es cau~e by the final uplift of the Pir Panjal peak
' revents monsoon to enter the valley.
-· ,.
- - - . .... _ , "i. ...
PART - V
MICROFOSSILS
' ~ ·l
Chapter 31
INTRODUCTION TO MICROPALAEONTOLOGY
31.1 HISTORICAL DEVELOPMENT OF MICROPALAEONTOLOGY

Micropalaeontology is a branch of palaeontology t.hat concerns with microfossils. It is self-
evident that the difference between micropalaeontology and macropalaeontology concerns
primarily objects and methods while both branches contribute towards the solution of same
general scientific problems. However, from historical point of view micropalaeontology has taken
its own course of independent development and in recent times it has been proved that
microfossils can be used more successfully in stratigraphy at least, in some cases, t.han
megafossils. A rock apparently appearing unfossiliferous may contain a large number of
microfossils that can be used in finding out the age of the rock concern.
The foundation of micropalaeontology was laid down by Alcid d'Orbigny ( 1802-1857) whose
work in other branches of palaeontology and stratigraphy was equally outstanding. His work
was centered on description, classification and distribution of foraminifera. It was Ehrenberg
( 1854) who studied on the influence of microscopic organisms in nature and tried to raise the
study of rock forming microfossils to the status of a branch of geology for which he proposed
the name 'microgeology'. He described many groups of microfossils in addition to foraminifera.
Cushman ( 1948), a leading authority on foraminifera became one of the first to establish the
value of foraminiferal studies in the field of Economic Geology. Practical value of
micropalaeontology has led to its application in all countries for the exploration of oil. Study
of microfossils took a new turn after Glaessner. His book, 'Principle of Micropalaeontology'
published in 1944, has been taken, even now, as the basis of all types micropalaeontological
works, especially on foraminifera. In his book he gave a comprehensive morphological
description of different groups of microfossils and discussed on their taxonomy, classification,
stratigraphic distribution, ecology · and their application in petroleum exploration. Recent work
on micropalaeontology includes statistical treatment of morphological features found in
successive populations. This helps to understand the palaeoecology of the fossil assemblage,
variation of its spatial distribution caused by change of temperature and depth of basin, phases
of rapid and fundamental changes in its course of evolution and also the succession of
microfaunal zones of worldwide or intercontinental geographic range. Micropalaeontology,
'academ.ic', as well as 'applied', is now turning towards more analytical methods. A detailed
morphological analysis now becomes the basis of taxonomy; ecologic analysis of population
reveals facies relations and a stratigraphical analysis provides the clues for necessary evaluation
of a species or an assemblage of fossils in its application in stratigraphic use.
In the present chapter, the author intends to give u details of morphology of foraminifera
and an outline of morphology of some other important groups of microfossils, their ecology
and broad subdivisions with special emphasis on the foruminifera.
433
434 PALAEONTOLOGY
31.2 MICROFOSSILS : A GENERAL ACCOUNT

Fossil world is commonly divided into larger macro or megafossils and smaller microfossils,
each kind with its own methods of collection, preparation and study. By definition 'microfossils'
are those· which can ·be studied and identified only by their study under microscope. Many
microfossils (such as larger foraminifera) are visible in naked eye but their proper generic and
specific identification requires their microscopic investigation. However, parts of shells, bone
of larger animal-fossils or wood-fossils of larger plants are often studied under microscope to
get their detailed structures and compositions. They however do not constitut_e microfossils.
Microfossils include microscopic skeletons of the individual microorganisms like foraminifera,
radiolaria ostracoda, diatom or they may represent disarticulated/discrete microscopic elements
of some larger or macroorganisms such as spicules of sponge, scolecodonts, echinoid ossicles
(spines) etc. Fossils of spores and pollen, representing reproductive elements of nonflowering
and flowering plants respectively are in this sense also microfossils. There are another group
of microfossils which are minute organic elements whose actual affinities are still uncertain
such as acritarchs, conodonts chitinozoas etc~ Study of microfossils constitutes a separate branch
in palaeontology called Micropalaeontology. Study of spore-pollen however forms a distinct
part within micropalaeontology, known as Palynology. However, it should be emphasized that
macropalaeontology, micropalaeontology and palynology share identical aim : to reveal the
organic history of the past earth and this can be done more speedily and efficiently when they
proceed together.
31.3 ADVANTAGE OF STUDY

Any fossil ·whether micro or macro has following importance to a student of palaeontology.
(i) It helps in recognition of the sedimentary bed in which it occurs. (ii) It helps in correlation
~f fossillif~rous bed~ occ.urring a~ different localities. (iii) It helps in dating the bed in which
~t occurs: (1v) It prov1d~s mfo~atl~n about environmental condition of that-time. (v) It provides
mfo~atlon about the hfe habit. (v1) It helps to reconstruct the evolutionary history of the past
organisms. ·
Ho~ever, the study of microfossils are preferred to megafossils in some cases due to
followmg reasons :
(a) Abundance: As microfossils are. minute in size they usually occur w1·th·m a roe k m · Iarge
be I
num r. n many cases, a small piece of a rock sample can yield a la be f fi ·1
individuals. The present-day trend is to study fossil-population wh· ~ge nu~ r.o o~s1;
information about the environmental cum ecologic control o • I~ ml ay g1v~ impod al n
bo t th · · f . n a part1cu ar species an a so
a u e mterpretatmn o evolution of a species and reco ·t· f . . h
t d i ·1 d . . . gm 10n o species lineages. Sue
as u yon oss1 s nee s detailed statistical analysis of diffe .
This becomes quite possible for microfossils for their a rent mor~ho.log1cal charact~rs.
in a succession. Collection of hundr d f . d" . bundance w1thm rock beds lying
e s o m 1v1duals of a · · ·n
case of microfossils. This is however species 1s not uncommon 1
• a rare event for megafossils
(b) Widespread occurrence : Due to very min t . . . · .
geographic dispersal even in global , Fu e si.ze. microfossils usually exhibits a wide
1
are easily transmitted or migrated fsea e. or their small m1croscop1c
· · size.
· the organism· s
barriers. Many microorganisms arero:/~e place t~ other overcoming several types of
~---
P agic by habit (mostly planktons) and this also
"'-
·-:'""'·
•--=-------
435
results their wide dispersal. Spore and pollen disperse through wind-current and by this
way spore-pollen of land plants may even be preserved within the sedi':'1e~ts of the. se~.
Thus microfossils exhibit a lot of index fossils which are globally dtstnbuted w1thm
their short-life tenure and help us successfully in regional, intercontinental correlation
and finding out the age of the beds in which they occur.
(c) Even distribution in sediments : Megafossils usually-exhibit patchy distributi?n within
rock beds. Depending upon the composition arid texture, some rocks may not yield well-
preserved fossils. Some rocks may not yield fossils at all. But because of the minute
size, microfossils usually tend to show an uniform/even distribution within a rock and
it is possible to obtain many identifiable specimens of a microfossil from any part of
the rock concern.
·(d) Occurrence within all types of rocks : A marine microorganism if occurred within a
particular geological time, its fossils would be available usually from all types of marine
rocks of that time in greater or smaller number.
(e) Occurrence of microfossils in rocks associated with coal .and oil deposits : The source
of such fossil-fuels are organic bodies as coal is derived from p!ant-debris and_the. sour~e
of oil is possibly some animals and in many cases, this are microanimals. Thus coal
and associated rocks become enriched in spore-pollens while various groups of
microfossils are found associated with petroleum bearing beds. Many oil-companies and
coalmine-authorities therefore employ a team of micropalaeontologists to learn more
about distribution of the fossil-bearing rocks they are handling. This commercial aspect
of micropalaeontology has undoubtly been a major stimulus to its growth. Apart from
these, there are however some philosophical and sociological sides to the subject. Our
understanding of the development and stability of present global· ecosystem has much
to learn from the microfossil record, since inany microfossil groups have occupied at
or near to the base of the food web. The importance of understanding microfossils is
further augmented by recent discoveries of microfossils from many Precambrian rocks
which were us~ally co~si~ere? unfossili~erous. With development of advanced techniques
of stu~y of m1crofoss1ls tt will be poss~ble to use these earliest records of organism to
estabhsh the ages of many Precambnan rocks whose ages so long are considered
controversial.
31.4 LIMITATIONS OF STUDY .

Though microfossils have more advantage over the megafossils in palaeontological t d
yet there are some limitations in using them. su Y
(i) The first difficulty is concerned with the proper identification of
. a m·scro&-,,0· ss1
. ·1· b
ecause,
~n most of the cases, they are very small, often not well preserved. This mak d" ff.
v1sua 1·1zat1on
in · · o f thetr . an d ot her morphological features es h"I hICU. 1ty
· exact shape, size
ca use d"ffi . . "d ·ri .
1 1culty m their proper I entt 1cat10n.
, w 1c may
(ii) Fragments of mega-organisms often treated as microfossils, exhibit variable types of

sh~pe and size, and it often becomes difficult to correlate these fragments with the
animals or plants to which they were related.
•
PALAEONTOLOGY
436
, (iii) Microfossils, as they are minute in size, are rarely prese.rved as in situ fossils; rather
most of the microfossil assemblages are thanatocoenose type. Thus taphonomic factors
should be kept in mind before any type of conclusion about their morphology or ecology.
(iv) Microfossils, for their small size often act as elastic grains within a sedimentary rocks
and like other elastic minerals they may occur as reworked products i.e. may suffer more
than one cycle of sedimentation. Caution should be taken to identify these reworked
fossils within rocks. Again because of their small size, microfossils of an overlying
horizon may percolate through the rocks along with subsurface water downward to be
collected within the older underlying rocks creating confusion about the age of the host
rock.
(v) Most of the microfossils include lower grade of organisms which frequently show
alternation of generation and two separate morphologic types corresponding to each
generation. In such a group now extinct, these two morphologic-variations are often
erroneously identified as two different species. This frequently happened in case of
benthic larger foraminifera, many of which are now extinct, like Nummulites.
31.5 CLASSIFICATION
The entire microfossil-world can be classified into three broad subdivisions :
(a) Microscopic organisms - Protista
(b) Microscopic parts of mega-organism
(c) Microfossils of unknown affinity.
_Protista group forms a distinct phylum in organic kingdom that includes II · ·
ammals and plants and some still primitive groups behaving both like . a I m1cdros,cop1c
Whittaker ( 1969) l"k . . . am ma s an p ants.
• e to propose a d1v1s10n Monera which in t d · ·
organisms i.e. have cells lacking true nucleus, cell vacuoles and co~h es s;~glle-celled ?rokaryot1c
pseudopodia etc. It has two subdivisions. er ce e ements hke flagella,
1. Cyanophyta (Blue green algae)
2. Schizomycophyta (Bacteria)
All other microorganisms are showing eukaryotic cells wi . .
e~ements. They are mostly mobile unicellular organisms wi th d1s~mct nucleus and other cell
dmoflagallates, have whip-li~e flagella for locomoti d th _vaned body plans. Some, like
autotrophic like plants and they may lie close to thon an contam photosynthetic pigments thus
radiol · are protozoans (sarcodina) which develo
. . ana e ancestral .I ine of amma
· Is. Foraminfera and
(Cihophora) have a coat of bristle-like cilia a d P mobile pseudopodia while tintinids
are all heterotrophic protistas which are more·n k_mouth ~urrounded by small tentacles. These
a 10 to ammals th 1
A. Microfossils (Protista) : an P ants.
(i) Pyrrhophyta - Dinoflagellates
(ii) Chrysophyta - Silicoflagellates d"
C") c· . , •atoms and coc 1· h
111 1hophora - Tintinids a d C . . co It ophore
(iv) s . n alp1onelhds
I
arcodma - Radiolarias and Foram· .r
~ m11era
- ,~ - ... . . --==- " . . , ..... - ·
....:r' . ., .. _..
JNTRODUCTION TO MICROPALAEONTOLOGY
437
B. Microfossils (parts of mega-animals and plants) :
t. Multicellular : green algae (chlorophyta), red al ae (rhodo h ta ,
(phaeophyta); bryophytes and fungi. g P Y ) and brown algae
2. Tra~heophyta : Spore/Pollen of algae-fungi, bryophytes, pteridaphytes gymnosperms and
angiosperms. '
3. Microscopic parts of annelids : Scolecodonts.

4. Microscopic anthropods : Ostracodas and Estheriids.
5. Microfossils of uncertain affinities : Conodonts, Chitonozoans, etc.
31.6 MICROSCOPIC ANIMAL FOSSILS

31.6.1 Foraminifera
A. General characteristics :
Foraminifera belongs to the phylum protista and includes a group of single-celled animals
almost all of which living either on sea-floor as benthos or within the sea water as planktons.
Commonly most of the animals of this phylum are soft-bodied and are uncommon in fossil
record. But foraminifera and a few other protistas secrete some sort of hard skeleton called
test enclosing the soft protoplasm. The test is usually composed of minerals (calcite/aragonite)
organic matter (chitin, tectin) or agglutinated particles. The test consists of a single or multiple
chambers interconnected with each other by an opening (/oramen) or several openings
(/oramina) from which the group takes its name. Foraminiferas range from Early Cambrian to
Recent. Foraminiferas are important constituents of marine sediments. Many organic; oozes are
composed of more than 90% of planktic foraminiferal dead-skeletons especially of Globigerina
(called Globigerina ooze). Bulk of many Tertiary marl beds are made of abundant fossil
skeletons of foraminifera such as Nummulitic limestones. Foraminifera fossils have been
successfully used in biostratigraphic classification and age-determination of rocks especially of
Cenozoic times, largely because they are abundant, diverse and easy to study. Planktonic
foraminiferas yield several index fossils and many of them are geographically widespread and
have short vertical ranges as they had rapidly evolving lineages. These factors greatly help in
intercontinental correlation of beds. As their developmental stages of life are preserved within
the test, they could be used as excelJent tools to evolutionary studies. Ecological sensitiveness,
especially of benthic groups make them useful in the studies of recent and ancient environments.
B. Living foraminifera (Fig. 31-1 A) :

A living foraminifera comprises a single cell with protoplasm divisible in two parts : an
outer light coloured layer of clear ectoplasm and an inner darker portion called endoplasm.
The outer ectoplasm surrounds the test of the animal and gives rise to numerous thread like
branching pseudopodia which usually function in locomotion, anchorage, food capturing and
excretion. Endoplasm portion of the cell occurs within the test and exhibits food vacuoles and
nucleus. Number of nucleus may be one (uninucleate forms) or many within the cells of
multichambered forms (multinucleates forms). Foraminiferas feed by trapping very small
organisms or detritus/organic particles by their sticky pseudopodias. The food materials are
digested outside the test within food vacuoles which then pass to ectolasm through the aperture
PALAEONTOLOGY
438
~~~~~~~~~~f---;;p~~~=== Test-wall
Pse udopodia
;y___...,..,tflli&:..--- Outer ectoplasm
. - - ~ ~ - - ~ - - - - - - - - - Food-particle lrnppcd
, t,y pscud9podiu
A . A LIVING FORAMINIFERA
Megnlospheric fonn
Microspticric form (Gumont)
Ma1ured
(Schizonl)
Microscopic
Large i n i t i a l ~
chamber~
Equatorial Sec1ion
Equatorial Section
Growing gamont
Young schizont
Ma&µn:d 11chizont
R. LIPE CYCLE AND ALTEMATION OF GENERATION
FIG . 31-1 : FORAMINIFERA AND ITS LIFE CYCLE (DIMORPHISM)
INTRODUCTION TO MICROPALAEONTOLOGY 439
of test ·md absorbed by protoplasm. Undigested and excreta-products along with food vacuocles
then pass outside the test.
A formninifera, like many other lower groups of animals and plants: exhibits alternation of
two forms in its complete life history : a form showing gamont generation reproducing asexually
and a form of scliizont generation reproducing sexually (Fig. 31-1 B).
The schizont representing diploid generation, asexually reproduces when the cell within the
test splits meotically into nmnerous daughter cells each with a nucleus with half the number
of chromosomes originally found in parent cells. These daughter cells are then released into
the water to disperse and eac~1 forms a young gamont. The production of gamonts from schizonts
usually takes place in winter months. Within the coming summer · months the gamonts become
matured when cell of each gamont again divided but this time through mitosis thus producing
numerous haploid gametes each of which bears a pair of whip like flagella. and-they are released
from the parent test. Usually a gamete produced from one individual, sexually unite with a
gamete of another individual and forms a zygote. The zygote grows into a new diploid schizont.
Morphologically, gamonts and schizonts are dissimilar (hence dimorphic). Gamonts are
usually more numerous and become small-sized but with a relatively large initial chamber hence
called megalosplieric forms. Schizonts on the other hand are less common forms, large in size
and with a very small initial chambers (hence called microsp/zeric forms). This has caused a
lot of difficulty in the identification of these two dimorphic form·s in their fossilized stage. This
results in many cases separate names for the dimorphs O.~ a single -s pecies.
C. Skeletal Morphology of fossil foraminifera :

1. Larger and Smaller foraminifera
Foraminifera fossils are usually ·g rouped as (i) Larger foraminifera and (ii) Smaller
foraminifera. Larger foraminiferas are usually visible in naked eye and may range from a few
mm to more than 5 cm in diameter but their identification upto generic and specific level can
properly be done only by observing them thin sections under microscope and for this, oriented
thin sections of their fossil tests have to be prepared. Smaller fon.uninifera are mostly of
microscopic size ( 100 to l 000 µ) and are seen and- ide1itified only .':Jnder a microscope as three
dimensional individuals.
2. Composition and structure of test-wall (Fig. 31-2)

Composition and structur~ of test-wall is an important basis of classification of foraminifem.
On this basis test-wall may be of three types.
(a) Proteinaceous or pseudochitinous : Test wall is thin, non rigid, composed of oroanic
matters often called tectiu. Tectin is present in all har<l-w.,lled tests as u 1inino 1~ the
0
chambers acting as a template for minernlization e.g. Lage"''· Allogromina.
(b) Agglutinated : Here the test-wull is made up of organic nn<l mineral mutters derived
from sea-floor which are bound together by some organic, culcnreous or ferrugint!ous
cement e.g. Textularia.
(c) Calcareous test : Here the test is made up of entirely inorganic caco 3 secret ti by the
animal itself. There are three types of calcureous tests.
Pah,c(Gcu)WP S6
PALAEONTOLOGY
440
Outer layer of
foreign particles
Inner organic lining

Flexible wall of organic skeleton
b. Agglutinated wall
a. Tectinous (Pseudochitinous) wall
1 , 1 1 1 1 Ii I f:At ordered outer layer ft c? O~ O ~;~~'4

~P~O ~jJM, [.__ Middle layer with dI)!"!)~ ..!! \'i4~0
~ Q~ 0\l ;j.Ji~lJ ~ random CaCO:\ cf')'.stals
U\t ~I) '\"'.ffl10°.,'lJ in organic matnx
~~~A \ ff. y
~\ ii I
c:,
I ~
~.~iii j \liiii( . __--. .
J1?
Innermost organic lining
a........_Ordered inner layer Innermost organic lining
c. Catcareous porccllaneous wall d. Pseudopunctate wall
Microgranular layer
-Mic'."granular
calcite Fibrous layer
Nonlamellar Lamellar
c. Microgranular wall (lnpcrforated)
• att~~::i~;~:i,
f":Porc~
Organic lining
f. Hyalinc/glassy (Perforated) wall
FIG. 31-2: MICRO~STRUCTURES OF TEST-WALL (a-f)
Unilamellar
Multilamcllar
FIG. 31-3: MODE OF ADDITION OF CHAMBERS WALL
--·
INTRODUCTION TO MICROPAI.AEONTOLOGY 441
~
r;;~
0 Sac-like
Pleurophuy:c Flask-shaped
Lagt/lCJ
~® ....,,,,ho,
Planispiral-discoidal
Ammodiscu.f 4 ~
0
~~~bular
Rhi za mmina
Globose
Glomospiral
Hemispherical
Irregular radiate Zigzag (Vennicular) (Sessile)
A.urorhizu A111111011erte/lu Hemispherumminu
a. Different types of Unilocular Tests
Curved
Nodmaria
With equitant
Aperture terminal
chambers
Kn,hopyxu
Nodosariu 1 Uniserial
Triserial
Eggerella
Biserial
Bolivina Biserial & Uniserial combined
Bige11eri11u
b. Uncoiled/Multilocular Tests
@~
Transverse section ~
Triloculine
~ Apertural view Triloc:uli11a
Quinquelm:ulina Biloculioc
Spiri,culimJ
c. Coiled Multilocular Tcsts-Miliolinc Coiling
FIG. 31-4: TYPE OF TEST AND COILING
442 PALAEONTOLocy
~iA
A:: - Transvcrse sectrnnal
. view
Umbilicus~ Apcrtural
view .i chamber
(Basal .
symmetrical)
External
appearance
A:
Equatorial
.
A: .:
section -~ - A x i a l section
chamber
~chamber
iA
Pla11ispiral-Evolute Pla11ispiral-Co11vo/ute
a. Planispirally coiled tests (A-A : Axix of coiling)
. ~ - A ; r t u r e ..
A-~ 1..4·• ..,""""'-+- Aperture .

Apertural-side
Spiral/dorsal view
view Umbilical/ventral view
b. Trochospirally coiled tests
Initial biserial
later uniserial
Initially coiled but
later annular chamber
Early low Trochospiral

later planispiral
Planispiral-Uniserial
c. Abnormal ·type· of lest
FIG. 31-5: TYPE OF TEST (MULTILOCULAR) AND COILING
••
.,-.,.----Later chambers (equatorial)
Nepionic/Periembryonic chambers
l
2nd chamber (Deuterocanch) Embryonic Nephrolepidine
-(-~"Pr-- lst chamber (Protoconch)
a. Growth of chambers from embryonic apparatus

f
apparatus
lsolepidinc I <I Polyl<pidi"'
(Equatorial section) b. Type of embryonic apparatus
FIG. 3 J-6 : ONTOGENIC DEVELOPMENT OF CHAMBERS
INTRODUCTION TO M!CROPALAEONTOLOGY 443
(i) Porcelaneous : Test wa11 appears milky white in reflected light and amber coloured
in transmitted light and composed of tiny needles of magnesium-rich calcite
randomly arranged; usually lined by an outer and inner layer where the needles
are orderly arranged (impunctate/Pseudopunctate) e.g. Triloculina.
(ii) Imperforate-microgranular : Wall of this test appears dark in thin section and opaque
in reflected light. This wall is composed of minutes granular crystals of calcite
arranged randomly or at normal to the surface giving the wall a fibrous appearance.
e.g. Fusuli11a.
(iii) Glassy or perforated : Many foraminferal test-wall appears glassy (hyaline) when
viewed in reflected light. Here calcite crystals either randomly or radially arranged
with surface. Wall of most of these foraminifera are traversed by small, straight
pores or branched alveoli through which pass fluids linking ectoplsm and endoplasm
e.g. Rotalia.
3. Arrangement of locula/chamber
Foraminiferal test may consists of a s_ingle ~hamber called unilocular or more than one
chamber (multilocular). In a multilocular form when the wall of the previous chamber is not
overlapped by the wall of the new chamber the arrangement of chambers is called unilaminar
type. But in other cases wall of each newly form chamber overlaps the wall of all the preceeding
chamber and that gives a multilaminar arrangement of chambers (Fig. 31-4 ).
Unilocular tests : A single chambered tests is usually simple tubular (opening on one or
either side) (e.g. Bathysiphon) or branched tabular (Rhizammina) or coiled planispirally
(Ammodiscus) or trochospirally (Ammovertella) showing a spherical or subspherical shape (e.g.
Lagena). This single chamber may be stellate, or variously branched (Astrorhiza) (Fig. 31-4a).
Multilocular tests : (a) Uncoiled (Uniserial and Biserial) : In a uniserial test chambers
are arranged along the growth axis in a single series. Each chamber has a terminal aperture
which results in growth of successive chambers one above the other along the line of axis (e.g.
Nodosaria). In biserial test two chambers are found 180° away from each other forming two
series. Aperture of each chamber is basal and lies alternately in opposite direction thus producing
the two series chambers lying in a zigzag manner e.g. Textularia (Fig. 31-4b).
(b) Coiled test: These are of two types (Fig. 31-Sa-b).
(i) A planispiral test arises when the successive chambers are arranged spirally around the
growth axis but all lie in a single plane and the rate of translation of aperture is practically
zero. A complete revolution ~round growth ?xis forms a whorl and each whorl may contain
more than one chamber. With the format10n of successive whorls, chambers on outer
whorls go more and more away from the axis. Apertures in successive chambers in this
case, lie ~ym~1etrically at t~e base of each chamber. A perfectly planispiral test is bilateral
symmetrical m apertural view.
(ii) A helicoi~/tr~h.ospiral test _arises when chambers are arranged spirally around the nxis
but there 1s distinct translation of the plane of coiling after each revolution so that the
test becomes asymmetrical. Such a test has always a spiral/dorsal side (i.e. the side from ,•.
r
444 l'ALA EONTOLoc y
which growth starts) and a ventral/umbilical side (i .e. the side where lies the last
with last rows of chambers including the aperture). Aperture in this case, lies ;h;i
base of each chamber but in a slight assymmetric manner causing lateral transla· t· e
, ion of
each chamber.
A biserial or triserial test may be derived from a trochospirally coiled test where each wh 1
. . I or
is composed of two/three chambers with much decreased va Iue o t sp1ra angle or from planispiral
test by turning the successive chambers through an angle 180°.
The growth plan of all these tests may be modified further by the rate of chamber expansion
and the successive chamber-height increases with continuation of growth. Depending upon the
tightness of coiling, the test may be evolute, involute or convolute (semiinvolute).
In evolute test, all the whorls and chambers are visible externally. But in an involute test, the
preceeding whorls are partly enveloped by a newer whorl. But in convolute test the last whorl is
only visible externally and that usually covers all the earlier whorls and chambers. In a semi-
involute test, the earlier formed whorls become involute/convolute but the later whorls are evolute
(Fig. 31-Sa).
(iii) There is a special type of coiling which is found only in some formaminifal test. This
is called streptospiral or milioline coiling (characteristics of miliolidae group). In this
case, the test is made up of a continuous spiral and chambers are added at angles of
72° leaving five chambers of a whorl visible from outside (e .g. Quinqueloculie). The
chambers are added at angles of 120° when three or two chambers are visible from
outside called triloculine and biloculine. In milioline coiling, the aperture of successive
chambers lie at the base but alternately in opposite side to maintain the planispiral plan
(Fig. 3 l-4c ).
While in a trochospiral form chambers are aligned along growth axis, in a planispirally coiled
test the chambers are usually aligned along a plane at right angle to the axis often called
equatorial plane and the chambers in this plane are called equatorial chambers. In case of
convolute type of test, where each earlier whorl is completely enveloped by the next younger
whorl, there is sometimes a space in between two whorls which are just lateral extension from
each equatorial chamber often called lateral or afar prolo11gation as found in the group
nummulitidae. Here sometimes the marginal/peripheral wall of each whorl become excessively
thickened by shell-material accumulation forming a thickened marginal cord (Fig. 32-4).
Besides, there are uncoiled test where chambers are themselves annular or often they are
arranged in annular rings. One ring completely or partially encircling the proceeding one.
In a planispirally coiled test, sometimes increase of height of chambers in successive whorls
result in the development of an axial depression towards the growth axis which may be shallo~\I
of deep depending upon the rate of increase of height of outer chambers. This depression is
called umbilicus which sometimes may be filled up by shell-materials forming an umbo on
either side of the test (Fig. 3 I -5a).
4. Different growth stages in ontogenic development

From ontogenic point of view, a foraminifera test usually exhibits 5-different stages whi~h
can be recognised as (i) Embryonic (ii) Nepionic (Periembryonic) (iii) Neanic (iv) Ephebic
and (v) Gerontic (old).
445
These stages can be e~si)y distinguished where there is some difference in morphology of
type of chambers and their arrangement and usua1ly 3 types of multiform growth are noted.
(i) Initia11y planispiral/trochospira) chambers but biserial to uniseria1 at adult stage.
(ii) Chambers initially coiled, later show rectilinear arrangement.
(iii) Spirally arranged earlier chambers followed by annular, branching or other irregular types
of growth in adult stage.
However, all possible types of combination are not always found. For example ontogenic
change from initial rectilinear to uniserial and from biserial to spiral growth have never been
observed. Again annular growth never leads to uniserial arrangement. The different ontogenic
stages of a particular species may indicate the ancestral form from which it is derived and that
could be a clue to its evolution. In this respect microspheric forms are more useful as they
record more completely these different stages.
The first chamber developed from embryo is called protoconch which is followed by a
deuteroconch or the second chamber. Protoconch and denterconch together constitute the
embryonic chambers/apparatus representing the initial stage of ontogeny. Chamber/chambers
just overlying the embryonic apparatus is/are called nepio11ic/periembryo11ic chamber/s. All other
chambers represent growth of adult stage (Fig. 31-6a).
According to relative size of protoconch and deuteroconch and the area of protoconch
encircled by the deuteroconch, the embryonic apparatus of some orbitoid groups such as
Lepidocyclina exhibit a few types viz : deuteroconch smaller than protoconch (polylepidi11e),
equal to protoconch (isolepidine), larger than protoconch encircling more than half .of its
circumference (neplirolepidine) and enclosing the protoconch almost entirely (eulepidine) (Fig.
31-6b).
S. Secondary chambers/lateral chambers

Very often seco11dary chambers are formed by subdivision of primary chambers and in many
cases it is possible to distinguish primary and secondary chamber in a foraminifera test. It is
often found that annular chambers at its initial stage remain unsepted but later chambers become
transversly septed and develop small secondary chambers or cliamberlets. In some forms
equatorial chambers are also longitudinally septed producing chamberlets.
A number of larger foraminiferas belonging to orbitoid group such as discocyclinids,
lcpidocyclinids and miogypinids develop a subsidiary system of lateral chambers over and below
the chambers lying in equatorial/median plane where normal equatorial chambers are found
sandwitched between the two zones of lateral chamber-layers over and below them (Fig.
32-5). Within the test, lateral chambers are usually arranged in multiple tiers, the number and
size of which are usually increasing from periphery to centre of the test· muking such test nearly
lenticular in vertical section. Such lateral chambers are clearly seen in vertical section as elliptical
or slit-like or rectangular in outline but three dimensionally they are tabular with irregular in
outline as seen in tangential section (Fig. 31-11 ).
. . ~ ....
- ·- - ... . ,;, .
44fi PALAEONTOLocy
0 Oo,lollular
( Orlml i1111 >
Snc•likc
(A I lo>:n 1111i1w)
< ~
~
Tubular
(Buthysiplum)
))
Tubular-branched
~ Vennk,lo,
(Amoverfel/u)
Discoidn 1-stl!l lntc (Rhizammi "")
Oo o=·· li""O Triangular-

elliptical
t llit S\11dal Oiscoidal-circular ~ (Textularia)
(A h·eo/i,w) (Discocyc:lina) Lenticular Fusiform
(N11mm11lites) ( Q11i11queloc:11/i11u)
Conical (Rora/ia)
Oiscoidal-elliptical
--->
~o
Saddle shaped-circular
6
(Miogy/>.fina) (Discocydina)
Flask-shaped
(Lage,w)
FIG. 31-7 SHAPE OF TESTS
~ 8 ({])
Globular Subspherical Tubular
( (l]fl)
Tubular
rectangular Pyramidal
a. Shape of chambers in three-dimentional view
Q) ~
c;re,n,(\ s,bcire,n"
~.~ C~ohc • Ogoval
~~ Spatulutc Hexagonal
RcctnogulM ILinear
Annular
b. Shape of chambers in 1hin seccion
c. Change of cqua1oriul
eharnbcr 's shnjXI with Lepid0<:ydi1u1
n101cnic developmcn1
FIG. J l-8 SHAPE OF CHAMBERS
JNTRODUCT'ION TO MICROPALAEONTOLOGY 447
6. Septa and scptal suture (septal filaments)
In all multilocular test one chamber is separated from the adjacent one by means of a
common wall lying in between them which is called septum . There are one/more foramens
across this septum joining the two adjacent chambers. The external junction line of each septum
with the wall of the test is called septal suture or septa/ filame11t. Normally the suture lines
may be depressed or elevated, straight or curved. Sutures between the two adjacent whorls is
called spiral suture and those between the chambers of the same whorl are called .teptal sutures.
The suture marking junction of apertural face with preceding whorl is called basal suture.
Septal filaments in Nummulites (31- tob) are variously modified and show a great taxonomic/
evolutionary significance. In earlier species of Nummulites these are simple radial lines on the
surface of test but the Middle-Late Eocene forms exhibit sinuous filaments often called sigmoidal
suture. Still younger species exhibit meandrine type and subreticulate type of septa! filaments.
In meandrine, type the filaments are grouped in bundles running fairly in a straight line mostly
ending poleward abruptly against another bundle. The subreticulate and the typically reticulate
septal filaments are characteristics of Oligocene Nummulites where the radial filaments show
repeated bifurcation producing polygonal meshes on the surface.
7. Shape of the test (Fig. 31-7)

External shape of the test is usually a function of the arrangement, nature of growth of
chambers and its rate of increase in 3-different dimensions. Accordingly, there are quite variable
types of shape of the test of foraminiferas. Some common types are : lenticular (circular in
outline, biconvex in side view), globular, subglobular, ellipsoidal, fusiform, discoidal, tubular,
conical, subconical, stellate, flask-shaped etc.
8. Function of the test

It appears that the test of a foraminifera mainly protects the inner enclosed endoplasm
reducing physical, chemical and biological stresses. Biological stresses include attack by
suspension and deposit feeders, infestation by parasitic nematode, worms etc; physical stresses
include harmful radiation of sun, water turbulence etc. Chemical pressure comes from fluctuation
of pH, CO 2, 0 2 and inorganic nutrient levels in the sea-water. In unfavourable condition
protoplasm can be withdrawn into the inner chambers. Additional advantage of the test includes
the stability it gives to the organism especially for spinose, thick-walled or agglutinated forms.
Without a shell, the animals find it difficult to control the biomass and regular cell-division.
Surface sculpture may also help in various way to assist buoyancy, to improve adherence to
strengthen the test against any mechanical stress and also helps to channel ectoplasm to flow
to and from the apertural pores and umbilicus.
9. Shape of chambers (Fig. 31-8)

Shape of chamber is also a variable feature that depends upon the pattern of growth of
each chamber along its three dimensions. In three-dimensional view chambers are recognized
as the following types : spherical, subspherical, tubular, annular, pyramidal, ovoid, cylindrical.
prismatic, stellate etc.
More commonly chambers are viewed and described from the study of oriented thin sections
especially in case of larger foraminiferas where chambers show u wide range of varia1ion in shape
Palae(Gco)WP-57
448 PALAEONTOLocy
8
0
Terminal circular
(Nodo.rn ria)
® (})
Terminal necked Terminal radiate Term inal slit Terminal long slit
( UviKerina) ( Le11tic11 li11a) (Pyr1:o ) (A rriculina)
Q) m
(Bu;
Basal-looped
Basal-hooded
(Paraftw,ri11a)
Basal-cruciform
(Cruciloc:ulina)
Basal-arched
(Nonion)
Basal-toothed
(Qui11<J 1telorn I ina)
Basal-slit like Tenninal-cribrare

(Melonis) (Pomep,mides)
Basal-multiple circular
(Elpltidium)
Basal-umbilical Basal-toothed Basal-circular Basal-u mbi Iical "Basal-extra-umbilical

circular circular umbilical with with umbilical plug tear-shaped
(G/obi,:erina) (Ammonia) umbilical lip (Parmta/ia) (Rotalia)
(Romrbitoides)
- - ~ - - Terminal
rear-shaped
Basal-slit like (multiple) . . , . - - - Supplimcntary

extra-umbilical apenures
(Di.fcoxbi.f)
Dcndrilic
(Demlritina) ( \Ii 1;~11 Ii"""")
Multiple-areal
apenurcs
(Orbuli11a)
f!E
Extra-umhilical double slar-shnpcd apenurc
(D11o.ft1>111i11a)
FIG. 31-9 : APERTURE OF FORAMINIFERA
~ -.
INTRODUCTION TO M/CROPAl.AEONTOLOGY
449
in two dimensional view. Some common shapes of equatorial chambers are as follows : circular,
subcircular (mos~ protoconch and dcuteroconch). arcuate, ogival, rhombic, spatulatc, hexagonal.
retangular etc. (F,g. 3 l-8b). Lateral chambers appeared in vertical section as rectangular, elliptical
or slit-like while in tangential section they appear irregular or polygonal in outline. It is often
found that shape of chambers is changing with ontogenic development as found in Lepidocycli11a
where in some forms initial equatorial chambers are arcute/ogival or rhombic but at adult stage
they become hexagonal or spatulate (Fig. 31-8c). In larger foraminifera, actual shape of a chamber
should be understood from their appearance in different oriented thin sections.
Internal structure or shape of chambers also provides taxon~mic clues in identification of
different genera of the same group. For example, in Discocyclina equatorial chambers become
only rectang1,1lar whereas in Lepidocyclina, equatorial chambers may be of different shapes bu1
never rectangular.
10. Aperture (Fig. 31-9)

The morphology of aperture of foraminifera has a great taxonomic value. The term aperti,re
should be applied only for the single or multiple openings found in the distal wall of the last
chamber. This usually connects the ectoplasm bearing pseudopdia with the endoplasm lying
within the test allowing passage for food and excreta and release of daughter cells. Its position
usually remains more or less constant in a particular species throughout its ontogeny. The term
foramen (single) or forami11a (multiple) on the other hand, is used to denote the internal passage
between successive chambers. The term stolen is restricted to a type of foramen which is a
passage with tubular projection as found in orbitoidal foraminifera. The position of the aperture
is the most important character in relation to formation of new chambers, and hence uhimately
in determining the type of growth plan of the chambers whether they will be uniserial, biserial,
planispiral or trochospiral or form an uncoiled test.
As regards the position, the aperture may be terminal, peripheral, basal, or central. Its shape
may be circular, subcircular, elliptical, slit-like, dendritic, bottle-necked etc. Multiple apertures
may be arranged radially or may occur along a line. Sometimes aperture may be represented
by a large number of small perforation on aperture face often called cribrate. as found in Eocene
genus Cribrohankenina. Multiple aperture obviously result the formation of multiple chambers,
all at a time from a single parent chamber.
In most simplest forms like most of the uniserial tests, aperture is a circular opening at the
terminal on distal face of the last chamber. Successive chambers then grow one after another
along the single growth axis producing an uncoiled uniserial test. Basal aperture may result in
biserial forms when they situated alternately in two opposite directions. In spirally coiled
chambers, aperture is placed at basal or peripheral margin of apertural face of each chamber
which develop in the same direction. When this basal aperture is symmetrically placed, the
result is the planispirally coiled chambers but if it shifted from the medial plane it causes
translation of each chamber from the plane of coiling resulting trochospirally coiled chambers.
Thus in a trochospirally coiled test aperture may be slightly or strongly eccentric often lying
near umbilicus (umbilical), as found in Globigerina). Aperture may be further modified by the
presence of several accessory structures such as a lip or flap-like feature (often called labiate
aperture), a tooth-like structure (dentale aperli,re), a central cover plate (b11llate aperture) or
a central elevated boss.
- a
450
PALAEONTOLOGY
Cancellate Fine spines Peripheral flange
Central boss -t-";"'11~~'L·~··I"

(mamelon)
~ Perforation
Sipho11i11a ~ Polygonal depression Di.fcocydi11a
Globigerina
//"';~,{"'\...,,, Longitudinal
costae
Thick spine
Calcarina Hanrkmi11a
Hastigerinoides
Rectobolivina
a. Ornamental features of foraminifera
Sigmoidal Reticulate
Radial Meandrine
b. Variation of septal suture (filaments) in Nummulites
FIG. 31-10 : (a) ORNAMENTAL FEATURES AND SEPTAL FILAMENTS OF FORAMINIFERA
- - Pillar ending to
surface spine
Inflation pillar
Incised pillar Texturnl pillar

Pillar Cavity Later.ii chambers
~:5
I !
Ci£=f
Pillar
Equatorial
Residual pillar chumbcr layer
Pseudopi llar
FIG. 31-1 J PILLARS IN THIN SECTION (Vc::rtical sectional views)
INTRODUCTION TO M!CROPALAEONTOLOGY 451
11. Surface sculpture or ornamentation and pillars (Fig .. 31-10)
The external surface of foraminiferal test may bear variable types of ornamental features
such as spines (spinose) keels (carinate), fine straie (striated) and coare costae (costale), granules
(granulate) flanged and reticulate suture etc. These features may be used for taxonomic purpose
with caution as they frequently vary through ontogeny and with change of environment.
Small, raised and elevated bodies on the test surface are mainly of three different size :
pustules, granules and papillae. A pustule !~ the largest type of granular structure usually found
at the central part of _the test as a single elevented body often called central or polar pustule.
Granules and papillae are still finer raised bodies usualiy uniformly distributed over the test-
surface. All these granules are again of two types : superficial (simple external features) or
external expression of some internal structures called pillars. Spines may occur as tine hairs
all over the test. Calcarina and Hantkenina develop thick peripheral spines. He.rtigeri11ella
exhibits needle-like spines from periph~ral zones. of each chµmber. Keel or flange is usually
found as a ridge or extension on peripheral zone as found in some lepidocycli11a, Di.rcocyc:lina,
and also in Lemiculina, Discorbts, Gi~botrunana,· Globorotalia etc. Costate of tests are shown
by Rectobolivina (longitudinal ~o~tae). H~terohelix exhibits longitudinally straited test. In ma~y
cases, the septal suture lines become thickened and appear as elevated ribs. Most of the
ornamental features specially keels and spines help the planktons in their flotation.
Many granules on the foraminiferal test are the external manifestation of some internal feature
often designated as pillars which are seen only in thin section of the test. These pillars are of
five types (Fig. 31-11) :
(a) Inflation pillars : These are forme~ by local thickening of wall along a radial
· (transverse) line where each laminae contributing to this extrnthickening. This inflation
pillars often appear on surface as pustules or as pointed spines.
(b) Incised pillars : Incised pi11ars are formed by portion of shell-material isolated from
.the . rest of the test developing into anatomosing fissures thus separating tine tubular
blocks of shell which ext~rnally are raised forming granules in between the fissures.
(c) Textural pillars : These pillars are marked by difference in texture from the rest of
the test and usually externally they do not produce granules.
(d) Residual pillars : These piJlars are cylindrical masses arranged in between lateral
chamber tiers. They are also devoid of any external granules.
(e) Pseu~opillars : Sometimes external loca·l thickening of the wall of lateral chambers
gives· the appearance of pillar-like structures, called psetidopillars. These are specially
observed in orbitoidal foraminifera (Lepitlocydina and Discocycina).
Combination of these pillars ar~ also found within a test ~uch as inflation-incised piJJars, .
inflation-textural pillars etc. ·
A special type ornamentation is found in Globigenina test surface which is marked by some
polygonal depressed areas separated by smooth zones. There polygena~ areas are uctually su~~e-
.
depress1ons . a small circular
surroundmg . .
perforations , " . ) . Th'.
(punctate sur,ace ·. called canceual•
1s 1s
452 PALAEONTOLOGY
type of ornamentation. Minute perforations on many pJanktonic test is a characteristic

ornamentation which make the test lighter and helps in buoyancy.
12. Canal and fissures

Canal and fissures are typicalJy observed in the wall of perforated calcareous test of
foraminifera.
Canal systems are comprised of essentially some fine tubular cavities bore within cell-wall.
Radial canal connects an aperture or opening of the chamber with the exterior where this radial
canal is later used as an exit for the protoplasm that will supply the content of a new chamber
while the radial canal of the earlier chamber is converted into a stolon.
It is assumed that formation of a radial canal is due to the emergence of a protoplasmic
current from the test. Ramifying canals are typically finer openings than the radial canals. They
consist of short passages across the shell laminae but elongated more or Jess parallel to the
laminae. This results from non-deposition of shell materials for .the occurrence of their fine
protoplasmic currents from the external layers of the protoplasm. If one such protoplasmic
current remain active the canal may continue from one chamber to the next but if it stops, the
canal would be terminated. Often such protoplasmic current may anatomise increasing the
gradual complexity of the canal system.
Regarding taxonomic significance of the canal system in foraminifera, there is diversity in
opinion. The assessment of phyletic significance of canal system is not always easy. However,
it is usually used as the basis of classification of higher taxonomic unit such as a family or
above the family.
Classification
Earlier classifications of foraminifera were proposed by several poineer workers such as de'
Orbigny ( J826), Brady (1884), Galloway · (1932), Cushman (J 959) and Glaessner ( 1944).
Galloway (1932) proposed 35 families within the order.foraminifera based on his interpretations
about the phylogeny of this group. Cushman ( 1948) in his last classification proposed 50 families
within foraminifera. Glaessner (1944) divided this order into 7 superfamilies and 37 families.
These superfamiJies are : (1) Astrorhizidea (2) Lituolidea (3) Endothyridea (4) Miliolidea (5)
Lagenidea (6) Buliminida (7) Rotallida. Loeblich and Tappan ( 1964) took into account some
features of foraminifera in their classification. These are (a) wall structure and composition (b)
shape, number and arrangement of chambers (c) position of aperture and its shape (d)
ornamentation of test. Brasier ( J980) recognised 5 suborders within foraminifera. These
suborders are (i) Textulariina (includes superfamilies Astrorhizidea and Lituolidea) (ii) Fusilinina
(includes superfamily Endothyridea), (iii) Rotalliina (includes superfamilies : Lagenidea,
Bulininidea and Rotalidea) (iv) Milioliniina (includes superfamily Miliolidea and Allogromiina).
The last one is a new divisions that includes some tectinous skeleton-bearing foraminifera,
mostly unknown as fossils. Some diagnostic features of each of these suborders are given below:
(i) Allogromina (Cambrian-Recent) : Benthic, organic test; unilocular with terminal aperture;
chamber tubular/globular/ e.g. : Allogromina.
UCTION TO MICROPALAEONTOLOGY 453
tNTROD
(ii) Textulariina (Cambrian-Recent) : Smaller benthic forms; non-laminar, aranaceous,
agglutinated wall, unilocular/multilocular, chambers tubular, globular, uniserial/biserial/
coiled/branching, aperture terminal/basal; e.g. : Textularia.
(iii) Fusuliinina (Palaeozoic-Triassic) : Benthic, non-laminar calcareous microgranular/fibrous

wall, imperforated or finely perforated; multilocular, chambers tubular, subdivided into
chamberlets; chamber-wall secondarily thickened e.g. Fusulina.
(iv) Milioliinina (Cretaceous-Recent) : Benthic, calcareous, porcellaneous, imperforated;

proloculus initially planispirally coiled, subsequently may be uncoiled, streptospiral/
milioline; In the last case, 2, 3 or 5 chambers may constitute a whorl, chambers tubular
with terminal aperture, often divided into chamberlets e.g. Triloculir1:a, Quinqueloculina
etc.
(v) Rotaliinina (Mesozoic-Recent) : Benthic/planktic, larger or smaller, calcareous test,
glassy/vitreous, multilaminar, perforated, unilocular/multilocular, coiled planispirally/
trochospirally, aperture along basal suture; often multiple apertures; canal system and
pillar in larger group; some exhibit lateral chambers. e.g. Nummulites, Globigerina,
Rotalia etc.; largest superorder with 12 super families. Benthic Forms are included in
the superfamily Rotaliacea and Orbitoidacea and planktic forms within the superfamily
Globigerinacea.
D. Ecological studies on foraminifera :

Ecological sudy of foraminiferas constitutes an important aspe~t of oceanographic
investigations. They are preferred to other microfossil groups not only for their abundance but
also for their usefulness in studies related to the climate of the present and th~ past, petroleum .
exploration, biostratigraphy, palaeolatitudinal reconstruction, marine transgression and regres~ion
and also in study of other oceanographic aspects such as chemic.al and physical envir~nment
•
,,
of the sea, type of sediment transfer and in tracing the major oceanic currents. ·
. The principle
. . · behind the role of foraminifera in palaeoecologic and palaeoenviron men t a I
mterpretat1ons mvolves a combination of basic concepts of sedimentolo gy, eco Iogy an d
~eanography supported by abundant data about the life-habits of the numerous 11·v 1·n ·
f h" h" h h · g species
o t 1s group w 1c ave been investigated through wide and intensive researches in th
few decades. e 1ast
In general, foraminifera occurs in all the different types of marine e · d

rarely also in freshwater (such as those belonging to suborder Allogromia~v;:~me:t :;1· ve~
the recent oceans is of the order of about I000-2500000 in I sq met . f e1r _a un a~ce m
•
Mot i · ·& • • er area o marme sediment.
s oramm1aeras are benth1c and hence are controlled by such fact l"k d h
and nature of the ocean floor etc. ors I e ept , temperature
wa~; :~ey s~ow ch~n~e of their distribu.tion. pattern with change of temperature of oceanic
diff ' ey a so exh1b1t a remarkable latitudmal variation as regards the abundance of ·their
i con:;:~:l~oups from equator _to_ pole. The distribution of planktonic foraminifera species is
l Y temperature, turb1d1ty and turbulence of water. Apart from these physical factors,
PALAEONTOLOGY
454
chemical environmental factors controlling the distribution of fora~inifera inclu?e salinity, ~ome
dissolved gases and also dissolved CaC03, Biological factors include associated organisms,
symbiotic association and nature of food supply.
t. Role of temperature on distribution . . 0 0 0

The horizontal temperature from pole to equator 1s ranging fr?m 2. C-0 C (pol.e) to 25 C-
370C (equator). On the other hand, in tropical zone temperature 1s gomg down with depth of
ocean in the order of 25°-28°C at the upper surface to as low as 4°C at great depth.
Overall observation has shown that if other factors remain normal, agglutinated foraminiferas
tend to dominate more in the cold water sea where calcareous forms remain underdeveloped
and are fewer in number. Likewise, the large calcareous forms like rotaliids, miliolids would
be the most dominant forms in the warm water zone. This observation is more or less true as
regards the distribution of recent foraminiferal forms in both horizontal and vertical direction
in the sea.
2. Influence of temperature on shell morphology

.It has been found that temperature variation in sea water may cause variation of size, number
of chambers, .size and spacing of pores, presence and absence of secondary pores and type of
coiling of many planktonic foraminiferas.
Frerichs et. al (1972) have found that planktonic foraminiferas living in more cold and hence
dense water have fewer buoyancy problems and thus have heavier tests with test-porosity, less
than those forms living in warm and shallow water (hence of lower density) where they increase
the buoyancy with development of high test-porosity and several other accessory features like
fine spines, peripheral keels, fine papillae etc. On the other hand, deep water planktons ·to cope
with the problems of high water pressure and low pH secrete extra glassy calcite. Warm and
cold water population of some planktonic species such as Globigerina pachydetna can be
distinguished by a predominance of right-handed dextral forms in warm water and left-handed
~inistral~y coi~ed tests in relatively cold water. The..Jequence of Pleistocene glacial and
mterglac1al penods has been often established from the study of such fo~aminiferal fossils from
Pleistocene rocks in deep sea core.
3. Role of water depth

~at~~-depth controls many ~t~er factors of ~arine environment such as temperature, light
availab1h~, water pressure, sahmty, oxygen availability, carbonate solubility and proportions
of other dissolved gases like co .
2
. Sa~dov~ (1967) has e~tablished ~he dominant benthic and planktic foraminiferal assemblage
m successive depth at different latitudes showing different temperature range (Table 27).
As regards the planktons, the tests of Globigerina pachyde,n h . .. . . · the
Id . a s ow smistral co1 1mg 1n
co temperate polar region (temp 0°C to 18°C) while dextr· . . · al
regions (temp 100c to 24oc). ally coiled forms m the subtropic
455
TABLE-27
FORAMINIFERAL ASSEMBLAGE IN SUCCESSIVE DEPTH AT LATITUDES
Depth(m) Latitude Domonant foraminiferal
No.
groups/species
l. 0-200 20°N-20°S Miliotidae/Nummulitiae
2. (a) 200-500 40°N-40°S Amphisteginids

(b) 200-1500 20°N-20°S
3. (a) 1500-2000 20°N-20°S Trochamnina
(b) 200-1500 60°S-80°N Cyclammina
4. (a) 2500-4500 20°N-20°S . Alabamina
(b) 2000-3500 60°S-80°N Uvigerina
5. > 6000 Tropical & cold Rhabdammina, Bathysiphon,
temperate latitllde Hamosina, Haplophragmoides
The proportion of benthic to planktic foraminifera in the modern sea provides a method for
determining water-depth gradient in ancient rocks in a general way at least. It has been observed
that the present-day planktonic foraminifera increases in number sharply away from the shore
reaching a f!laximum at continental slope and deep sea. On the other hand calcareous benthic
foraminifera predominates in sediments of continental shelf and near shore zones: Arenaceous
foraminiferas are dominantly found in near shore brackish water zone such as in marshes and
lagoons (Fig. 31-12).
Lagoon
Marshes Contiental shelf Contiental slope
Nearshorc Foraminiferal group
e Plank tonic
~
'i: fo"!minifern
·~ r----i:==-------1-----======t-------_J
,!=
....0
~ Calcareous
foraminifcra
Arenaceous
fonuniniferu
FIG. 31-12 : CHANGE IN PROPORTION OF RECENT PLANKTONIC c

. AND ARENACEOUS BENTHIC FORAMINIFERA WITH ALCAREOUS BENTHIC
DEPTHS FROM SHORE ACROSS CONTINENTAL SHELF DOWINNCTOREASING WATER
SLOPE. CONTINENTAL
Pct1
ile(Geo)WP-58
456 PALAEONTOLOGY
4. Light and water depth

Most of the benthic larger foraminiferas such as Miliolina prefer to live in photic zone where
sunlight is available. This zone becomes the main thriving zone of pelagic and benthic algae
making this zone rich in organic matters and 0 2. This makes this zone attractive to benthic
foraminifera. This is due to the presence of greater algal food sources and also often a symbiotic
association of some foraminifera with benthic algae.
5. Salinity
Usually salinity of sea water increases with depth. Majority of the foraminiferas are adapted
to normal salinity of sea water (about 35%) and under such condition they exhibit highest
diversity. Hypersaline condition favours porcellaneous. miliolids and rotalids to grow but deters
other groups. Hence rich assemblage of Miliolina, Textularia, Rotalia may be used as indicators
of high degree of palaeosalinity.
6. Substrate type
Silty and muddy substrate are preferred by the foraminifera as organic debris, especially
bacteria (food of foraminifera) are present profusely in such areas. As mechanical stress is less
on such substrates, foraminiferas growing here are mostly thin-shelled and delicate. On the other
hand, a coarse sand and gravel-rich substrate which contains larger pore-spaces and fewer
amounts of organic nutrients supports fewer foraminiferas for greater mechanical stress in such
condition. Thus foraminiferas from such substrata become fusiform, biconvex, thick-walled
. exhibiting various surface ornamentations. Those foraminifera which prefers hard substrate like
rocky bottom and sea grasses are normally attached either permanently or temporarily by their
flat lower surface of the test and such a test often becomes planoconvex, concavo-convex,
dendritic or irregularly branched.
7. Chemical environment
(a) Dissolved gases : In general dissolved oxygen content in sea water is decreasing down
the depth below the photic ~one due to absence of green plant. Local oxygen deficiency in sea
wate~ may be caused by high organic productivity in some particular zones of sea. Oxygen
deficiency however
. does
. . not greatly affect the smaller organ 1·sms as th ey reqmre
· a_very 11·tt1e
oxygen
. for thetr
. resptratlon.
. However,
. poor oxygen
. content may aftiec t a &,oramm1,era
• ·&· l popu lat1·on
m two ways . reducm~ the size of population and decreasing the size of tests and developing
unomamented
. . . tests
. (either
. calcareous or agglutinated)
. . However, a &,ew genera bel ongtn · g to
bohvm1dae may hve m a oxygen depleted restricted 20 ne smce ·
theu
· appearance.
Presence of H2S produced through so b · I •
. . me actena action (sulfate-reducing bacteria) frotll
decay of organic matters m an oxygen d f · · · 1
f t f th f . .c I e ic1ent environment however becomes inhosp1tab e
or mos o e oramm11era groups. However below the . . reen
plants but presence of other orga · ( ' . . ~hotic · zone with absence of g
. . . msms performing resp1rat1on) CO pro ortion in sea water
1s found gradually mcreasmg. Excess content of th' CO . • 2 p h' h
in normal condition ranges between 7 8 is 2 m depth lowers the pH of water w ic
· - 8·3· Normally at pH value 7 oo d' I · f caC01
from organic skeleton starts and thus above th'1s value. th - . · . . ' isso utton o. · ·rera-
begins to decrease in sea water. e proportion of calcareous foramint
457
(b) CaC03 dissolution-Calcium Carbonate Compensation Depth (CCCD) : The amount of
dissolved CaC03 in sea water primarily depends upon the water temperature which is greater
in warm water than in cold water. Hence solubility of CaC01 is indirectly proportional to water
temperature. So there is more soluble CaC0 1 in sea water in warm tropical zone at shallower
depth. This causes increase of proportion or" calcareous organisms including foraminifera and
also proportion of limestone in this zone. 'Fominifera tests become larger and robust-sized
because here there is more precipitation of excess CaC0 3 from water (CaC03 supply rate is
higher than the power of its dissolution by water).
However, in cold water at greater depth in lower latitude or cold ~ater at shallow depth in
higher latitude, the amount of soluble CaC03 is much less and hence the rate of soluting of
the CaC03 is higher than rate of supply of CaC03. CaC03 solubility also increases with water
pressure. The lower content of CaC01 in deep sea with cold water produces a condition
unfavourable for development of rich population of calcareous foraminifera. Such zones are
dominated by planktic foraminifera or forms with agglutinated and arenaceous tests.
Furthermore with progressive increase of depth, the ratio of dissol'ved COif02 in water
increases because of decrease of photosynthetic plants although animals continue to respire.
This leads to decrease of pH of water with depth from normal 8.00 to as low as 7.00. The
level at which rate of CaC03 dissolution becomes equal to rate of its supply is called Calcium
Carbonate Compensation Depth (CCCD) or simply Carbonate Compensation Depth (CCD).
This level is extremely variable from latitude to latitude with the variation of water temperature
as in the Pacific Sea it is lying between 4000-5000m in lower latitude but only at 400-500m at
higher latitude. As the limit of CCCD is difficult to locate, the 'Concept of Lysocline' i.e. · the
level of maximum chang~ in the rate of solution of CaC03 is often preferred. As beyond. this
limit CaCO from the skeletons of all .calcareous organisms including foraminifera begins to
dissolve, su~h a condition becomes quite unsuitable for such organisms . .
8. Food habit .
Like many other macro-organisms foraminifera play an important role in marine ecosystem.
They are micro-omnivorous organism feeding on small bacteria,. algal protista as well as on
microscopic invertebrate larvae. Some of them scavenge but mostly they are predators. Some
foraminifera even show a symbiotic association with algae much like that shown by hermatyp'ic
corals in photic zone. The algae provides n~trients from photosynthe~is supplying Caco to
3
the animals.
High diversity and abundance of foraminifera population may suggest easy and wide-range
availability of food. Local and seasonal function of food supply may cause variation of its
diversity. Planktonic foraminiferas tend to thrive in regions of oceanopelagic zone where
phytoplanktons are dominant which become the chief source of food of the animals. .
Benthic foraminifera. have great chance of being ingested by other marine predators such as
worms, g~stropods, ech.moderms .and many fishes, which are mostly deposit feeders browsi.ng
o~ the sediments a~d m1cr~-organ~sms on the sea floor. The effect of such organisms is however
difficult to ascertam. But m fossil recqrd original abundance of a particular assemblage may
greatly be reduced by such selective destruction of the foraminiferal tests by the deposit feeders.
. .
PAUEONTOLocy
9. Salient features of foraminiferal ecology
(a) Marshy fauna is dominated by agglutinated taxa such as, Peneraplis, Textularia etc
(b) Lagoonal facies of rocks is dominated by Ammonia. j:;lphidium and Quinquelocut
. h I .
wh1c are ca careous 1mperforated foraminifera.
tna
(c) Brackish water faunas are consisting of cosmopolitan species. 'outer bay' and 'inner bay'
faunas may be demarcated by the i;atio of calcareous/agglutinated forms such as Ammonia/
Ammobaculites. Such a delineation 1s necessary in petroleum exploration in order to find out
the rate of sedimentation and the --ctegree of reducing condition in which organic matters are
converting into petroleum.
(d) Coastal shelf is dominated by benthic species. Planktonic forms begin to outnumber them
in outer shelf zone and about completely replace them in upper bathyal zone. Their ratio of
about 20: 1 is indicative of a depth not less than 200m.
(e) Forms with simple agglutinated species indicate bay and lagoonal environment but
agglutinated forms with-tabyrinthic wall structure and siphonate chamber may be found in outer
shelf and also in bathyal environment.
(f) Diverse miliolids dominate in inner shelf but biloculine forms dominate in bathyal zone.
(g) Calcareous tests with smooth perforated wall are the dominating forms of inner shelf
while the simple ornamented but large and thick forms are abundant in deeper zone.
(h) In abyssal and hadal zones CCCD level prevents accumulation of calcareous foraminifera
and thus beyond this level most forms bear agglutinated tests.
(i) Majority of the planktonic forms living at surface water within the photic zones are
spinose, e.g. Globigerina; while the forms living in lower water zone are usually thick and
keeled. e.g. Globotruncana.
(j) Change of coiling direction of some planktonic forms as shown by some Globigerina
_species may be a good indicator of change of temperature of sea water. Abundance of dextral
forms indicate normal warm water and sinistral forms grows in cold water temperature.
10. Foraminifera test as contributor to marine sediments

Planktonic foraminiferal tests are important contributors of deep sea sediments often
associated with such other organisms like coccolith. They account for more than 80% of the
modem carbonate sediments in deep sea as organic oozes. Three factors are controlling the
depositions of Globigerina ooze (Globigerina forming more than 30% of the sediment) which
are climate, depth of lysocline and supply of terrigeneous sediments. Berger ( 1971) has estimated
that 6-10% of the living populations of plankton leave empty tests everyday, mostly as a res!llt
of reproductive process. These tests settle to bottom to accumulate as ooze. These tests are
less susceptible to dissolution than that those of coccoliths for their skeletal outer layer, except.
when they approach lysocline level, usually lying between 3000-5000m depth. Fluctuation of
depth of lysocline thus can cause cyclic deposition and dissolution of these test. Even the
conditions remain otherwise favourable, oozes cannot properly accumulate where there is greater
influence of tcrrigenous elastics and hence such oozes are rarely found on the shallower
/f{{RODUCf!ON TO MICROPALAEONTOLOGY 459
continental shelf zone. At present, such oozes are producing at 50°N and 50 S latitude at depth
between 200-500m especially along mid-oceanic rift zones.
E. Geological history :
There is no distinct evidence of occurrence of foraminiforn fossils from Precambrian rocks.
The first report came from Early Cambrian rocks represented by ammodiscaceans resembling
Bathysiphon and Tolypammin.a. It is generally held that the uniloculur fonn was the most
primitive. It is also held that the lagynacids (the foraminifera with organic skeleton), a lor1g-
ranging form probably continuing from Precambrian is the ancestors of agglutinated
ammodiscaceans. The Cambrian occurrence is however disputed by many workers who held
the view that true skeletal foraminifera did not come before Ordovician. Foraminifera with hard
test of the group fusulinids appeared from ammodiscaceans ancestors and began to flourish from
Silurian onwards culminating in the complex test of Fusulina in Late Carboniferous-Permian
and the group died out at the end of Permian. Lituolacea and Milioliids appeared in the Early
Carboniferous possibly from ammodiscaceans (agglutinated test).
Rotaliinids however made their first appearance with the beginning of Triassic possibly from
Fusulinid ancestry. They began to diverse in the Jurassic-Cretaceous Period when appeared most
of the families of th~s group including the first planktonic foraminifera in Jurassic.
Milioliids, textulariids and rotaliids flourished heavily by Cretaceous-Tertiary helped by the
newly opened Atlantic Ocean. The planktoic group Globotrucanids ~f Mesozoic became exinct
at the end of Cretaceous and gave way to other groups such as GlobOrotalids and Globogerinids
which made their appearance in Paleocene-Eocene times. Along with this, appeared Nummulitids
in the Old World and Orbitoids in the New World seas and soon they became worldwide
abundant. Most of the Nummulitids died out after Oligocene and Orbitoids after Miocene. Since
then. larger foraminifera stocks show progressively decrease in diversity and number.
F. Use and application of foraminifera :

{a) Biostratigraphic use :
Being small-sized but abundant in number, foraminifera fossils (both benthic and pl·lnkt ·
& • • • • • • omc
,orms) are found widely d1stnbuted especially m rocks of Tertiary age when they show , ·
d" · . . m,tx1mum
1vers1ty
. and . abundance. For this reason,
. marme
. Tertiary rocks of various, part,
. s o f th e wor Id are
bostrat1graph1cally zonated on the basis of various zone fossils of foruminife ~ M·-
h · . 1•1· any o f t he species
. .
s ow evolutionary changes of some morphologic character with time thu. d'I . . .
f ·1 h 1 · · ·· · s rea I Y tracable w1thm
oss1 s e pmg m recognition of several lmeage groups. Planktonic f . · 'f . . . .
large n~mber of index fossils since Cretaceous are successfull ~r~n.11m er~s which ~1eld a
correlation of Mesozoic-Lower Tertiary rocks (Bolli 1966) Ben ~ use_ I~ .t~e mtercontmental
somewhat restricted distribution may be used f · . thic forum1mtera, although show
• · or 1oca 1and m ~ · , . . .
rocks. These studies coupled with radiometric d· t' h· . som~ cases regional correlation ot
high resolution biostratigraphic zonation and a /n~ ave helpe~ m establishing the concept of
faunal zones of nearby or a d "tstant .corre ation. Correlat10n has been done by comparing
. . section more a . •
foramm1feras are involved Late ti . 'ccurnte 1y especially when pf ankroruc
· r, a re med radio t · . • .
proposed to which planktonic foraminifer· me nc tune scale tor the Cenozoic has been
a zones have been matched. Marine Tertiary rocks based
-
460 PALAEONTOLocy
on benthic f~ra~inif~r~ have b~en zona~ed in Ir~dia and abro~d. ~ome index fossils of benthic
and planktomc toramm1fera which help 111 zonat1on and drawmg time boundaries such as tho
between Cretaceous-Tertiary (Maastrichtian-Danian), Paleocene-Lower Eocene, Lower Eoce/~
Middle Eocene, Middle Eocene-Upper Eocene etc may be listed in Table-28. e
TABLE-28
ZONATION OF MARINE TERTIARY ROCKS OF INDIA
Geological European
-
Leading index fossils
period time scale
,
Miocene (Lower) Burdigalian Taberina malabarica, Austrotrilina howchini
Miogypsina globulina,
Lepidocyclina (Nepholepidina) sumatrensis
Aquitanian lepidocyclina (Nepholepidina) tani, :I:
Miogypsina dehaarti ~
;:>
::(
Oligocene Up Chattian M iogypsina complanata
Mid Rupelian N. fichteli, lepidocyclina (Eulepidina) dilalata
Lr Lattorfian N. fichteli
Eocene Up Priabonian Pellatispira, Heterostegina; N. fabianii (Assam),
Hantkenina alabamensis (Assam)
Mid Nummulites atacicus :c
u
f-4
N. acutus; N. obtusus, Assilina exponrns/ ::>
Lutetian ~
0
Discocyclina sowerby; D. dispansa/
~
.Alveolina elliptica ~
<
(/1
Lr Ypressian N. atacicus, Assilina granulosa/ (/1
Globorotalia rex (Kutch and Assam) <-
Palaeocene Up Montian Globorotalia whitei, Miscellanea miscella, 0
z~
lockhartia conditi., N. thalicus, Discocyclina < t.tJ
ranikoti (Assam and Pondicherry) ~>
Lr Danian <~
Globorotalia uncinata, Globorotalia ~u
trinidadensi.,· (Pondicherry) <
Uppermost Maastrichtian Abathompha/us mayaroensis
Cretaceous Glohot ru11ca11a gansseri (Pondicherry and
Trichinopoly)
Globot run.cana t ric:a rina ta
-
(b) Environmental use :
Applications of foraminifera in environmentnl interpretation are bused on comparison of data
colle-et.ed from the study of recent fomminifern. For example, dramatic change of depth, salinity
.. -
/NTRODVCT!ON TO MICROPALAEONTOLOGY 461
d climate may be traced during the Pleistocene time from the different studies on foraminifera
; : Pleistocene beach deposits. (Scott & Medioli l 97~) Fluctuation of palaeotemperature during
01
;leistocene glaciation is also traced from the study of relative dominance of dextral and sinistral
tobigerinid tests from Pleistocene deposits. (Ramsay, 1977; Funnell & Rhedel 1971)
ialaeosalinity of earliest peri,ods may be traced from the ratio of textulariina, milioliina and
rotaliina as well from the diversity of foraminifera populations. The value of foraminifera as
indicator of the depth of deposition has been outlined by Funnel ( 1967) by comparing genera
and species. Another useful guide is 'the ratio of planktonic to benthic individuals in a sample,
the former increasing gradually as the water is more and more away from shore with increase
of depth.
Living planktonic tests may be used as indicators of current circulation within a mass of
water. Creteceos current pattern can even be reconstructed from the distribution of their fossils.
Calculation of the rate of sedimentation involving knowledge of the length of foraminifera life
cycle have been also suggested by Uchio ( 1960).
However, little use has been made to correlate the morphologic character of foraminifera
with the environment in which they live although there are some preliminary studies relating
to depth (Bandy 1964), substrate (Brasier, 1975) and general environmental factor, (Schafer
Pelletier, 1977).
31.6.2 Radiolaria (Fig. 31-13)

Radiolarias are exclusively marine planktonic protozoans, each characterised by many
radiating filamentous pseudopodia (hence the name), a central capsule and a silicious skeleton.
The animal's body is spherical, subspherical or bell-shaped, composed of protoplasm which
has two distinct parts : a relatively thick outer extracapsular layer and a central capsule (the
feature that has differentiated it from all other protozoans). The extracapsular cytoplasm is
concerned with floatation, nutrition and secretion of silicious skeleton. It is again divisible into
three layers; an inner assimilative layer, next to central capsule, an intermediate layer called
calymma (helping in floatation and secretion of skeleton) and an outermost permeable
enveloping membrane called sarcodictyum. The central capsule has porous membrane of
chitinoid and mucoid materials. It is a permanent structure with a nucleus surround d b
cytoplasm with vacuoles. It se~ves in reproduction and food storage. Pseudopodia are e;ittiny
from central capsule and passing through all the three outer layers. g
The skeletons of radiolaria, radially or bilaterally symmetrical show d. . .
forms. They are generally composed of opaline silica but in one grou' it is m~ great 1v_ers1t~ m
sulphate. The skeleton is mainly external. In some genera the bp ade up of strontium
the outer skeleton; in some forms there are more than o' lmem rfanous central capsule bears
· ne ayer o skelet l T · ·
the other. The skeleton typically consists of an intricate lattice-work ·. . a s1 ica, on.e m.s1de
stellate or conical (bell) shapes bearing radial spines or lackin th having glo~ular~ d1sco1d~I,
classified into four suborders namely Actipylea, Peri lea to em. The rad10l~na group .•s
classification is based on the number of .. PY ' nopylea and Tnpylea. This
pores m central capsule th .· . f h k I
arrangement of spines and some other characters. ' e compos1t1on o t e s e ~ton,
The fossils of radiolaria are known f o p · b ·
number · th . r m recam nan rocks and they are still living in large
m e present sea. The dead skeleton of this animal have constituted many silicious
PALAEONTOLOGY
462
Spherical Test
-----Outer Silicious Skeleton

- - - - Pseudopodia
_ _ _ Outer Cytoplasm
...--.....--
. - - - Middle Skeleton
li:.1ii~...--- Inner Cytoplasm

U,1;,1-,ii!Ht---nner Sphere Skeleton
~ - - Inner Or Central Capsule
A Cross Section Through a Spherical Test
HG. 31-13 : RADIOLORIA
rocks of Precambrian times. Radiolarian cherts are commonly found associated with geosynclinal
sediments in many old and young orogenic belts.
31.6.3 Ostracods (Fig. 3 l-14a)

Ostracods are microscopic, mostly marine invertebrates belonging to class Crustacea and
phylum Arthropoda. They bear a bivalved calcareous carapace (enclosing the soft part), hinged
along the dorsal margin. They distinctly differ from bivalved mollusca in having soft part wi~h
several paired appendages like all other arthropods. They are mostly marine and ranging ,n
age from Ordovician to Recent. The size of the animals is ranging between a fraction of a
millimeter to about two millimeter. They show variable shapes of the carap: ~e, but mostly are
ovate or reniform.
The bivalved carapace is composed of two almost equal and identical valves dorsally hinged,
called right valve and left valve . Valves consist of calcified chitin divisible into outer and inner
-""'-'
lamella. Soft parts including the appendages are enclosed by the carapace. The growth pattern
like other arthropods is ecdysis for which the valves lack any growth marking.
Head and thorax are fused forming a single segment called cephalothorax. Among the five
pairs of cephalic appendages the first two are atennae, second pair mandibles , third' pair
maxillae and the rest are legs. Appendages perform several functions including locomotion,
food-capturing and mastication of food materials.
Two kinds of canal penetrate the carapace. A radial pore canal, subparaJlel to the outer
surface of carapace, lies in between outer an inner lamella and free margin . Normal pore canals
are simple, sieve like, usually scattered over the carapace, perpendicular to the surface. The
connecting adductor muscle in between the two valves, often marked in fossil as scar, are located
at the central part of carapace-interior, externally marked by sulcus . The nature of these scar-
markings are different in different groups.
Hinge of an ostracod possesses two types of tooth elements terminal and median that
function in articulating the two valves. Most primitive ostracods have weakly developed carapace
without teeth (adont) where the median element is consisting of a Jong ridge on hinge of right
valve and a corresponding groove on the hinge of left valve. Merodont hinge exhibits elongated
and strongly crenulated terminal elements on right.value which fit into terminal sockets of left
valve Amphidont hinge has short terminal elements that consist of well developed teeth and
sockets in both the valves together with median elements.
Outer surface of carpace shows variable types of ornamentation. An eye tubercle occurs in
many valves as a transparent lens of calcite, directly outside the internal eye, helping in
penetration of light through carapace. Carapace surface shows reticulate, spinose, granulose
types of ornamentation.
Several carapaces are left by an animal during its successive stages of growth and those
empty carapaces are called instars which are often preserved along with the adult form. The
two valves of an ostracod carapace may ·be identical, equal-sized but in some cases one valve
may be slightly larger ·either along dorsal or along ventral margin or all along the margin of
the valve. The wider side o(the valve is generally indicating anterior and narrower side posterior
direction. The characters which are given special attention for identification and classification
are (a) nature of valves (its shape and convexity) (b) structure of hinge (c) line of attachment
of two valves (d) surface ornamentation and (e) nature of such structures as brood pouch, hinge
pattern, adductor scars, eye-lid lens etc. A few generic forms are Cytheridea, Cypris, Tetradella,
Digygopleura etc.
Most of the ostracods live in shallow sea but some are known to occur in river and lake.
Ostracods with thick carapaces are considered animals living in a high-energy environment while
ostracods with thin and fragile car~paces are characteristic of quiet water. Many of them are
simple mobile benthic forms crawlmg on the surface by their legs while other are swimmers;
many prefer hard bottom; some _are sand dwellers and some are epibiont. This is a long ranging
group (Ord.-Rec .) whose fo~stls are found within variable types of marine sediments and ,•
occasionally they constitute bulk of the sediment.
464 PALAEONTOLOGY
Digestive Tract -----Hinge Ant.cnnule
. - - - Ligament Antmna
Calcified Outer
Adductor Muscle
Eye
· - - - Lamclla Branchial Sctae
Adductor
Muscle
Appendages
Ventral
Dimension In 25J·. .iw-- Calcified
Side view Shell Inner Lamella
Radial Pore Canal

-----Free Margin
Morphologic Features In C.S
Main Appendages
Overlapping Valves -----Terminal Teeth
Hinge Arca
In Dorsal View
Median
Dental Bar
Adont Hinge Merodont Hinge Amphidont Hinge
,._
Type Of Hinge & Other Features
Eye Tubercle Longitudinal Ribs
Pitted Ornamentation
Ribbed
Surface Sculptures
(a) BROAD MORPHOLOGICAL fEATURES OF 0STRAC00S
Multiple Thoracic Segments
•
Rivalved Shell
Dorsally Hinged
Anterior
Caudal Furcae
'----Thoracic Appendages
Inner View
(b) CfZJCUS (cstheria)
Eun/cites
Upper
Side Upper Lower
lldraites Side Side
/cl SCOI. F.COOONT FOSSIi S Oenonites
Ff(i . 31 - 1-1 ()\11(\('()f)"i . l ·"il"lll:l<ll>S :\~I) S(.'1)1.i-:('<>l>O\'rs
31,6.4 Estheriid (Fig. 31- I 4b)

Estheriids are a microscopic arthropods belonging to subclass Branchiopoda and order
Conchostraca. It has a bivalved carapace like ostracods which encloses the soft part. Unlike
ostracod, the animal's body is well segmented consisting of 20-27 somites with numerous
appendages. The last segment is bifurcated producing a caudal /urea. In the absence of soft
part, it is difficult to distinguish their carapaces from those of ostracods. However, as the name
'Estheria', allotted to the genus is pre-occupied, it is renamed at present as Cyzicus. Mostly
fresh water, the animals are ranging from Silurian to Recent. In India, estherid fossil E.
mangliensis has been reported from Deoli bed of Panchet Formation in Ranigunj coal-field.
The morphology of this very primitive microscopic arthropod remains almost unchanged since
its appearance upto the present day and hence it is also taken as an example of living fossils.
31.7 MICROSCOPIC PARTS OF ANIMALS-SCOLECODONTS (Fig. 31-14c)

Scolecodonts are a group of chitinous, horny or silicious microfossils possibly representing
jaws and denticles of worms polycliaetes (Phylum Annelida). In living polychaetes there is a
large stabbing tooth, or a circle of small denticles, and more commonly a pair of large grasping
and tearing jaws. Living Eunice carries its numerous denticles and jaws in a special pouch below
the digestive tract. Comparing the size, composition and morphology of denticles of living
polychaetes with those of fossil specimens, the affinity of scolecodonts to worms-teeth is more
or less confirmed. Scolecodonts show a considerable variation of their size, shape, structure
and number both in living and fossil gen~ra. But it is difficult in many cases to isolate
scolecodonts that actually belong to a single individual. Scolecodonts are found in all the
geological horizons from Ordovician but they are more common within Palaeozoic shales and
impure limestones. They appear in rocks as black microscopic objects, commonly in large
numbers. Sometimes they superficially resemble similar tooth-like fossils, the conodonts, but
the latter show much wider variation of size and shape, different composition (calc-phosphate),
structure and colour (light brown). A comparison between these two is shown in Table-29.
31.8 FOSSILS OF UNCERTAIN AFFINITY-CONODONTS (Fig. 31-15)

Conodonts are a group of microfossils of uncertain biologic affinity. Most of the
palaeontologists believe that they represent some sort of denticles (dental structures) associated
with some ,animals (may be a fish or an annelid). They range in size from I to 2 mm
and show variable shapes. The colour of conodont is typically brown with sometimes a glossy
lustre.
Most commonly, conodonts are tooth-shaped structure, single or multiple pointed, almost
~lw.ays found attached. with a basal bony pla~e ~f ~ame compos~tion. X-ray spectrography has
md1~ated that compos1ti~n of the conod?n~ 1s s1m1lar to the mmerals of apatite group (more
precisely carbonate-apatite), somewhat similar to skeletal composition of vertebrates.
Geometrically, conodonts are grouped into three types of elements; (a) coniform, which is
cone-shaped consisting of an elliptical base and a pointed cusps very often curved
. rami"o
(b) tt• rm, w h'1c h ex h'b' · fl k d ·h 'd · '
I its a mam cusp an e on e1t er s1 e or one side by processes bearing
'
smaller denticles that are similar or dissimilar-sized (c) pectiniform, which has smaller denticles
on a platform bearing a number of radially directed processes. There are variations within eac.h
( gro~p. such as : 2 types within coniforms, 7 types within ramiforms and 15 types within
l)ectin1forms.
PALAEONTOLOGY
466
Distacodid
Non-Fibrous
Fibrous Lameller
(a) DIVISIONS BASED ON INTERNAL STRUCTURES
Apcx(upper)
Main Cusp
,......~--Internal
Central Cavity
Basal Plate Anterior Posterior

Ramiform Pectinifonn
Ba.sc (Lower)
Coniform
(b) DIVISIONS BASED ON FORMS
- - - White Matter
Lamell~
Contact Zone
-....:i~'I----Bml Plate
{c) INTERNAL STRUCTURE OF A CONODONT
Conodcnt Elements Inside

The Sac Like Body
~
Conodont Elements
Conodont Predator Near Head
(d) CONOOONT ANIMAi. Cl YOAC.NATHUS
l·lli .. ~l - 15 : CONODONTS
'I
f .....
_:f,
'
I
467
Internally, these elements mostly compo · d f . . b ·
. . b . se o some car onatc-uput,tc.~ having u basal ,,art of
same compos1t1on ut with more organic · tt Th ' . b· . · ·
. ma ers. 1s asal plate may rcpres ·nt' t·he pl ace of
attachment of the elem~nt to the host body. lnternally, conodonts may sh ow u/ibrou.r, (compo cd
o~ bundles _of fibrous minerals), lamellar (parallel lamellae of mineral s) or a di.vtacodld (non fibrous
with cone-m-cone s~ructure) structure. In lamellar type of conodonts, occur some bubble like light-
coloured, non-laminated _stru~tures called white matters which arc opaque and dark e r than
lamellae. In .s~me groups mtenor of the element contains a cavity in the shape o f the element. In
others (pect1mds) there may be a small internal pit centrally located .
Recently, from some Carboniferous sediments a few fossils have been reported which exhibit.
impressions of animals like worms or arthropods or fishes bearing sac-like bodies of 5cm- 8cm
long with possible muscle or organ impressions within which conodonl elements are found in
groups. These elements are interpreted as a part of digestive system of a filter feeding animal
and the sac-like body itself represents a conodont. Or, the elements may represent the stomach-
content of a condont predator (may be a fish/annelid or arthropod). In 1983 Briggs et al. has
reported a fossil from Lower Carboniferous sandstones near Edinburg. The fossil is an impression
of a slender segmented warm-like body about 4 cm long having a terminal tail-fin and internally
possessing conodont-elements in the other terminal part (opposite to tail) . This may indicate that
conodonts may represent some denticles of an eel-like animal. Detailed study has also revealed
that segments of these animals are chevron-shaped like those of chordates rather than worms. So
conodont elements may represent denticles of some very primitive jawless fish-like vertebrates.
With this present knowledge, a search for a conodont animal or its actual affinity could lead
to some organisms having some carbonate-apatite elements in their skeleton . It should be
remembered that carbonate-apatite mineral is not a common element of skeletal parts of organisms
but some brachiopods, worms, arthropods and even some vertebrates have such mineral in their
skeletons. Scolecodonts are similar elements, generally assigned to some annelid denticles, that
are distinctly different from conodonts.As no other comparison is possible except this mineral,
relationship of conodont elements with any other animal is obviously not confirmed. At present,
they may be recognised as a separate phylum (Conodonta) with no other relatives. However, with
the recent discovery of more specimens similar t~ that described from Edinburg in 1983, idea of
accepting conodont-elements as tooth-like structures associated with some primitive jawless
vertebrate is gaining ground (Aldridge et al. 1993). These fossils indicate that a conodont bearig
animal had eyes, otic cap·s ule and traces of branchial bars and possibly a notochord. Conodont
elements are found beneath the head region of the animal probably functioning as a complex
feeding busket helping in feeding, in capturing preys and in cutting them into pieces. At present,
this animal is named Clydagnathus. Many worker, have considered it as an extinct but very
primitive Agnatha ranging from Cambrian to Triassic.
Stratigraphically, however conodont fossils are common within marine Ordovician-Permian
rocks associated wih many other invertebrate fossils of contemporeneous times, but they are
,
more abundant within black shales (Ordo-Silurian age) where they occur along with graptolites.
Many conodonts are recognised as valuable index fossils and are used as age-markers of rocks
I -.
where other significant megafossils are absent. Some common genera identified are, Distacodus,
Acodus, Prioniodus Lonchodina, Lonchodus etc.
LL g a P .b
• . a; I Aft. . ·,-· ,, .,
.. -- • ..,. "' - ... J ...__.
I ,W .__ . ·- .. _, .._
p I\ I ,A EONTOLOG Y
468
....--
- · Distal Shield
Plates -~~~v--,,-,...... Proximal Shield
~~r----,,. (Two Layered)
Pcrforations
Funow Distal
~_,,,,,..~:::t
Calciadi~ll11111 (Tertiary) Shield
Grnuwdinium (Jur-Rec) (b) COCCOLITHS

. ta) l)INOFLAGELLA 11 :S
Terminal Nodule--,,'11
-!,----- Raphc
Granules
Two
. . - - - - Unequal
Valves
Central
=·---e-~
Pore--~~....
Connecting
Band
Pennate Diatom
Diatoms (dorsal view) Centric Diatoms

Centric Diatom
Cross Sectional View Of Valves
Pcnnate
Valve View Of Diatoms
ll'l OIATOMS
Top View
OfOogoniwn
-----Leaf
Nut.uk (oo~oniu1111 I .onttiludinal Srclion Of Nucule

(, idl' , 1.:w ) :\ u,·uk /<l. 11," 11111u111 of Chura Chara Stem
Oogonium And With
Anthcridium
1111c1t,\l( I\ \ • AND
FIG.31-16 :
, ~---
SOME PLANT MICROFOSSILS (CHARA, DIATOMS, D1NOFLAGELLA1E
COCCOLITH)
..
.. Scanned by CamScanner
!NTROD CT/01\ TO MICROPAL.AEONTOLOGY 469
TABLE-29
DISTINCTION BETWEEN CONODONTS AND SCOLECODONTS
Conodont Scolecodonts
Composition Mostly Ca-phosphatic Mostly chitinous and
soluble in inorganic acids. soluble in organic acid.
Colour Brown with glossy lustre. Jet black with dull lustre.
Host rock Common in black shales. Found in all types of
shallow marine sediments.
Structural features Almost always associated Basal bony ridge
with a basal plate, usually absent.
Geological Range Cambrian (?) Ordovician Ordovician to Recent.
to Permian.
31.9 MICROSCOPIC PLANTS

31.9.1 Dinoflagellates (Fig. 3 l-16a)
Dinoflagellates are often treated as some borderline creatures with combine features of the
most primitive plants and primitive protozoans. Considering the possession of chlorophyll-
pigments as one of the decisive features for distinction of plants from animals, most of the
authors prefer to consider them as a group of algae belonging to the group pyrrhophyta or
dinophyceae. They seem to have originated in Palaeozoic (may be even older) and are still
existing today as a group of phytoplankons living in upper water of the ocean in the photic
zone, where sunlight is available for their photosynthetic process. The living forms show two
stages of life, a normal planktic stage bearing two flagellae and an uncommon cyst-stage,
probably formed under adverse condition. Skeletons of such cysts are made up of organic or·
inorganic hard parts (calcareous or silicious) that make them sufficiently hard and most of the
dinoflagellate fossils represent the cyst-stage of the animal, usually consisting of two or more
articulated plates with furrows at their junctions. There are three basic types of cyst but each
bears an exit pore called archeopyle . It is via this hole the organism escapes upon the return
of favourable condition. Although, many dinoflagellates are planktic, some are found in
symbiotic association with some invertebrate groups encrusting on the outer side of their
skeletons. The most conspicuous example of this phenomenon is the symbiotic association of
living reef-building seleractinian corals and dinoflagellate Zooxanthelle. Association with these
algae has forced the reef-coral to grow within the limit of photic zone of the sea.
Dinoflagellates are widely used as biostratigraphic tools in Mesozoic and Cenozoic rocks
but they are also useful in past-environmental analysis; Cyst-type, for example, may reflect a
near shore or offshore condition. In India, dinoflagellates ure reported from marine Mesozoic
beds of Kutch, Trichinopoly and Assam.
31.9.2 Coccoliths (Calcareous nannoplanktons) (Fig. 3 I- I6b)

c 1·ths are the minute calcareous skeletal elements secreted by some microscopic
occo I ·d· ·d pelagi·c flagellate algae. l'hey occur on tI1e outer surface
. of. this
. smg
. 1e-ce 11ed
Ch rysomonam ma
470
PALAEONTOLOGY
plant, forming a sort of armour around the cell. The function of these calcareous element .
not well understood. In a single cell, the coccolith elements may show one or variable type:~~
morphology such as : elliptical imperforated plates with thickened margins called discolith·
perforated single disc called tremalith and others. All these plates are very minute-sized visibl~
only under electron microscope. After the death of the animal, cocolith elements become
disaggregated from their parent body and are accumulated with the sea-sediments. In rare cases ,
however, encystment prior to death can preserve the entire armour in its original intact condition
and such a fossil is called coccosphere. The classification of fossil coccoliths is based on their
nature of perforations on the plates.
Fossils coccoliths are known from sediments as old as Upper Cambrian but they became
common in.Jurassic, Cretaceous and Tertiary. A cubic centimeter of a rock specimen of Eocene
marl of southern Bavaria yields about 5000 forminifera but about 800 million coccoliths. Such
diversification has made them one of the-principal micropalaeontological zone fossils of
Mesozoic and Te~_iary. A recent coccolith is Cycoccolithia. The fossil f9rm Coccolithus is found
in Oligocene of France. -In India, coccoliths are known from Cretaceous of Assam and Tertiary
of beds of Kutch. Silicojlagellates are marine planktonic organisms related to coccolithophorids
but with silicious instead of calcareous skeletons consisting of a simple system of rings, arcs ;
and spines. Their fossils are known from Cretaceoll$ to Tertiary, commonly found within silicious
sediments associated with diatoms. Recently all these planktic microorganisms having affinity
to both plants and animals are grouped as nannoplanktons.
31.9.3 Acritarchs
Acritarcha is a newly erected group of phytoplanktons including some spore-like or cyst-
like biomorphs of unknown origin. The recognition is based on some very minutes fossils similar
to cysts of dinoflagellates whose living mobile forms are not known. They form an assemblage
of microfossils of uncertain affinity but most probably represent some sort of algae-like
dinoflagellates. The record of fossil acritarchs is much older than dinoflagellates as their fossils
are reported even from Archaean crystalline and schistose rocks. In India, Dharwar rocks are
found to contain such fossils. This group is found to dominate in Palaeozoic but decline after
Mesozoic. Some oldest Indian form are laeosphaeriada, Archaeofavosina etc.
31.9.4 Diatoms (Fig. 31-16c)

Diatoms are microscopic yellow green algae (Chrysophyta). In addition to chlorophyll, the
cell possesses other pigments like xanthophyll and carotine; they together form a mixed colour
(yellowish green) called diatomin. Some of them are found in fresh water but mostly they~
phytoplanktons of sea. Often these unicellular organisms are joined with one another fornung
a colony.
8
· The most characteristic feature of the diatom is its possession of a cell waJJ composed of
bivalved skeleton (often called frustules) made up of opaline silica. The two valves of the
skeleton are unequal, one larger outer valve (epitlieca or epivalve) partly enclosing the smalle;
inner valve (hypotheca or hypovalve) like a petridish or a box and its lid. The edge of e~cg
valve is inwardly curved that causes tight attachment. Their junction zone forms a connecll\
band often called girdle. In some cases, a v-shaped notch is found along the midway of ea~d
valve called raphe. There are three nodular growths, two at polar ends (polar nodules) a
-~· -- _ ..
471
one at central part (central nodule). Due to unequal accumulati'on f ··1· h
. . . o st tea on t e cell wall
the latter exh1b1ts different types
. of surface sculpture like stri·ae , punc t ae, granu 1es, etc.
Structurally,
. there
. are two. baste groups. of diatom skeleton . Centri·c t ypes are ra d'1a 11 y
symmetncal and m valve-view appear c1rcular, polygonal or triangular and their cell wall is
without raphe. Pennate grou~ exhibits ~ bilateral symmetry of skeleton and it appears lenticular,
needle shaped, or s-shaped m valve view. The valves are asymmetric in girdle view.
Among the phytoplanktons, diatoms are so abundant that they often form the main constituent
of food chain of many marine animals including fish and whale. In fact, each diatom cell is a
house of reserve food especially of oil. Thick accumulation of diatom skeletons on the sea floor
produces a silicious sediment called diatomaceous earth or fuller's earth. This hard sedimentary
rock is used commercially in various purposes as abrasives, lining within blast furnace, making
hollow bricks, backelites role etc. Most fossil diatoms are centric· types. The pelagic habit of
the animal enables them to disperse widely and some diatoms become good index fossils helping
in long-distance correlation of rocks. Fossil diatoms are known from rocks ranging in age from
Jurassic to Recent but they are most common in Tertiary deposits particularly within fine grained
silicious sediments. In India, numerous diatom fossils are reported form Cenozoic rocks of
Andaman (Gupta, 1974). Some common diatom fossil are Amphipleura, Calonies, Campylonies,
Cymbella, Pleurosigma etc.
31.9.S Chara (Fig. 31-16d)

Chara is a freshwater green algae (Charophyta) characterised by a plant with a central axis
often called 'stem' divisible into nodes and internodes. From each node develop a few needle
like branches often called 'leaves'. There are rhizoid-like structures from the basal end of the
axis. The internode portion of the axis contains a central elongated cell, surrounded by narrow
elongated cell layers, often called cortex. The plants are lzomothalic producing both sperm-
producing organ globule or antheridium and ovule-producing body oogonium or nucu/e,
attached to the node region, the former underlying the latter. The oogonium develops a hard
cel\-wal\ that protects the ovule and matured zygote and germination of latter leads to a new
plant. From palaeontological point of view Chara oogonia are important as they are commonly
preserved as microfossils.
A mature nucule or oogonium is a small, often petiolate, oval-shaped body covered by five
elongated tube cells. They lie side by side spirally covering the inner organ. The terminal end
of each tube cell is free and together they constitute the terminal crown or corona. From India,
several species of Chara such as C. coeleta, C. subglobossa, C. malcomsoni are reported from
Deccan intertrappeans (Kateru beds) of Andhra Pradesh.
31.10 MICROSCOPIC PARTS OF PLANTS-SPORE-POLLEN

\ Spore and Pollen
. .r study : Study of fossil spores and pollen forms a separate and important branch
Ad vantage o,
· h' · I ontology known as 'Pa Iy,ro I ogy ' . S.mce the Iast f-I fty years, extensive . work
wit m m1cropa ae, ..,·1 spores and pollen and the data Imve been success tu · II y app 1·1ed in
· vanous
· ·
has been done on I oss1 · d
· e such as in geology, glaciology, archaecology, systematic botany an
ot~er_ branc~e.s ~f _sc1enc Because of their smaller size compared to other microfossil~, r.hey have "'
d h . ~ sils are available in large number wuhrn rocks.
.I
also m medical science. .
a greater chance of preservauon an t eir os.
Palae(Geo)WP-60
l'AI .A 1':0NTOI.OQ y
472.
Male Spores
\
llAC()S\S
Microsporanglum
\With Male Spore • •
.0) -
• ••
• •: 0
i!_,)
0 ~
1IAPL 10
\ ~
/ " \ "1\C~"· Fcnutle Spore
' : t1 ~ Ol:NFRATI N
·· Mcgasporangium
\ W'th
•
Female Spore's \
n---t---Fcnilc Stem , J'

', I~
, Fcmatlc
l'- ~ \. DIPLOID ', Oamcrophytic
Adult Sporophytic Plant(2n) GENERATION \ Plant(n) (n)
\ Egg(;}
,~,~ .
b ( S ~.Y ll'S·
~
Embryonic
~
(2n)
Zygote\
.,,·l,1\\\01\
r,ert'
·_ , , /
Sporophytc (2n)
(a) LIFE CYCLE OF A PTERIDOPHYTE ANO ~RMATION OF SPORES
•
\
\
....
Pollens (n)
' • • • • ISi
\ • : ••~AMETOPHYTE STAGE
' 0 •
'
'\
\ Pollen (n)
\
Carpet---~
(Female Reproductive
Body) Pollen Tube
SPOROPHYTE STAGE
(h) LIFF. C'YC'I.F OF AN AN<llOSPERM SHOWING FORMATION OF POI.I.ENS
FIG . J 1-17 : LIFE CYCLE OF PLANT AND FORMATION OF SPORES A NI POLLF.N
... -
473
Not onlys?, becau~e of their ~reat power of dispersal through air, many of them become good
index fossils used m correlations of rocks. Moreover, this mechanism of dispersal of spore-
pollen make their fossils available in all types of rocks deposited in basins formed under different
environments. For example, foraminifera fossils are available only with marine rocks, but fossils
of pollen of a continental plant may be found not only within continental sediments but also
in contemporeneous marine rocks.
In geological science spore-polJen are mainly used in :
(a) regional and intercontinental correlation of sedimentary rocks of Devonian and younger
age.
(b) tracing strand lines or shore lines as spores and pollen always tend to concentrate in a
large number along the shoreline with other microplanktons.
(c) interpreting the changing pattern of past climate evident form change of types of spore
and pollen, thus indicating· a change of vegetation.
Spore-pollen in the light of life-cycle of a plant : In general, spores may be defined as
reproductive units of non-flowering plants (cryptogams) while pollen are male reproductive units
of flowering plants (angiosperms). Gymnosperms also produce similar elements which are
however inferior to pollen often called pre-pollen. To understand them clearly, it is essential to
have a knowledge of life cycle of lower and higher group of plants (Fig. 31-17).
A plant of lower group usually exhibits two generations in a complete life cycle, one
generation alternating with the other. Such phenomenon of alternation of generation is also
found among lower group of animals such as in foraminifera.
In pteridophytes, a sporophytic plant (Zn-chromosome) asexually produces millions of spores
(n-chromosome) through meotic cell division of spore motlier cells formed within sporangium.
Germination of these spores gives rise to gametophytic plants that bear male and female
reproductive organs either in same plant (for homosporoes) or in two different plants (for
heterospores). These reproductive bodies on maturity release numerous male and female gametes
(n-chromosome) through mitosis cell division. Sexual union of two gametes produces a zygote
(2n-chromosome) which germinates to a new sporophytic plant. The Haploid generation (n-
c_hromosome) starts with the formati_on of sp?res (n) and end~ before the formation of zygote
(2n) while the short deploid generation (2n) mcludes the portion of life from the formation of
zygote before the formation of spores. In low~r group of plants like mosses and pteridophytes,
the gametophytic generation represents the ~am plants and this stage of the life cycle becomes
much pronounced. In mosses, the sporophyt1c form sometimes is reduced merely to a parasitic
outgrowth on the body of a gametophyte. Moreover, such plants in gametophyt· t ·
.. . . 1c s age reqmre
marshy condition for keepmg their male gametes alive during its moveme t t d "
" f · I h · h n owar s aema 1
i · ·
gametes aor ert1 1zat1on. n 1g er seed-bearing land plants this g t h · · e
becomes h1g . h . . ame op yt1c generation
ly reduced and the mam plant 1s represented by spo h · · h" h
.
be ars fl ower t hat pro. du.ces mobile rop yt1c generation w 1c
pollen or male gametes (n ch ) d h ti I
. . - romosome an t e ema e
g~metophyte remams captive on the body of a sporophyte, represented merely by the ovary
with one/more ovules each representing a female gamete ( h ) A 11 -
h. ti · n-c romosome . po en upon
reac mg emale stigma, grows as a long tube (pollen tube) that extends down through the tissue
- - -
474 PALAEONTOLOGY
......,.'""1"""tC----Wa\l
~ - - - Wall Internally
Striated
Densospori tes
lycospora
- - - - - Border
Spore (Bilateral Syn~metrical)
Spore (Radially Symmetriyal) Lael'ig11 r, ,s1u ,,., res
Colpa
Monocolpate
Tricolporate
Tricolpate
Triporate
(a) MORPHOLOGY OF SPORES
- - - - - - - P s i late
W~I { :::: _ _ _........i11e--;1
------Vermiculate
------Spinose
Pollen : Various Types Of Exine Ornamentation
Bi-Winged Pollen
(b) MORPHOLOGY OF POLLEN
FIG . 31-18 : MORPHOLOGY OF SPORES · AND POLLEN
-. .,
•
of the female organ towards one ovule and fertilizes it. This pollen tube represents a highly
reduced male gametophyte. After fertilization the new sporophyte is produced with 2n
chromosome, but it takes the form of an embryo or seed. Seed is essential for the life on land
to protect it form adverse conditions. Under favourable condition the seed germinates to produce
a new sporophytic plant.
Morphology of spores (Fig. 31-18) : In lower group of plants such as in mosses, spores (n-
chromosome) which on germination give rise to gametophytes are identical looking, called
lwmospores. But in higher plant like in pteridophytes they are different in appearance
(lieterospores) : the large-sized female spores, called megaspores and the small-sized male spore,
microspores producing male and female gametophytes. It is not possible in fossils to
differentiate a male and a female spore in case of a homosporous plant. In fossils, however
spores which are larger than 200µ are tentatively called miospores and smaller group of spores
are called microspores. Fossils of spores and pollen are together ca11ed polospores.
Normally, four spores develop from a single mother cell called spore-tetrad, formation of
which may take place in two ways. It may develop by division of the mother cell all at a time
and the tetrads will be arranged tetrahedrally. The spore produced by such a tetrad has a tri-
faceted proximal surface when the faces are meeting along a tri-radiate depressed suture line
called trilete marks and the spore is called a trilete spore. In other case, a mother cell may
spit first into two daughter cells, each of which splits again forming a spore tetrad. But here
the spores are arranged vertically joined along an axis (tetragonal-tetrad) and each spore derived
from such a tetrad shows a single depressed suture (parallel to the axis of attachment) called
monolete mark and the spore is called monolete spore. In some cases spores are found without
any scar or Iete-mark which are called alete spores. Accordingly, a spore usually exhibits a
tri-radial or bilateral symmetry. Lete-mark is also the zone along which the spore germinates
giving rise to a new plant. Spore wall in many cases exhibits granular, reticulate, spinose type
of ornamentation. The thickness of wall is also variable and a spore with a thick wall internally
may exhibit radial striation. Sometimes a flange or air sac (cavate) may be found in a spore
helping in its dispersal by wind. Spores are mainly classified based on its symmetry or type of
germinal aperture.
Morphology of pollen (Fig. 31-1 Sb) : A pollen is also formed by division of mother cells
like spores and its basic features depend upon the manner of this subdivision. But unlike lete-
mark, a pollen grain bears one, three or more circular or elliptical apertures called pores and
colpi respectively through which the pollen germinate at the time of fertilization of ovule. Pollen
are classified and named as monoporate, triporate, polyporate or monocolpate, tricolpate
or tetracolpate based on the number of pores or colpi present in the body. There are some
pollen which may bear both pore and colpus and they are called colporate (such as
monocolporate, tricolporate or polycolporate). A pollen grain without any colpus or pore is
called aporate/acoplate and in fossils it is difficult to distinguish it from an alete spore. Like
spores, pollen also exhibit radial or bilateral symmetry and show their shapes accordingly.
. Pollen grain has a two-layered wall; the lower thin-layer called inti11e and the outer relatively
thick
. ,
layer. called exine. Exme
· 1s
· a homogeneous membrane but my be deeply sculptured on
its o~ter side (tectum) by various types of structures like granules, punctae, reticulations,
vermicules, lobes, spines etc. A smooth-surfaced pollen is found rarely. Exine layer is too hard
~
476 PALAEONTOLOGY
to be destroyed by common natural hazards and that is why pollen are preserved beautifully
and in large number within rocks.
Besides these, many pollen (especially those of conifers) bear a pair of wing-like structures
or air bladders that help them to disperse in air for a long distance.
Pollen grains are comparatively smaller than spores having a size between 1Oµ to 80µ and
rarely above I00µ. Usually, they are classified mainly on such characters like symmetry, nature
of germinal aperture, nature of exine layer or presence of other accessory features like air
bladders, wings etc. Quite obviously, the classifications of fossil spore and pollen are artificial
as they are difficult to compare them with the spores and pollen of their living counterparts.
However, Neogene spores and pollen have much similarity with those of recent plants. Numerous
TABLE-30
A COMPARISION BETWEEN SPORE AND POLLEN
SPORE POLLEN
Definition
These are reproductive units giving These are male-reproductive units of a
rise to new gametophytes of flowering plant. They sexually unite with
non-flowering plants. ovule (female reproductive unit) to form a
seed that germinates to a new plant
(sporophyte).
Number of chromosomes
Haploid number of chromosomes in Haploid number of chromosomes in the cell.
the cell.
Symmetry/Shape
Symmetry radial or bilateral; shape Symmetry radial/bilaterial, shape variable,
variable spherical, pyramidal, spherical, pyramidal, ellipsoidal,
ellipsoidal etc. bi-winged forms are common.
Outer surface
Outer surface smooth sometimes Outer surface normally ornamented by
coloured/ornamented. structures like granules, hairs, spines,
reticulations etc.
Nature of wall
Wall single-layered, thick/thin Wall double-layered, an outer thick exine
internally straited.
Germination areas
and in inner thin intine.
--
Present/absent; present in the form of Present/absent; present in the forms of
a line of depression called 'lete markes' pores or depressed elongated zones ·
may be one or three in number. 'col pus'; both may be present in some forms;
the number of pores or colpi variable.
*S : Spore; P : Pollen
JNTROl UCTION TO MICROPALAEONTOLOGY
477
fossils of spores and pollen are reported from Upper Palaezoic to Quaternary rocks from various
parts of peninsular and extrapenisular India. A few spores and pollen from Lower Gondwana
rocks arc : l eiotriletes (S)*, Punctosporites (S), Parasaccite (P), Platysacccu.\· (P) etc. A
comparison of morphology of spore and pollen is given in Table-30.
Ecology and distribution :

Ecology of spores and pollen reflects, of course the ecology of their parent plants, but rarely
spores and pollen are preserved in sediments with the fossils of the parent plants because of their
great dispersal. So ecology deduction from spore-pollen fossils becomes more uncertain unless
one can identify the parent body. Wind dispersed grains are produced in greater number than those
dispersed by insects mostly because of uncertainty of their utilization. Small-sized miosporcs arc
often found in wind 350-650 meter above the land surface during day but they sink during night
or may be brought down by rainfall. Under favourable condition pollen have been known to drift
more than 1750 km. but mostly they will settle within I km. from the source. A few may reach
the ocean by aerial dispersal. As they settle, they may fall directly into lakes and swamps and
may be preserved within sediments in these basins or they may be carried by water into rivers,
estuaries and ultimately into the oceans. During this transportation there may be sorting of the
pollen-grains Larger megaspore tend to settle in rivers, estuaries and shallower deltas and shelf
areas, and smaller microspore may reach even in deep sea.
Being" both light-weight and resistant, spores and pollen may suffer several cycles of
reworking and redeposition leading sometimes to confusion about the fossil record. Experienced
palynologists are able to detect these reworked forms by observing differences in colour,
corrosion, size, stratigraphic inconsistencies etc.
31.11 APPLICATION OF MICROFOSSILS IN PETROLEUM EXPLORATION

The application of microfossils to practical problems of petroleum geology resolves itself
into a number of distinct stages. One important observation is that most of the world's petroleum
deposits are located within marine Tertiary rocks rich in foraminifera. That is why in any
petroleum exploration work study of foraminifera becomes essential for establishing local and
regional stratigraphy of th~ area under investigation.
In all oil exploration work two facts should be clearly realized. First, the oil genesis that
requires several physical-che~ical-b.iological and temporal _factors to be favourable resulting in
transformation of organic debris t_o 011 and the seco_nd, the 011 accumulation which again requires
1>eve~al _fact~rs .lik_e rock properties, structure, fac1es change etc. to ~e fovourubk for trapping
~he liquid 011 w1th111 the rocks. In nature, howeve~, a ~~rfect_ c~o~dmat1on among all these factors
1s not a common event and hence also the ava1lab1laty of 01) m the •sub"lla·t··,c~· ., · ' ....
Tll"'C hUSIII
· ·In
which oil exploration work have to be done may be a vargin busin or u basin which is I , I
. Id. ·1 J . .
y1e mg 01 . n a v1rg111 area
h t· I . -
I un llaItrcu, y
. t e sur ace. geo . ogy 1s first to. be .studiccl c·,r~·t·aall
( .. o
y unl emp 1
should be made to ascertam• the age of different outcro1Jp1r1g "o,· , t·
' 111 ,1 1011 •
s bnscc. on macro· t·oss1·, s
• •
1
1
and megafoss1ls and/or rad1ometnc data for purpose of general ori ·nt t' F hi . • ·h f
· Id b · · d. . c u 10n. or t s, cue o
the .formations shou c rnvcst1gate
. . 111 detu1ls and their rcSJJecti've
· d'. 1s·t·mc,1vc
,· macropa
· 1a ·o rt>
1< -
logical features arc established. This may help to establish the biostrnta· ,pti,·c subdivisions of
i'I.
· · I b' Th ' · gu .
the format.ions mto severa 1ozones . . . 1s 1s the most time absorbing part of this pllm11ion
work, particularly when a large area 1s involved showing rnpid focies change. Th, contrihutio11
PALAEONTOLOGY
478
eld and success of this exploration largely depends upon the time
to the deve lopmen t Of the fi · ·b · f · f
l study of horizontal and temporal d1 stn utlon o micro auna within
spent on the fun dam e 1lta·
the rocks in the area of investigation.
Shallow drillings at intervals are then carried out which provide_ materials for local
correlations and finding out the structures exhibited by the rocks. B.efore gomg for a deep drilling
a preliminary seismic survey may be quite helpful to ascertam the nature of rocks lying
subsurface. Deep drilling operations are then started in an effort to puncture the c~p rocks and
to strike the oil reservoir. From the start of deep drilling a constant cheek on mtcrofauna in
the mudflow of rotary wells, in well-cuttings and cores has to be kept according to a prearranged
plan of sampling. Analysis of microfossils data from surface and new data from subsurface
core materials will help faunal correlations between the well section and surface sections.
Whether a deep well will strike a oil reservoir or not, is difficult to predict due to the
uncertainties about the nature of structure and facies of the rocks lying deep below. Whatever
may be the fate of the wells, the microfossils available from the different core samples and
wells should be investigated and from the available stratigraphic cum palaeontologic data,
correlation of the subsurface rock of different well sections will be established. At this stage,
micropalaeontologists will be able to identify some litho and biomarker horizons. With increasing
knowledge of the distribution of fossils in the marker biozones of the oilfield and with gradual
improvement of the technique of examination and correlation of local fauna, the originally
established sequence of zones should be constantly checked and if necessary revised.
If the basin contains oil and if a well strikes a oil reservoir, the entire attention should be
focused on that particular well to understand the local criteria as the detailed knowledge obtained
from that well helps in the placement of future productive wells.
The location of the future well then will be guided by the special distribution of the
planktonic biozone/s corresponding to which the oil has been located. Similarly, in an already
oil producing field, location of future wells should be planned according to the spatial
distribution of the recognised oil-bearing formation.
For a large field, for getting maximum speed in the investigation work along with greatest
~ccur~y, oil companies often carry out work by setting branch laboratory where the entire work
1s earned out by trained assistants under the supervision of some experienced palaeontologists.
In. most of th~ cases, generic identification is sufficient and the new genus is then numbered
with the mention of the formation from which it is obtained. Thus Nodosaria Tmt-1 stands for
~ definite species of Nodosaria from Tertiary-Miocene Timpan Formation of Assam. For rapid
1denti~cation of genus/species, type speciments should be mounted along with their photographs
and with short morphologic description within a small card.
~ ~s there is a cl~se relationship ?e~ween environment and the organisms living in i!·
environmental analysis of host rocks of 011 containing in situ fossils may be made. Such analysis
of host roe~ and sou~ce r?ck may help to coordinate the entire case history of a particular oil
field. That_is, the ent~re h_istory of formation of oil, its paths of migration and its accumulation
can be. estamated. This wall serve as· use
· f uI document ·m the research and development w,ng· of
the oil-company which
. . will pl·ay a. m,tJor
, · role ·111 the placement· of future wells with greater
ti
con 1dence and scientific logic.
PART -VI
STUDY OF FOSSILS
..
I
1
I PRA Chapter 32
CTICAL WORKS ON FOSSILS
I. COLLECTION AND p
AND METHOD OF RTEHiPEJ\RATION OF FOSSILS
IR DESCRIPTION
32.1 COLLECTION OF FOSSILS IN THE F
Collection of fossil in the field-site . _IELD
· s is a maJor attraction f I ·
first questions to be asked by them is h d or pa aeonto 1og1sts. One of the
O
the amount of preparation planning w dere :e look for fossils? The answer should reveal
professional palaeontologi;t or a ded·ant dcare t at goes into the work of finding fossils . A
tea e amateur has to spend many lo h d ·
published reports, maps and field data whi h I h ng ours stu ymg
occur, t es of the rocks . . c can revea t ~ places where outcrops of interest
yhp , . m which they occur and the equipments and techniques needed to
co. 11ec t t e ,osst 1s from those sites
. · E qmpments
· ·
or techniques · · obviously are different for
quite
differe~t rocks types and also different for different groups of animal and plant fossils. Collection
of fossils fro~ a ~ard sandstone, cherty rock or hard limestone needs quite a different method
than th.at which ts necessary for collecting fossils from softer rocks like shale and marl.
Collect1on of samples from a highly weathered shale or marl requires other techniques. But a
good field palaeontologist will not set off without geological hammer, chisel (small and large),
brush, pocket knife, hand lens, pencil, a note book, a climometer compass, specimen boxes
(large and small) measuring tape, marking pen, camera, paim ·brush, loose papers, satchels, glue,
plaster of paris and a geological map of the area. (Fig. 32-1 ).
Collection is easiest where fossils have been weathered out and rest on the surface such as
on river sand and cultivated land or in desert sand. In case of microfossils they may be collected
within small cotton satchels or polythene bags. Larger fossils from such localities may be picked
up by hand. Fossils may be entombed within relatively harder rocks like limestone, sandstone
or shale (less weathered). It may be possible to break off the rock enclosing the specimen using
the blunt head of a hammer, or splitting the rock using the sharp chisel-edge of the hammer.
Failing this, one may use the hammer and chisels to cut the host rock around the specimen,
carefully detaching it from the rock. For a still harder fossiliferous rock, collection of fossil-
bearing rock is preferred. It is better to lift a large block of rock containing a fossil than to
damage the specimen to be collected. For a large but a beautifully preserved specimen which
is impossible to be removed from the host rock, plaster of paris may be used to take a mould
of the specimen in situ that can be reproduced in the laboratory by a cast. In any case,
destruction of such a fossil trying to separate it from rock, is quite unwanted. Let other see the
fossil. Plant fossils are commonly found within carbonaecous shales or other shaly rocks that
are usually finely laminated, fissile and soft in nature. So _special. care should be tak~n for
collection of such fossils. Plant fossils are mostly found as 1mpress1ons, usually occurring on
the upper surface of each bedding plane. So rocks have to_ ~e slitted along the_ different bedding
planes to get more and more fossils. For this, sharp kmfe, sharp edged-chise l may be used.
481
f·
. ! . lJ . \.,; •\St.A. ·. ,$ ,~Jti !·. ..
~
I i t µ . ..,,. QJ .
PALAEONTOLOGY
482
~
~ Chisel
Paint Brush
Hammer
Note-Book & Pencil
Cloth-Made Bag
Papers
Collection Box
FIG. 32 - I : FIELD EQUIPMENTS RELATED TO FOSSIL COLLECTION
• '.. · J_.1 ,
'RODUCTJON TO MICROPALAEONTOLOGY
ffei ~3
wever, petrified wood or fruit may be found in embedde .. . ·
1-l~ding planes. For silicified fossils it is easy to s . dhcondition often cutting across the
t,e . I eparate t em from rocks by using hammer
and ch1se .
Fossil skeletons of a large vertebrate (say dinosaur) may be & d . h'
d • 1oun wit m rocks as fragments
ich may occur scattere over a considerable area Coll t' f .·
wh . . . . · ec mn o one or few skeletal fossils
of a dinosa~r m a parti~ular area IS Just a clue of possible occurrence of the entire skeleton of
the animal. m that localtty.. The
. probable area in which the.· bones. 1·1e 1s
· gn'dde d on topos heet.
The locat10n of eac~ fossil. is. ma~k~d on respective grid. It shows exactly where and in what
pasiti~n e~ch bone ltes. Gnddmg 1s important because it shows which bones Jay close together
and this will help to recon~truct the skeletons in museum or laboratory. Intensive work for several
days/months ~ay be req~tred for exposing a complete skeleton or a large number of skeletons
of different animals. Taking out of fossil bones from a rock depends on its nature. If the rock
is soft the bones can be easitr dug · out, carefully of course, to avoid damage. If they are
associated with hard rocks, a slow and careful process of removal by chipping around each
individual bones is preferred. Sometimes bone fossils are too fragile or weakened to be removed.
In this case, it is necessary to strengthen the fossil by adding some liquid plastic substances,
which seeps through fractures of the fossil giving it extra-strength. Fractured and broken bones
are treated just like a hospital treats broken limbs; they are given plaster cast.
All fossil specimens removed and collected from rocks should be properly numbered
mentioning the locality and number of the fossil. For example, a number may be as such
4/12 meaning that the specimen is collected from the locality 4 and is the 12th specimen of
this locality. Besides this, geographic and stratigraphic position of the exposed fossilferous rock
should be noted in the field note book. Specimens should never be prepared or cleaned in the
field for the field tools are unsuitable for this purpose and this needs delicate and patient work
in the laboratory.
32.2 SEPARATION OF FOSSILS FROM ROCKS AND THEIR CLEANING

Fossils collected in field, often need challenges when studied in the laboratory. Mechanical
and chemical methods of preparation can be employed to reveal morphological details of fossils
unseen during field collection. Normally, the methods chosen must be suitable for the sediment
and the fossil to be prepared. Specimens may be extracted from clay or sand only after a few
hours of soaking in water; other may needs a few days. Others need several weeks of careful
works by knife, fine needles, small hammer and chisel, drills, and abrasives. Obviously these
fossils and rocks are hard enough to be fractured or chipped. The common chemical method is
to use suitable acids. Care must be taken in choosing the type of acid (which would be different
for different rock and fossil materials), and selecting concentration of the acid failing which
th~ ~ossil may be destroyed or becomes highly fragile or delicate by the acid itself. Siliceous,
c~ihnous and calcium phosphatic fossils embeded in limestone may be extracted by the use of
dllute hydrochloric or acetic acid. The former is best-used during the preparation of invertebrate
fossils impregnated by silica. Specimens should be immersed within acid having less than 10
percent concentration. Acetic acid is preferable for vertebrate fossils for their Ca-p~osphatic
com.position. The problems becomes critical with the calcareous fossils for which very weak
acetic acid or solution of sodium carbonate may be used.
;;:&, " · - , . ; ,: . I 1119 ...
Pi\LAl:-ONTOLOG't
4R4 . . 'l'h ·
· · ·.d ·11 Ould h· 1.t, 11 ow .(
. I I·, y w·i•·hi'
P•
n the foss il in water. c specimen, then
Immersion Ill nci s . d .. • , uurt··H.:e should be protected by application of
b d . d ,111 d th " n ·wly ·xpos. 1I s1• " •
should e ne · • . b. . . .. l ·d until the fossil is completely exposed. Finally
thinn ·d down glu ·. This proc ·ss may r pc,1 . I· . . . . . ·h
. I I I u co·1ted by ·1 p asl.lc v.irni s .
the specim ·n is washed, dncc ' c can ·t an • • .
· .1 · hbontory or museum by several waytL Ultrasonic-
leaning of v ·rt ·brute bones is uonc II1 ,. • ' h h . b
.'b . . h, ,· . .·, .. 0 that the loose rock ,rain s arc shaken off. Ca-p osp at1c ones may
bath v1 r,,tes t c oss1 s s b . I d b
· .. • . k . by using light acid. After crng c cane up, ones are
be sep·u·,tcd trom cu 1l:,1reous roe s . f . F. II
· ' • . d .· . . ~ ' ial g•luc Holes are filled up hy plaster o pans. ma y for
repair•d and protccte . using a spec · · . . h h
a tong-time preservation u protective coating of plastic varnish has to be put over t e w ole
bone.
For separation of microfossils from rocks two methods are usually followed._ Disag~reg~tio_n
method is suitable for fossils of foraminifera, radiolaria, ostracoda etc. For thi s, fossils ~1thm
loose and weathered rock-samples are preferred and collected. Slightly .weathered ~ock-s~1mens
have to be crushed upto a preferable size. These crushed rock maten~Js are boiled w1.th equal
amount of sodium carbonate or hypo for an hour. Finally the materials are washed m water
repeatedly and dried on hot plates. For siliceous microfossils, embeded within calcareous
sediments, dilute hydrochloric acid may be used within which the materials should be immersed
for several days. The dried up materials is then allowed to pass through sieves with different
sized-meshes and these different sized materials, may contain different groups of microfossils.
Each sized-materials will be kept on a glass tray or porcelain plate for observation under the
stereomicroscope and fossils are picked up by a sharp needle using a light glue at the point of
needle. The fossils are usually collected in an assemblage tray for further study and for
separation of different groups. For detailed study and observation ·of some microfossils such as
larger foraminifera, palaeontologists have to prepare oriented thin sections of specimens on glass
slides. This work need a considerable and thorough knowledge of morphology of the fossil
concern.
Spores, pollen, dinoflagellates, diatoms and such other microfossils are separated from rocks
(usually black shale or very fine grained sandstone) by maceration method. For this the rock
specimen is crushed and then immersed in 52% (N) hydrofloric acid for 16 to 17 hours. Then
the acid is removed by filtered water within the centrifuge apparatus. Then the material is kept
within s.chultze's solution (KCJ0 3 : HN0 3 - I : 3) for 2-12 hours and again is made acid free
by previous method. Coal or carbonaceous shale where plant materials (spore-pollen) occur.
they ~ave !o be kept within I 0% KOH for 2 to IO hours and again centrifuged. The obtained
material will be rich in microfossils. Usually for spores and pollen, type of maceration depends
~~n age of the r~cks. For ex~mple, for rocks younger than Quaternary there is no need of 3rd
st~ge of macerat1~n. After this~ o.ne drop of macerated materials is put on a glass slide and
dn~ up by a des1cator. Then 1t 1s covered by a covership after addition of some mounting
m~daum s_uch as ~lycerinc .or je.lly. These are usually observed under a biological microscope
using lenses of high magn1ficat1on (200-1 OOOX).
32.3 DE~C_RIPTION OF FOSSIL SPECIMENS

p Description
d fi . . of fossils
. · ·.
is one of th~e fun damenta I task of nearly all palaeontological wor ks.·
or e mmg a new species
• •
or for 1'den t'li · o f a specimen,
1 1cat10n . . ·
description should be systematic
,, •
.,. ~ . ' . . -.,
INTRODUCTION 1'0 Ml UOPALAHONTOLOGY

48.5
s
I s
'
8-Valve
!S
Brachiopoda (Bilateral) I .s
•s Cephalopoda (Bilateral)
s. Pelecypoda (Bilateral)
I
I
s I
Is Irregular Echinoid
Arthropoda (Bilateral) (Bilaleral)
~
•
s
Tetrapoda Vertebrate ' .
(Bilateral)
sl•
Regular Echinoid
(Radial- pcntamerus)
liexacora1 (Radial)
. .J
FIG. 32 • 2 : NA'" r
S I u_RE OF SYMMETRY WITHIN DIFFERENT GROUPS OF ANIMALS ·,.;L
'
-S Lme Represents Trace Of Symmetry Plane . ,#-
; -•
~-----_.._._.,.• •1_;0Sii9~
~
.•,,
• !
,
486 PALAEONTOLocy
Hemispherical
Conical Cylindrical Tubut.
HatSbaped
Top Shaped
Pyramidal Vennicular Ceratoid
Acicul• Slipper- Shaped
CJ
Heart-Shaped
O· c
DiscoidaJ
:>-G Kidney-Shaped
Fusiform
~ Hoot-Sblped
~ V7 Boat-Shaped
(a) SHAPES DENOTING THREE-DEMENSIONAL FEATURES
ooaDo oo<>Q..667
Circular SubcircuJ• Triangular Rcctangulor
Hexagonal
Elliptical Rhombic Lanccolatc Cllordalc WedlC
shaped
(b) SHAPES DENOTING SOME TWO-DIMENSIONAL FEATURES
FIO. 32 - 3 : DIFFERENT TYPES OF SHAPES FOUND WITHIN ANIMAL SKELETON
,,,rRODVCTION TO MICROPI\LAEONTOLOGY
487
and as complete as possi_ble. Th_is enables a worker to compare his specimen with other similar
fortll~ _kno_wn from published ~tteratures. Before a systematic description of the specimen its
id~ntthcat1on upto the level ot phylum and class is necessary from accurate observati on and
examination as this is a ~rec~ndition for identification of a genu s. After thi s, it is po sible to
follow a scheme appropriate f'or proper description of that genus and that will lead the worker
to identify the genus/species by comparison with other known similar form s. For example, by
careful observation of a specimen, it is quite probable to identify, wheather specimen is a mollusc
or brachiopod, and if it is a mollusc, whether it is a grastropod, cephalopod of pelecypod.
[)e-Scription of the specimen follows accordingly. Say, the specimen is a pelecypod with taxodont
teeth, then for its generic identification comparison will be limited within such pelecypods only.
And, if the species is identified as Arca, then for specific recognition it is necessary to compare
the specimen with description of other known species of Arca. If it appears totally di s. imilar
from others, the worker may think of erection of a new species of Arca. Erection of a new
species based on a single specimen is usually not preferable as any species usually shows a
considerable variation of its character which can be studied only by observation of a large
number of specimens. For a new species, the author should select the type specimens and
preferably also a holotype and he must be guided by other codes of nomenclature.
One of the main problems of description is to choose the attributes. Many features that are
seen in living specimens are not available to a palaeontologist who has to base his description
within the restriction of fossilized materials. For example, weig~t, volume, skeletal composition
of the animals may be greatly changed after fossilization. Generally, skin or muscle characters
are also not visible. It is often difficult to differentiate between an amphibia and a reptile from
their skeletal morphology only, because their difference is mainly biological. Palaeontologists
thus has to select such attributes which appear to be significant both biologically and
palaeontologically. Selection of these significant features is a matter of subjectivity, the effect
of which may be partly eliminated by observation of a large number of specimens using so me
stati stical methods.
There is no single format to be followed to produce an ideal description. However, some
common aspects of the description of a fossil-specimen have been discussed belm .
A. Symmetry (Fig. 32-2)

Leaving aside plants, most of the animals exhibit a definite symmetry in their body often
show~ by their skeletal pattern (normally preserved as fossil). An animal's body usually ~xhibits
t~o kmds of symmetry : radial and bilateral. In the former type there me more than one p_lane
of symmetry, while bilateral symmetrical forms show only one such plane. A_mong _the _unmml
gro~ps showi11g radial symmetry are some protozoas, poriferas, worms., annelids, cmdunus und
echm~derms. Animals which are mainly bilaterally symmetrical include urth~·opods'. molluscs,
brachiopods and chordates. However, in some cases the primary symmetry ot an anmm~ gro_up
ma~ be superimposed by a secondarily developed symmetry. For exnmpl~. corals ure pnmanly_
radial, but rogose corals show a secondary bilaternl symmetry as shown by th~ nut~re .of
~;;eio_pme~t of their primary and other major septa within the;, embryonic caly~. Ag~un ,echmo1~s
. . primarily radially symmetrical but this is superimposed by a secondury b1luteial _symmeuy
1 ~llocylic echinoid (irregularia). Primary bilateral symmetry of soml! molluscs is lost and
~:~.n.
Pal castropocJ shells become asymmetric. Symmetry character O t· s·ht: ti s· O f pe
sp1ral o· . lt:cypods·
ae(Geo)Wp_62
488 PALAEONTOLocy
and bivalved brachiopods is also different although both of them are bi~aterally symmetrical
In a brachiopod shell this symmetry-~lane pas~es a~ross the two valves m a posterior-anterio;
direction (length) cutting each valve mto two 1dent1cal halves. In a pelecypod, the symmetry
1 ne is passing in between the two identical valves.
Pa ·
In both arthropods and chordates th
"d · I h I · e
bilateral symmetry plane cuts the body laterall~ mto two I e~t1c~ ~ ves ~assmg across head
to tail. However, it should be remembered that internal orgamz~t1_on mcludmg soft parts of an
animal may not always conform with its external symmetry. This 1s true for most of the higher
animal groups where the nature and position of internal organs are very complex.
B. Size
Size of an animal is also an important character. This can be referred to qualitatively as
large. medium or small and quantitatively by direct measuring the dimensions (length, width,
thickness etc.) Broadly, the whole organic world may be classified into two groups based on
the size: microorganisms and rnacroorganisms or megaorganisms. The former group are visible
or at least recognisable only under microscope. A new phylum 'protista' has been sometimes
recognized which includes all such microscopic animals, plants/bacterias and others. Such
subdivisions are also accepted in fossil world, named us microfossils and megafossils. However,
microfossils may include another group such as microscopic parts of some larger animals or
plants which are separately fossilized. Fish-teeth," insect-appendages. echinoid-spines, spores-
pollens of plants are such fossils. Size of an individual fossil species/genus may not be its
diagnostic property. This is because all animals usually exhibit increase in size with their gradual
ontogenic development and size of an individual may indicate a particular stage of this ontogeny.
Juvenile form of an animal are always smaller than its adult form. Comparison of size of two
individuals is only possible when one is sure that both of them represent same ontogenic stage
of development which is however difficult to ascertain from study of one/two fossil specimens.
Size may be an important character when a palaeontologist works with the fossil populations
of different species and tries to compare them. Here size of a large number of specimens in a
population of a species is measured directly to find out the value of the average size. To get
accurate value of size one has to measure the three dimensions. length, width and thickness
separately. For better comparison one can plot all these data graphically which may show
difference in size range between the two species.
C. Shape (Fig. 32-3)

As an animal possesses a definite symmetry, it also usually exhibits a suitable shape which
will conform with its symmetry. Normally. the shape of an animal should be described
considering it as a three dimensional object having definite length. width and thickness. For a
pro~r visuali~!ion of shape of an animal it should be viewed along three different pla~es/
sect1o?s ~ontammg length-width, length-thickness and width-thickness. A perfectly sphertcal
bod~ ·~ circular in all these sectional views; hemispherical body is circular in one view and
sem1-~1rcular in other two views; conical body is circular in one view, but triangular in ot~er
two views and so on. In many cases, shape of an animal is described observing its similarttY
with ~other com:°1on ge~metric forms having approximately the same shape. The animals, ofte~;
rnu:
desc~abed as. lent1cul_ar (hke ~ le~s), oval (like egg), tubular, cylindrical or ellipsoidal,. all
be circuiar m top view. Again, it may be described from its overall resemblance with sorn
common objects like pyramidal. acicular (needle-li~.e), vermicular, horn-shaped, lanceolate,
. ..,, ,,
JNTRODV TJON TO Ml ROPAIA/:.'ONTOL OGY

489
spatulat • (tongu ·-shap ·~). s:.mdle-sha~cd, to1~-shaped, lull-shaped, boat-shaped, hook-shaped,
heart-sha? •d e~c. Som ·t1111cs shape ?t a particular genus may be distinctive and the shape of
all such forms Ill other genera are referred to by the name of the genus, for example, volutiform
(shape of Voluta), mytiliform (shape of Mytilus), turrilifrom (shape of Turritella), crypreform
(shape of Cypr 'll) etc.
D. Description of other chuructcrs

Apart from the three fundamental characters, symmetry, size and shape, a fossil of an animal
may possess various other characters diagnostic for it and for a systematic description of the
fossil these characters should be described one by one. These characters may be chosen in order
of their importance in animal's body or serially following any other systematic methods.
However, choice of characters in order of their importance would be different for different
groups. The choice of such characters become somewhat subjective as an author may consider
one particular character more important than the other and others may have different opinions.
Thus it will be more appropriate to select character by some other means. For example in
description of a solitary coral fossil one may start from the features on external surface followed
by the internal characters. For brachiopid fossils, description may follow the features serially
from posterior to anterior direction, ending with its surface sculpture. For palecypod, shell
features of external side may be described first followed by internal features in a direction from
umbo to ventral margin.
In all the cases, the features of a fossil which have been described should be illustrated by
sketches and/or line-drawings of the specimen with proper labelling. A good photograph of the
specimen may be sometimes highly illustrative. Besides normal conventional photos, x-ray and
SEM-photos may be shown for illustrating such characters which are usually not visible in naked
eyes, or in normal microscope (such as shell-microstructures, bone-microstructures, spine-
microstructure etc). Microfossils are described from microscopic observation and are usually
illustrated by common sketches, microphotographs or line drawings drawn under camera lucida.
II. DESCRIPTION AND IDENTIFICATION

OF SOME FOSSIL GENERA
Model schemes of description of fossil-genera of common invertebrates and plant groups

together with the description and illustration of some common genera of each group are given
below. This will be helpful for students of palaeontology in doing practical work with fossils.
32.4 PHYLUM : PROTOZOA

Class : Sarcodina
Order : Foraminifera
Scheme of description
5
1· Symmetry of test; 2. Shape; 3. Size; 4. Nature and tightness of coiling (for coiled form);
· Nrrangement of chambers (uncoilded form); 6. Shape of chambers; 7. Septa and septal suture;
8
· ature of aperture; 9. Surface ornamentation; I 0. Wall-structure.
490 PALAEONTOLocy
A. Larger foraminifera
Nummulites (Fig. 32-4a)
External view : Test bilateral symmetrical; large/moderately large/small; lenticular th' k
. I/s1gmo1
thin; planispiral, convolute; suture radia . .daI/ret1cu
. Iate; sur f ace smooth, a 'central
IC /
pustule often present.

Equato rial section : Circular in outline; protoconch small/large (microspheric/
megalospheric); whorls numerous/few; whorl height increases gradually; chambers
rectangular, septa curved backward.
Axial st: rion : Lenticular in outline; equatorial chamber-plane nearly straight, triangular
'quatorial chambers; alar prolongation from equatorial chambers separates one whorl form
other; whorl-wall peripherally thickened by marginal cord; pillars occasionally present.
Age : Palaeocene-Oligocene.
Assilina (Fig. 32-4b)
External view : Test bilateral symmetrical, discoidal, thick/thin; planispiral, involute;
whorls visible from outside; spirally arranged granules, bigger towards centre; septal suture
radial; central pustule often present.
Equatorial section : Circular in outline, whorls coiled planisperially; initial chamber
circular small/large, adult chambers squarish; septa thin, straight, slightly curved near
upper whorl; whorl-wall thick, aperture at the base of each septum; whorl height increases
rapidly from centre to periphery.
Axial section : Flattened, lenticular in outline; equatorial chamber-plane nearly hqrizontal;
coiling semi-involute (wall involute, chambers evolute); chambers semi-elliptical,
peripheral wall thickened by marginal cord; pillars few.
Age : Palaeocene-Eocene.
Alveolina (Fig. 32-4c)
External view : Test bilateral symmetrical; ellipsoidal with slightly obtuse poles; suture
lines straight or slightly curved, depressed, extending from pole to pole; surface smooth.
Equatorial section : Circular in outline with whorls planispirally coiled, convolute; whorl
divisible into narrow elliptical chambers with thin curved septa; embryonic chamber very
small; whorl-wall very thick due to flosculization, imperforate.
Axial section : Elliptical in outline, chambers long, narrow, tubular extending from ~le
to pole, slightly alternating; each chamber divided into very small rectangular to ellipucal
chamberlets by very thin partition-wall perpend.icular to septa; whorls wall very thick.
Age: Eocene
Discocyclina (Fig. 32-5a)
Exter11al view : Test bilaterally symmetrical, thick/thin, broadly lenticular, circular in
outline, often saddle shaped; surface finely granulate or papillate; a thin peripherial flan£~
may be present.
,------Marginal Cord
Sigmoidal Septa!
----Equatorial
Filaments
Chamber
Pru1m;o n1: h
1::..--1.~~.....q~,...~t----~ Large
f \1 c:g:i lm: ph..:ric 1
Sit.I\! View ~ - - - - - - - ~,;plUIII
Curved
Top View fatcrnal View Backward
~lilliliiiiiiil~....., ---Whorl Wall
Equatorial Section
Equatorial Chamber
Marginal Cord
-----Alar Prolongation
(a) NUM1\/Ul1T£S r------- Whorl Wall

- - - Marginal Cord
Granules
·..nlt.'lllt----Equatorial
Suture
ca:vw:2> Side View
Chamber
Pro101:0111:h
Large
Whorl (Mcgalospheric >
Visibh: External View
Outside Top View
1. .. ...-,;rr-::----- Septum
Vertical
, - - - - W a l l Involute
Chambers Evolute Equatorial Section
Marginal Cord Axial Section Protoconch

(b) ASS/LINA
Whorl Wall----.
Thick Due To
Flosculization
Tubular
Chamh\:r
Chmnberlct
Side View
Equatorial Section
Axial Section
External View
(c) .~LVEOLINA
FIG. 32 - 4 : LARGER FORAMINIFERA
PALAEONTOLocy
492
fine Granules - - - ,
Central Boss---
Peripheral Flangc·- ---1 Side View
Top View External View
Equatorial Chambers
Embryonic Apparatus--- Rectangular In Annular
Rin~
Embryonic
Apparatus (1+11)
Lateral Eulepidine Type
Equatorial Chambers
Chambers· -Layer
With Rectangular
Chambers Equatorial Section
(a) DISCOCYCL/NA
Peripheral Flange
Side View
Top View
External View
Equatorial Chambers ' Layer
Equatorial Chamber
With Eq uator1·a1 Chambers - Embryon ic Apparatus
~., -... . Arcuate To Rhombic
....
... " .....
~
. Embryonic ApparafUS
.... - -
- C r • i 'J " 11 lll
---
Nephrolcpidine Type
--.... - - Lateral Chambers
Pillar -
Vertical Section
Equatorial Section
(b) LEPIDOCYCL.INA
FIG. 32 - 5 : LARGER FORAMINIFERA
INTRODUCTION TO MJCROPALAEONTOLOG Y
Equatorial section : Embryonic apparatus bilocular prntoconch circular, attached with a

larger subcircular overlapping second chamber (dculeroconch) ofte n hy u Hmall Htalk
(eulepidine); equatorial plane may be horizontal or curved; cqualori:.il ·hamhcn,
rectangular,arranged in annular rings, sometimes irregularly overlapping ; c h,rn1 h ·r,
connected by annular stolon at the base; septa radial and strai ght; in case the cquat<1riul
plane curved irregular polygonal lateral chambers with traces of intersecting pillurs may
be visible.
Vertical section : Elongated biconvex in outline with equatorial chamber-plane al middle
and lateral chambers on either side; equatorial chamber-plane may be straight or curved
with rectangular equatorial chambers; height of chambers gradually increases towards
periphery; first chamber (protoconch) large, subcircular, completely covered by a larger
second chamber; lateral chambers in regular or irregular tiers, number of which decreases
towards periphery; chambers elongated rectangular in outline; pillars radially arranged
in between rows of lateral chambers, thicker toward the outer side.
Age : Palaeocene-Eocene
Lepidocyclina (Fig. 32-Sb)
External view : Test bilaterally symmetrical, thick/thin, lenticular, circular in outline with
a prominent central mamelon often surrounded by a thinner peripheral-flange which may
be broken in fossil; surface with fine granules and often with a central pustule.
Equatorial section : Circular in outline in ideal case; initial embryonic apparatus bilocular,
first chamber protoconch circular, partly or wholly enclosed by a larger subcircular second
chamber (eulepidine or nephrolepidine); chamber are thick-walled except the partition
between them; periembryonic or nepionic chambers (chambers surrounding embryonic
apparatus) arcuate, arranged in a ring; other equatorial chamber varying in shape from
ogival near centre to rhombic or hexagonal towards periphery but never rectangular; lateral
chambers (may be seen in case the equatorial plane curved), polygonal, thin-walled,
intercepted by pillars.
Vertical section : Biconvex in outline; equatorial chambers-layer at middle with lateral
chamber-layers on either side; first chamber circular, second one subcircular and other
equatorial chambers rectangular; lateral chambers rectangular, arranged in tiers; numerous
pillars present intercepting the tiers of lateral chambers.
Age : Oligocene-Miocene
B. Smaller foraminifcra (Microscopic description) (Fig. 32-6a-e)

Textularia
Test bilaterally symmetrical, broad at base, tapering towards apex, flattened conical,
elliptical in apertural ~iew; uncoiled; chambers alternately arranged in two rows (biserial);
ea~h chamber inflated with lobed margin; suture straight and depressed; aperture semilunar
lymg on the mid-inner margin of the last chamber; wall arenaceous with vitreous lustre.
Age : Carboniferous-Recent
L a4 ,,.t ·f h . i • .C a ;
___../
I
...... -- .... .....
"' l
494 PALAEONrolor.
,r
Aperture
Chambers
Biserial
Sucure Last Ch111tibcr
Al>Crture
AperturaJ View
Side View
(a) TEXTULARIA
_Aperture ---~~;;;;;~ Chamber In

Ventral View
Side View Ventral View

(h) ROTAL!A
Aperture Slit-Like
Apertural View
Side View Aperture With A

Tooth-Like Projection
(c) PENEROPL!S
Chambers S~~ng
Milioline Corhng
.
r--.~~~-- Aperture
~~~......_
(cl) TRJLOCULINA
Dorsal View Side View
(e) Gl0807'RUNCANA
FIG. 32 - 6: SMALLER FORAMfNJFERA
JNTRODUCTION TO MICROPALAEONTOLOGY 495
Rotalia
Test a. ymmetric, circular both in apical and basal view and nearly planoconvex in
apertural view; trochospirally coiled; evolute, chambers are rhombic in dorsal/apical view
but triangular in basal or apertural view; margin lobed; suture on dorsal side raised and
curved, but depressed and traight in basal view; umbonal plug prominent; aperture at
the base of last chamber, elliptical; wall calcareous, perforated .
Age : Cretaceous-Recent
Peneroplis
Test circular, compressed lenticular in side view; planispiral, involute to convolute;
chambers quadrate or curved elongated, triangular in outline; suture curved or straight,
depressed; margin rounded ; umbilicus shallow; aperture slit-like situated symmetrically
at the middle of last chamber; surface ornamented with fine ridges and furrows parallel
to coiling of test; wall calcareous, perforated.
Age : Eocene-Recent
Triloculina
Test triangular in apertural view, fusiform in side view; chambers coiled in three planes,
120° apart from each other (milioline type); chambers elongated, tubular, often ornamented
by transverse ridges; suture depressed, curved or straight; wall calcareous, imperforate;
aperture circular with a tooth-like structure at top of the last chamber.
Age : Jurassic-Recent
Globotruncana
Test vitreous, perforated (planktic); compressed trochoid; apex blunt, base flat; conispiral,
evolute; chamber subcircular; suture curved thick and ridged; margin lobed with irregular
marginal keel; aperture narrow umbilical, extending upto margin.
Age : Upper Cretaceous
32.5 PHYLUM : CNIDARIA

Class : Anthozoa
l. Habit (solitary/colonial); 2. Type of colony (for colonial coral); 3. Symmetry (for solitary
coral); 4. Shape of corallite or corallum; 5. Surface sculpture; 6. Septa and septa! features;
7. Axial structure; 8. Tabulae.
Order : Tabulata (Fig. 32-7a-c)
Halysites
Colonial, chain type of colony in top view, each chain bifurcating and joining repeatedly;
corallites tubular, elliptical in cross section; wall unfused; septa absent, spine-like false
septa may be found; tabulae concave upward; external surface of corallites with growth
lines.
Age : Ordovician-Silurian
Palac(Gco)WP-63
496
Fa\' sitt'S
. , ,· • cerioid· w·,11 among adjacent corallites unfused but perforated b t:
oloma 1. mnss1v • ' . . . . . . . Y11ne
•s· cc~r,llit ;>s polygonal m cross section, sltghtly elongated m side view· se
mura I pOr , • ., 1 • f h . • Pta
absent but spine- like pseudoseptn growing out from wal 1 o eac coralltte but not reaching
th, centre; tubulae horizontal.
Ag, : Ordovician- Devonian/Carboniferous
Syring / rn t
Colonial. fas iculated, phaceloid; adjacent corallites separated but nearly parallel and often
joint~d by latent! tubes; corallites tubular, circular in top view; septa absent but pseudosepta
may be present; tabulae horizontal or concave upward.
A~" : Ordovician-Carboniferous
Order : Rugosa (Fig. 32-8a-d)
alct·ola
Solitary, bilateral symmetrical; short, medium-sized, slip.per or sandle-shaped, one side
flat, other side convex; wall thick; calyx deep with several major and minor sepata, often
losed by an operculum showing internally a medium ridge and several other less
prominent lateral-ridges; surface with transverse growth lines.
Age : Silurian-Devonian
'Zaphrentis
Solitary. bilatemlly symmetrical, moderate to large size; elongated, conical or horn-shaped
(ceratoid); wall thick epitheca with growth lines; calyx deep with major and minor septa
and with a deep cardinal fossula; dissepiments poorly developed; tabulae convex upward.
Age : Silurian/Devonian-Carboniferous
Cystiphyl/um
Solitary; conical/cylindrical; wall thick, surface with growth lines; calyx shallow; septa
rudimentary; calyx filled up with numerous overlaping dissepiments; central parts filled
up by tabulae.
Age : Silurian-Devonian
Wagenopl,y/lumllonsdeleia
Colonial, usually fasciculated subparalled-branched; corallites conical, circular in outline;
~all t~in; septa numerous, major and minor, alternating; peripheral part with numer~us
d1ssep1ments, columella rod-like, associated with tabellae at centre giving an a~ial
structure; Tabulae convex upward; outer surface of corallites irregular with longitudinal
and transverse lines.
Age : Carboniferous-Permian
497
Wall Not fused
Massive Mural Pon:s
Honey-Comb
Colony ...___.Spine-Like
Pscudoscpta
Polygonal
Corallite
Colony (Magnified) Showing Horizontal & Vertical Scctioa
Colony
(a) FAVOSITES
r----Corallite
Elliptical In Outline ~~~~:a;;~~;L---i hain Colony
Bifurcating
Elongated
...______ Corallite In
Longitudinal Growth Rings
View
Colony
Colony (Magnified) With Transverse and Longitudinal Section
(b) HALYSITES
~ ~ ~ - - - Top View
OfColony
Longitudinal Section
Showing Tabulae
Colony
(c) SYRJNGOPORA
FIG. 32 - 7 : TABULATE CORALS
Calyx
Slipper shlpcd
Side View
Front View
(a) CALCEOLA
(b) ZAPHRENTIS
Corallites Circular
In Top View
alyx Filled
~~[1,i~~~:--- Up With-
Disscpimcnts
Dissepiment Radial
striations
Longitudinal Tahulae ---~o-\el!"""
Striations
''""_"_ _.....,,__Growth
Lines
Longitudinal-t---w.. ,
Section
,_.--Apex
(c) ·y~1·w11YI.I.UM
(d) IVA AGENOPH)'LLUM
FIG. 32 - 8 : RUGOSE (Tetracorals) CORALS
tNTRODVCT!ON TO MICROPALAEONTOLOGY 499
Septa
Exsert Septa
ut------Concentric
Growth Lines ~.....tF----"ynapticulated
Septum
(Magnified)
Radial Septal Groove
(a) MONTLJVALTIA
~ MajorSeptu
e----Minor Septum - - - - . . . - t i ' : : :

· Side View
·~.u.:J.:11.i.N=---- Continuation Of Exsert

Septa Toward Base Top View
- - - Synapticuk
Pore
Basal View
One Septum (Magnified)
(b) CYCLOLJTES
---Colony Massive
orallite Polygonal ·
Colony
Top View
Synapticulated Septa
(c) ISASTREA
FIG. 32 - 9 : HEXACORALS
PALAEONTOLOGY
500
_ _ _ _ Glabella
Anterior
---Glabella
--Facial Suture
,__---Caudal Spine
Dorsal View
(a) REDLICHIA AxiaJ Thorax
Anterior
Dorsal View
(b) PARADOX/DES
~ -·
Pleuron
labella
AxiaJ Thorax
CephaJon Eye
Facial Suture
l'ygidium [
Dorsal View
Plcuron
(c) PHACOPS
Axial 'fhora.t
Thorax
Dorsal View
(d) CALYMENi
FIG. 32 - JO: TRILOBITES
. ,,.....
/NTRODUCTION TO MICROPALAEONTOLOGY 501

Order : Hexacoralla (Fig. 32-9a-c)
Montlivaltia
- Solitary, radially symmetrical; turbinate, short conical; epitheca well-developed with
longitudinal septa} corrugations and some transverse growth lines; calyx shallow, septa
numerous, exsert, major and minor; septa} surface synapticulated; dissepiments abundant.
Age : Triassic-Cretaceous
Cyclolites
Solitary, radially symmetrical, hat-shaped; calyx convex upward; epithecal wall radially
corrugated; septa numerous, exsert, major and minor, trabeculated and fenestrate;
dissepiments numerous; no axial structure.
Age : Jurassic-Eocene
Isastrea
Colonial; massive, cerioid; wall unfused; calyx polygonal; six major septa and other minor,
synapticulated; no axial structure; dissepiments abundant.
Age : Jurassic-Cretaceous
32.6 PHYLUM : ARTHROPODA

Subphylum : '!!ilobitomorpha
Class : Trilobita (Fig. 32-1 Oa-d)
1. Symmetry and shape; 2. Features on cephalon : outline-glabella - facial suture - eyes -
genal angle - other features; 3. Thorax : number of segments - pleural extremity - other features;
4. Pygidium : shape, size relative to cephalon, number of segments, other features; 5. Surface
sculptures.
Order : Phacopida
Calymene
Bilateral symmetrical, subelliptical in outline; anterior and posterior broadly rounded; head
semicircular; genal angle rounded; glabella, inflated, broadest behind, partially segmented
with 3/4 lobes and furrows; eyes small, facial suture g_onatoparian; thorax with 12/13
segments; pleurae grooved, ;ngular and faceted; pygidium smalle; than head-shield,
subcircular with 6 to 11 segments· margin entire. • --"-le:--....,,,...._,,,.'--"_,,_
. . . ' 1\--UC...'10 f.id~
Age . Ordov1c1an-Silurian UO -
Phacops
Bilateral symmetrical, nearly elliptical in outline; anterior and posterior side broad and
rounded; cephalon large, semicircular; genal angle rounded, glabella inflated, anteriorly
broad and rounded • unsegmented; facial . suture propanan;· eyes very Iarg~compoun d·,
PALAEONTOLocy
502
------Umbo
- - - - Reticulate Structure
Node
a.+~~~y--Mediari Ventral
Sinus
Ventral View
Reticulale Structure
Anterior
Near Umbo
Side View
(a) PRODUCTUS
Spine Along Posterior
Beak-Ridge Umbo
Dclthyrium
Hinge Area Costae
Anterior
,I, Growth Line
Dorsal View Ventral View
B-Valve
P-Valvc
Side View
1h1 CIION/;"/'/-..\
FIG. 32 - 11: BRACHIOPODA
-
JN1'ROVVC'fl0N TO MICROPALAEONTOLOGY 503
thorax with eleven segments, margin rounded; pygidium smaller than head with 6-8
segments.
Age : Devonian
Order : Redlichida
,-
Redlichia
Bilateral symmetrical, oval in outline; anterior side broader than posterior; cephalon
semicircular, genal angle projected posteriorly in the form of two prominent genal spine§;,
glabella anteriorly narrow not reaching anterior margin, 3/4 segments, often partial;_eyes
large, cresentic, facial suture opisthopariar,1 thorax with 9/ 10 segments; pleurae grooved
w,tfispinose end, curved bacfiard; pygidium smaller than. head-shield with 7/8 segments,
margin spinose; first pygidial thoracic segment extends posteriously into a large caudal
spine.
Age : Cambrian
Paradoxides
Bilateral symmetrical; large, elongated, elliptical in outline, anteriorly broad, posteriorly
narrowed; head-shield semicircular with broad anterior outline; genal angles projected
into two long genal spines; glabella subglobular, broader anteriorly with 2 to 4 segments,
last one being the largest; facial suture opisthoparian; ~s,s moderaly larg~ arched; thorax
long with 16/20 segments, pleurae grooved and each produced into backwardly directed
spine; pygidium sm~ll (micropygu&), unsegmented or with very few segments (2 to 4)
each projected laterally into backwardly curved spine:
Age : Cambrian
32.7 PHYLUM : BRACHIOPODA

Scheme of description (morphology of external shell only)
l. Symmetry; 2. Thickness; 3. Shape (valve outline and sideview); 4. Dimension; 5. Features
of posterior side (umbo, hinge line, interarea, pedicle opening); 6. Anterior side; 7. Surface
sculpture. ·
Class : Articulata
Order : Strophomenida
Productus (Fig. 32-11 a)
Bilaterally symmetrical, shell thick, *L < B > T (transverse), planoconvex or concavo-
c~nvex,. semicircular in outline, hinge line straight, ~rophic, often olate: large ventral umbo
highly mcurved, (almost covering the ventral hinge area); anterior margin smooth; surface
ornamented with radial costae, intersected by concentric growth lines near the umbo
producing a typical reticulate structure; at points of intersection sometime develop spines
~nd/or tubercles. A prominent ventral sinus present. '
Age : Carboniferous-Permian
'
* L : Length; B : Breadth; T : Thickness; H : Hdght.
Palac(Gco)WP-64
\
l'/l / ,/\1\()(/ I ()I,( J( I
504
Anterior Margin---__;;:• Ventral Vit w

DorsaJ View lik VI w
(a) REFINESQUINA
Posterior
B-VaJvc---M ~ ·. J-- - -( :'II(},,,,.,

P-Valvc---1:-1~
~~~ i ,- - ..... < mwth Une
Knee Shaped-__,.,..
BendAt,c:!!IE~~
Anterior Side View Ventral View Anterior Margin
Margin
Anterior
(b) LEPTAENA
Posterior
-----Umbo-----
Hinge - - - - Dclthyrium I
Linc: J'-V,dve
.....all!~~
Capill11e
... -..--Rcliuplnatc In
DorsaJ View Side View
VcntraJ View
Side Vh.:w
Anterior
(c) STROPHOMENA
Posterior
Donal View
Side View Anterior
· (d) 0R7UIS
FIG. 32 - 12 : BRACHIOPODA
INTRODUCTION TO !t!_~Cf?.OPALAEONTOLOGY •
505
Chonetes (Fig. 32-1 J b)
Bilaterally symmetrical, shell thin L < B > T (t . . )·
I . h. 1. . ransverse , concavo-convex; rarely
p anoconvex, inge me straight (strophic) hinge area well de I d t · ·
t· I d Ith · I d ' ve ope , nangu 1ar with a
r~angu ar e ~rm~, c os~ ; beak _ridge (upper margin or hinge area) of ventr~I valve
with _a row of_ diverging spines that increase in length from umbo towards hinge margin·
anterior .margin ~mooth with a broad notch at the middle; surface with few concentri~
growth Imes, radial costae and often with granules and tubercles.
Age : Silurian-Permian
Rafinesquina (Fig. 32-12a)
Bilateral symmetrical, shell very thin, L < B > T; semicircular in outline, concavoconvex;
umbo upright; hinge line straight (strophic); hinge area small but well-developed;
delthyrium triangular closed; anterior margin smooth, surface with numerous radial
capillae and few concentric growth lines.
~ F i g . 32-I2b)
Similar to Rafinesquina, but near anterior margin the shell shows an abrupt knee-shaped
bending often at right angle to posterior part.
Age : Ordovician-Devonian/Carboniferous
Strophomena (Fig. 32- J2c)
Similar to Rafinesquina, but the shell is resupinate (s-shaped), ventral valve convex near
umbo and concave towards anterior (i.e. valve towards anterior convexiconcave but
concavoconvex towards posterior).
Order : Orthidina
Orthis (Fig. 32- I 2d)
Bilateral symmetrical; shell thick, L = B > T; subcircular, planoconvex to biconvex; hinge
line curved, short; umbo of ventral valve prominent, elevated; surface ornamented with
prominent costae.
Age : Ordovician
Order : Spiriferida
Spirifer (Fig. 32- J3a)
. Bilateral symmetrical; shell thick, L < B > T; traingular in outline; biconvex, strophic,
o~ten alate; hinge area well developed, often transversely striated, with a partly closed
tn~ngular delthyrium; umbo of both the valve incurved; anterior margin crenulated,
~niplicated; surface ornamented with prominent radial costae with few transverse growth
Imes; a prominent ventral sinus and a corresponding dorsal ridge present.
Age : Silurian-Permian
' ' ...
> 1
506 PALAEONTOLOGY
Posterior
1linac Arcll - - - - - , ~~- - - --~~ u,nbo~-------~-, Faint Growth Linc
- - - IJclthyrium
Hinge \.Inc----,
Atill~"l"I
Costac
Median Dorsal Ridge
Anterior
Anterior Margin
Ventral View
+4--B-Valve
Dorsal View
4-4-.......--- P-Valve
Side View
(a) SP/RIFER
Hinge Arca Posterior Umbo----,
Coslac----~~
Anterior Margin _ _ _ _ _.,....__._
___ Median Ventral
Median Dorsal ---"lie:ir,i 1
Ventral View Sulcus
Ridge Dorsal View
13- Valves
Side View
(b) SYRJNGOTHYRIS
Umbo - - - - - - ,
Posterior
Side View
Anterior
(c) STREPTORHYNCHUS
FIG. 32 - 13 : BRACHIOPODA
..
JNTRODVCTJON TO MICROPALAEONTOLOGY
507
Spiriferina
Similar to Spirifer, but with an internal median septum in ventral valv d h
e an punctate s e11 -
wa It .
Age : Triassic-Jurassic
Syringothyris (Fig. 32-13b)
Simil~r to Spirife~, but ventral valve with a large traingular hinge area (almost equal to
the width of brachtal valve) bearing a prominent, partly closed large.triangular delthyrium.
Age : Devonian-Carboniferous
Streptorhynchus (Fig. 32-13c) .
Bilateral symmetrical; thick, L = B = T; semi-oval in outline; dorsal convex, ventral
convex near umbo but almost plain towards anterior; ventral umbo prominent, outwardly
projected; dorsal umbo incurved; ventral hinge area large with a triangular delthyrium
bearing a semi-oval pedicle opening at the base; surface ornamented with strong costae
and fine radial lines, anterior margin plicated.
Age : Permian
Atrypa (Fig. 32-14a)
Bilateral symmetrical; thick, L = B > T; semi-oyal in outline, convexiplane or ventral
valve slightly convex near the umbo, dorsal much inflated; Hinge line long, slightly curved
near margin or nearly straight, sometimes alate; hinge area almost absent; ventral umbo
small, upright with a small partly- closed dethyrium bearing a foramen at the apex; surface
ornamented with concentric growth lines and fine radial ridges; a poorly developed ventral
median sinus and a corresponding fold .may be present; anterior margin nearly entire,
often projected at its middle as a lip.
Age : Silurian-Devonian/friassic
Arthyris (Fig. 32- l 4b) f· · .,.,.
Bilateral symmetrical, thick, L = B > T, rever~ ly oval i~ outli~e, biconve~; hinge line
curved, hinge area short; ventral umbo small, m rved with a circular ped1cle opening;
hinge area poorly developed; surface with numerous concentric growth lines; a median
ventral shallow sinus and a small dorsal fold present; anterior margin entire.
Age : Devonian-Permian
Order : Rhynchonellida
Rhynchonella .(Fig. 32- l4c)
B_ilateral sym'metrical thick L = B < T; rhombic in outline, biconvex (dorsal more convex),
htnge line short and curved; hinge area poorly developed; umbo small, sharp, slightly
~ur~ed with a foramen surrounded by a deltidium; surface ornamented with prominent
radial costae; a large median sinus in the ventral valve which is broad and deep in front
508 PALAEONTOLOr,y ·
Posterior
Capillae - - -
Dorsal View
Anterior Ventral View
Side View
(a) ATRYPA
Posterior
P-------- U m b o - - - - - Median Ventral Sulcus
Foramen
8-Valvc
4--1~-P-Valvc
Concentric
Growth Line Ventral View
Dorsal View Side View
Anterior
(b) ATHYRJS
Umbo----.1_ Umbo
Hinge Line Pedicle Opening
Median _ _ _ ___...,..
Dorsal Ridge
Anterior Margin
MuJtiplicated -----Dorsal View Anterior View
(c) RHYNCHONE!sLA
Hinge Line Ventral
Curved Umbo With
Ci•ular
Formen
~....,.~._-P-VaJve
P-Valve
.,~--B-Valve _ _ _ P-Umbo Strongly Incurvcd Umbo
Side View
Biplicated Dorsal View
Anterior 8-Valve
Margin (d) TEREBRATULA
Anterior
(e) PENTAMERUS
FIG. 32 • 14 : BRACHIOPODA
·' .. ' . ...
._,_
tNTRODUCT!ON TO MICROPALAEONTOLOGY 509
where it pro~uces a .to~gue-shaped extension; dorsal valve with a corresponding fold;
anterior rnargm rnult1phcated with prominent W-shaped plications.
Age : Devonian-Cretaceous
Order : Tcrebratulida
Terebratula (Fig. 32- l 4d)
Bilateral symmetrical; thick, shell longitudinally oval in outline (L > B > T); valves
equally biconvex; hinge line short and curved with poorly developed hinge area; umbo
of ventral valve prominent, upright, with a large circular foramen (often with a basal
closed delthyrium), surface ornamented with few concentric growth lines; anterior side
often with two folds on dorsal valve and two corresponding sinuses on the ventral valve
producing a biplicated anterior margin.
Age : Triassic-Miocene/Pliocene
Order : Pentamerida
Pentamerus (32-14e)
Bilateral symmetrical; shell thick oval or nearly pentagonal in outline (L > B > T);
biconvex (ventral more convex); hinge line curved, short; ventral area narrow; umbo of
both valve strongly incurved often touching each other; delthyrium present; surface smooth
or with a few concentric growth lines, anterior margin smooth.
Age : Silurian
32.8. PHYLUM: MOLLUSCA

Class : Pelecypoda (Bivalvia)
Scheme of description (Description of single valve where both are identical)
A. External features :
J. Thickness; 2. Shape; 3. Degree of inequilaterality 4. Umbo; 5. Hinge line; 6. Hinge
area; 7. Ventral margin; 8. External surface ornamentation and other features.
B. Internal features :
9. Teeth; JO. Ligament; 11. Adductor muscle scars; 12. Pallial line.
Order : Nuculoida
Nucula (Fig. 32-15b)
Valve thin, (L = H); triangular or oval in outline, inequilateral; posterior side smaller;
umbo prominent, beak opisthogyral; hinge line curved, hinge area poorly developed;
ventral margin smooth; surface nearly smooth or with numerous fine concentric lines.
Hinge pl~te wel_l developed, curved with taxodont dentition; a ligamental pit below the
umbo; D1mayanan muscle scar nearly equal·• pallial line ..s,·mp Ie: ·111 terna IIy nacreous.
Age : Cretaceous- Recent
PALAEONTOLOGY
510
Dorsal
Umbo-------:*lm,r...._
Hinge Linc
Taxodont--~~
Anterior--.+--.-.
Adductor
Muscle Scar Ventral Margin
Ventral
External View
Internal View (a) ARCA
Dorsal
Beak------
Umbo----___;a
l '," 1,·11, 1 1 ,\dd1i-·1i,r

\lu, d ..: Srar
Palhal Lmc----.:1~..;::..-..._ _.,.. Ventral
Internal View External View
(b) NUCULA
Dorsal
~-----Umbo-------.
---Growth Line
Anterior Adductor _ _ _.__. Ventral

Muscle Scar _ / Margin
Ventral
Internal View External View

(c) GLYCIMERIS
Dorsal
Growth Linc----, -----Umbo------. ---Beak
----Teeth (Schizodont)
Hinge Line
A-,.___ Posterior Adductor

Muscle Scar
Ventral
.___ Pallial Linc
External View
Internal View
(d) UN/0
---Beak
Ventral Mlrain---..:l -.--Pallial Line

External View Internal View
(e) TRIGONIA
FIG. 32 - 15 : PELECYPODA (Bivalvia)
!NTRODUCIION TO MlCROPAl.AEONTOLOGY
Order : Arcoida 511
Arca (Fig. 32-l 5a)

Valve thick, L > H; subquadran 1
I b b . gu ar, moderately or t I .
~ger, um o road, mcurved with proso rat . .s ron~ y mequilateral; posterior side
hmge area often with straight or .
rmes, mtersected
.
g~ beak, hmge lme straight with well devel d
hinge margin to bel~w the umbo~nverte~• V-sha~ed ligamental grooves converging
' ventr"' margm crenul ated' su rface with coarse radial
by fine concentric striae.
~:m
Hinge plate relatively wide and stout b .
(taxodonta) anterior adduct I earmg numerous serreted teeth and sockets
. . ' or muse e scar round d · ·
isomayanan; pallial line simple. e • posterior elongated nearly
Age : Triassic-Recent
Glycim.eris (Fig. 32-15c)
Valve
. thick, L -- H·, suborb'1cuIar ·m out1.me;· nearly equilateral; umbo nearly central slightly
t~curv~d, ~eak orthogyral; hinge· area smalJ, triangular, often with inverted V-shaped
divergmg hgamental grooves; hinge line curved; surface smooth or ornamented with
numerous radial striations and concentric growth lines; ventral margin smooth;
Cardinal plate well developed curved with taxodont dentition; adductors impression
subequal, anterior one subtraingular, posterior oval; paJlial line simple.
Order : Trigonoida
Trigonia (Fig. 327 15e)
Valve thick, L > H; subtriangular in outline, strongly inequilateral, posterior much larger;
anterior rounded, posterior part produced and angular with a ridge from umbo to the
posterior~ventral . border; umbo anteriorly placed but the beak opisthogyral; hinge line
curved with narrow hinge area; surface with strong concentric growth ribs but posterior
ridged-off portion with different ornamentation.
Hinge plate with two grooved teeth (schizodont) in right valve diverged from below the
umbo, and one v-shaped tooth in left valve; adductor impressions deep, anisomayarian
(anterior smaller, placed near the umbo) pallial line simple.
Order : Unionoida
Unio (Fig. 32-15d)
Valve thin, longitudinally oval in outline with a thin outer periostracum, L > H; anterior
and posterior margin rounded; strongly inequilateral, anterior smaller than posterior; umbo
broad, anterior; beak prosogyral; hinge line slightly curved; hinge area not developed;
external surface with concentric growth lines, ventral margin smooth.
Hinge plate poorly developed; right valve with one small anterior lateral and one long
posterior lateral and left valve with two anterior and two posterior laterals, teeth bifurcated;
512 PALAEONTOLOr,y
IJorsul
Umbc, - -- -.,..,ta
llingc I.inc
llc11k - --.....J11111 Umbo
I klnt1ifPIII frl'lh
1\111cn11r AJJuc111r - --..+~I Growth

Muscle Scttr Line
lnlcnud View
(a)CARDl7'A External View
Dorsal
,___ _ Radial Costae
Bcak~---,.,::;ii~~~~
He1erodon1---~""?1-~lt'-.:~'l)i
Anterior Adductor _ _...._

Muscle Scar
Ventral Margin _ _ ___;.,,,_.

-..ww~-
1nt em aI View
External View
(b) CARD/UM
External Ligarnental Groove -----1 Dorsal
Posterior Adductor--..._.,__,.,
Muscle Scar
Pallial Linc·----'
lntemaJ View External View
(c) CYRENA
Dorsal
--~---Umbo---A~~.a~-.::--- Ligamcntal oroove
Beak---iff-11:..v.
Hcterdont - - ~ • : . - " '
Posteri<'
-Adductor
.L.J.....
MuscfeSc,I
£,..-----r-Pallial SinUS
Ventral
Margin
Ventral
ExtemaJ View lntcmaJ View
(d) VF.NUS
FIG. 32 - 16 ; PELECYPODA
JNTRODVCTION TO MICROPALAEONTOLOGY
513
anisomaryarian, anterior muscle scar rounded, deep; posterior shallow and elongated;
pallial line simple, interior nacreous.
Order : Veneroida
<;ardita (Fig. 32- l 6a)
Valve thick; oval in outline; strongly inequilateral (posterior side larger), L = H; anterior
and posterior margin rounded; umbo prominent, incurved, anteriorly placed, beak
prosogyral; hinge line curved with poorly developed hinge area, surface with prominent
radial ribs, ventral margin crenulated.
Hinge plate not well developed, each valve with two long laterally placed cardinals,
parallel to dorsal margin and a very small anterior cardinal tooth; adductor impressions
anisomayarian, posterior larger; pallial line simple.
Age : Palaeocene-Recent
Cardium (Fig. 32-16b)
Valve thick; L = H; cordate or ovate in outline, slightly inequilateral (posterior larger);
umbo broad, incurved, beak slightly prosogyral; hinge line short, curved; hinge area
rudimentory; external surface ornamented with strong radial costae producing a highly
corrugated ventral margin.
Hinge plate poorly developed with imperfect cardinal and lateral teeth, right valve with
one/two cardinals and two anterior and two posterior laterals, left valve with one cardinal
and one anterior and one posterior lateral tooth; adductor muscle scars shallow,
isomayarian; pallial line simple.
Cyrena (Fig. 32-16c)
Valve thick; L = H; oval or cordate in outline, inequilateral (posterior larger); anterior
and posterior margin rounded; umbo broad, slightly incurved; beak prosogyral; hinge area
divisible into anterior heart shaped lunule and posterior elongated escutcheon; ligament
external posterioraly placed; surface with concentric growth lines, ventral margin entire.
Hinge plate thick with well developed heterodont teeth, right valved with two cardinals,
two anterior and two posterior laterals and left valve with three cardinal one anterior and
one posterior lateral; anisomayarian, posterior scar -larger; pallial line with a small sinus
at posterior; interior nacreous.
Venus (Fig. 32- l 6d)
~alve t~ick; oval in outline, L = H; shell strongly inequilateral (posterior larger) umbo
road, mcurved, beak prosogyral; ligament external, posterior; hinge line curved short
~ I
514 PA LAEONTOLOG 1.
Dorsal ----...-----Linear Teeth
Umbo----...
Hinge Linc ~il::~ftl;r===::.r~-- Ear Like Projection
L--li,l~----Resilifer
M.----4-~---Posterior Adductor
Muscle Scar
A-~---Pallial Line
,.,,.__ _ _ ventral Margin

(a) PECTEN Dorsal
Internal Li11,amental11-;~~:-=;;,;.--
Pit (Resilifer) r.
Hinge Line
lsodont Tooth-~~--
Posterior Adducto::::r~-.IIJ>-+-111'1r
Muscle Scar
Dorsal
Ventral Margin
Internal View
~.._-Posterior (b)
Fine Concentric
Adductor Dorsal Growth Linc
Muscle Scar
Beale

Ventral
Posterior_+---..i
Adductor
l l' ) OSFRl:R:\ Muscle Scar
External View
-r---Lcft Valve
Thick Larger
Jncurvcd---¥.RU
' Postaiorty Highly Incurvcd
Umbo Right Valve
Twisted Flat Smallcr
Umbo
Growth Linc
Left Valve Lid-Like

Unequal Valves
Internal View
External View
(c) EXOGYRA
Longitudinal
Lobes
Right Valve------
Costae---~
Apcx------
Extcmal View
(t) HJPPURJTES
FIG. 32 - 17 : PELECYPODA
515
with. distinct lunule and escutcheon , surface with concentric growth 1·mes; ventra1 margin
·
entire.
Hinge plate "':ell ~evelope~, wide; each valve with three thick cardinal teeth only; adductor
muscle s~ars amsoma~ana.n, posterior larger and elongated; pallial line with a sharp
angular smus at postenor.
Age : Oligocene-Recent
Order : Pterinoida
Spondylus (Fig. 32-1 ?b)
Shell inequivalved, right valve larger and more convex; valve thick/thin, oval or irregular
in outliner; L = H; i.lmbo of right valve broad, beak orthogyral; hinge line straight often
with ears, right valve with a triangular cardinal area; surface irregular, foliaceous often
with ribs, growth lines and spines of variable size.
Hinge plate imperfect, left valve and right valve with two cardinal teeth on either side
of a ligamental pit; mono~ayarian muscle scar, subcentral, (lying towards posterior).
Pecten (Fig. 32-17a) ·
Valve thin, ovate, L = H; almost equilateral; right valve more convex; umbo sharp, beak
orthogyral; hinge line straight extending on either side in the for~ of two ~qual or unequal
ears (alate); an anteriorly placed notch below the anterior ear (byssal notch); surface with
strong plications producing a crenulated ventral margin; Hinge plate poorly developed
with two linear teeth horizontally diverged along each ear on either side of a triangular
ligamental pit below the umbo; adductor muscle scar single, slightly towards posterior
(monomayarian).
Age : Carboniferous-Recent
Ostrea (Fig. 32-17 c)
Shell inequivalved, left valve larger, irregularly convex outer side, right valve, smaller,
flat or concave, valve thick, slightly/strongly inequilateral, posterior larger or smaller,
umbo rounded, upright, sometimes prosogyral, sometimes opisthogyral; hinge area poorly
developed or absent; surface of left valve foliaceous, very much irregular and uneven,
right valve less irregular with concentric growth lines.
Hinge. plate nearly absent, with~ut_. an~ teeth, ligamental pit broad, elongated, shallow
below the left valve umbo; no palhal lme; monomayarian scar posteriorly placed.
Age : Triassic/Cretaceous-Recent
Gryphaea (Fig. 32-1 ?d)
Shell inequivalved,. left valve l~rger and outwardly more convex, right valve flattened or
concave; surface with c~ncentnc growth lines, valve oval in outline L < H, ine uilateral;
umbo of left valve prominent and po· t d 1 · ·q ·
me , ~~rong y mcurved and twisted anteriorly; hmge
PALAEONTOLOGY
516
----Peripheral
Slit Band
t:t-~!ir-71--lnhalant Siphon
~ . - - Umbilicus
~ ~ r - - Peripheral
Slit Band
Peripheral View Apertural View
(a) BELLEROPHON
I
I
Seire I
,...
. I I
1 - ~ - Spiral And Spire :
Axial Lines , Spirals
. Body
,- mr-,.--- Spiral Granules
Wbor1 Body;
Whor1,•a-~
',_
(c) TROCHUS
(b) TURBO
Spire:-
'-
r-
1 Spirals
I
Body I
Wbort I ,-
I '
I
,_
I
Body
I
I
Whorl I
I
I
(d) NERITA ,_
I
(e) NAT/CA
FIG. 32 - 18 : GASTROPODA
I
JNTRODVCT!ON TO MICROPALAEONTOLOGY 517
area of left valve well developed, triangular with a shallow triangular ligamental groove;
Hinge line ~urved.' no hinge plate; teeth absent, adductor muscle scar single, deep
(monomayanan) slightly towards posterior; ventral margin smooth.
Exogyra (Fig. 32-1 ?e)
Similar to Gryphaea, but umbo of left valve is strongly incurved, more or less spirally
directing posteriorly.
Order : Hippuroida
Hippurites (Fig. 32-1 ?f)
Shell inequivalved, right valve (lower) larger, conical or subcylindrical in shape, left valve
(upper) small and flattened, . or slightly concave occurring as a lid or operculum on larger
valve.
Larger valve has a narrow apex; surface striated with three to four parallel longitudinal
lobes and furrows from apex to cardinal margin; hinge with two cardin~l teeth; anterior
adductor impression large, posterior one depressed.
Smaller valve with a central umbo and two prominent teeth, posterior one smaller.
Age : Cretaceous
Class : Gastropoda
Scheme of description :
I. Symmetry; 2. · Shape; 3. Type of coiling; 4. Length-Width ratio; 5. Spire and spiral
angle; 6. Whorl prophile; 7. Spiral suture; 8. Body whorl; 9. Umbilicus/Columella;
10. Aperature; 11. Ornamental features.
Subclass : Prosobranchia
Order : Archaeogastropoda
Bellerophon (Fig. 32-1 Sa)
Bilateral symmetrical, shell thick/thin globular, planispirally coiled, convolute: umbilicus
small, aperture subcircular or cresentic with a deep median peripheral slit-band; surface
smooth or with some growth lines.
Age : Ordovician-Triassic
Turbo (Fig. 32-1 Sb).
~h~e~I ~s~mmetric, thick/thin, turbinate L > W, conispiral, involute; spire slightly smaller
imperfo y Whorl, spiral angle acute; whorl inflated, spiral suture depressed; shell
orated or f . . . .
(holostom per orated; aperture dextral , large, circular; apertual face entire
atous); surface smooth or with spiral and/or axial lines. ·
Age : Jurassic R
· - ecent
/
518 PALAEONTOLOGy
,- r
'II I
I
I I
I I
I Spire :
Spirally Arranged Nodes
Spire I
I I
I
r
k
I
,_
r
-~---
I ..._._ • .,
..-
r----Posterior Caial
Body I
Whorl I
I
L
- - - - - - Anterior Canal
(a) TURRITELLA (b) CERITHIUM
Spire _ _...
Concealed Posterior Canal
Aperture
i
I
I
I
I
Spire I
I
Anterior Canal
,-~~
I
Dorsal View Apcrtural View

(c) CYPRAEA. Aperture
I
I
r I
Spire I I
r Bodyl
,-
J..
1
Whorl I
I
I
/ I
Body I I
Whorl I
I I
I
r 'I,_
I
I (c) FUSUS
I
L..
- - - - - Small Anterior Canal
(d)CONUS
FIG. 32-19: GASTROPODA
., _.,.:>
JNTRODUCTION TO Ml 'ROPI\LAFON'l'OI.OG Y
I <J
Trochus (Fig . 32- l 8c)

Shell asymmetric, thick, troc hif'or111/co11i ·ul ; L c: W; ·011i spiral, invol111c; Nf) ir · harp; eq ual/
larger than body whorl ; spirul an il c ac111 ·; whuf'I profile flull cn ·d with flu shed pi nt!
sutures; body whorl with flat base; apcl'lurc d •xtral (holostomatous), ap ·rtural fac:c ent ire,
rhombic in outline, she ll impcrforat e; surface with spiral lin ·s, often wirh small r,1n ulc .
interior nacreou s.
Age : Silurian- Recent
Order : Mcsogastropodn
Nerita (Fig. 32-1 Bd)
Shell asymmetric, thick, ovoid, L = W; conispiral, nearly convolute; spire almost nat ,
much smaller than body whorl, with very few whorl s, spiral angle obtuse, whorls nattcned,
spiral sutures flushed with the surface; aperture dextral, semicircular, entire, holostomatous,
outer lip thick, inner lip with tooth-like structure; shell imperforatc, surface smooth or
with spiral ribs.
Natica (Fig. 32-1 Be)
Shell asymmetric; thin, oval or globular (L = W); consipiral, nearly convolute with few
whorls; spire much shorter than body whorl, spiral angle obtuse; whorls inflated with
depressed spiral suture; aperture dextral, semilunar or oval in outline, entire, holosto-
matous, outer lip sharp. inner lip often thickened by callus; shell perforated with di stinct
umbilicus, surface with fine spiral and axial lin·e s or may be smooth.
Age : Eocene-Recent
Turritella (Fig. 32- I 9a)
Shell asymmetric, thin, turreted L > W, conispiral; evolute; spire much larger than body
whorl with many spiral whorls which are flat or inflated; spiral angle acute, spiral suture
depressed and dipping; aperture dextral, oval, entire, holostom~uter lip thin; shell
imperforate, surface with fine spiral lines. ·
Cerithium (Fig. 32-l 9b)
Shell asymmetric, thick, turreted, L > W, conispirnl, evolute; spire much larger than body
whorls with many flattened spiral whorls; spiral angle acute, spiral suture depressed;
aperture dextral, rhombic; outer lip thick, often reflected; apertural face with a short
posterior and a recurved anterior canal, siphonostomatous; shell imperforate, surface
ornamented with spiral ribs and sometimes with spirally arranged nodes. ·
Age : Cretaceous- · Recent
Cypraea (Fig. 32- I9c)
Shell asymmetr' .
spire .1 •c, thin, obconical, L'> W, flattened apertural surface, conispiral convolute;
. a most conceal d b h
Pala '(G e Y t e last whorl; body whorl inflated; aperture dextral, narrow,
c co)WP-66
/
PALAEONTOLOGY
520
_ _ _ Spiral Linc
i
l
Spire~
I
~
I
I
IJ'-'~~~Apertural Face
I Crcnulatcd/Spinose
1
Body I I
Whort I Spire I
I
l
l
'-
n
Posterior Canal
Long Anterior
l Canal
I ..+--SpiraJs And
I Axials
I ._,__ _ Aperture
(a) MUREX
I
Boby I
t- Whorl I
i I
. I I
SplJ'C I
I
',_ I
I
i I
I I
Body l '-
Whorl' (b) VOLUTA.
'
(c) PHYSA.
~'/i1.)w~_ Apcrturc Closed

By lncurvcd Lobes
Median Groove
On Face
Transverse Ridges
(c) TENTA.CULITES
(d) CONULA.RIA.
FIG. 32 - 20 : GASTROPODA
521
slit-like along the entire length of last whorl· inner and t ,. · tl
. , ou er 1p m ected· apertural fac
dentate or crenulated with short canals at each end ( · h ' . e
..& h d sip onostomatou ); shell imperforated ·
su1 .ace smoot an nacreous.
Order : Neogastropoda
Conus (Fig. 32- l 9d)
Shell asymmetric, thick, obconical, L > W; conispiral , almost con olute as last whorl
conceals most of the other whorls; spire much smaller than body whorl , spiral whorl s
flushed with surface, spiral angle obtuse; aperture dextral , long slit like along the , hole
length of last whorl with a short truncated anterior canal (siphonostomatous); shell
imperforate, surface smooth or with fine spiral and/or axial lines.
Fusus (Fig. 32-19e)
Shell asymmetric, thick, fusiform, L > W; conispiral, im olute; spire almost equal to body
whorl; spiral angle acute; whorls inflated; spiral suture depressed , dipping; aperture
dextral, posteriorly oval and anteriorly projected into a long narrow canal (siphonosto-
matous); outer lip thin, inner lip often with callus; shell imperf'orate, surface ornamented
with fine spiral lines often intercepted by granules or tubercles.
Murex (Fig. 32-20a)
Shell asymmetric, thin, fusiform; L > W; conispiral, involute, spire and body whorl nearly
equal; spire sharp with acute spiral angle, spiral whorls inflated often divisible into ramp
and shelf; spiral suture depressed; aperture dextral, oval towards posterior but with a long
narrow canal towards anterior (siphonostomatous); outer lip thick, spiny, inner lip smooth;
shell imperforate; surface with spiral lines and tubercl~s; each whorl. bear nearly three
major varises each of which bear rows of spines that are increasing in size from apex. to
the middle of shell and again decreasing in size towards the base.
Age : Eocene-Recent
Voluta (Fig. 32-20b)
Shell asymmetric; thick L > W; volutiform (fusif<:>rm); conispiral , involute, spire smaller
than body whorl; spiral angle acute; spiral whorl with angular profiles, divisible into ramp
and shelf; spiral suture depressed and dipping; aperture dextral, elongated, narrow, wider
posteriorly and with an anterior deeply notched canal and a small posterior canal
(siphonostomatous); outer lip thick, inner lip with thick callus; shell imperforate, columella
with obli_que folds; surface with fine radial and spiral lines, rows of spirally arranged
tubercles or nodes along the peripheral zone of spiral whorls (junction of ramp and shelt).
Age : Eocene-Recent
PALAEONTOLOGY
522
- - - - Siphunclc Opening
~---Aperture
~,es:~~~~-- Suture Nautilitic
111--.,._____,,___ Saddle
~~-=~t-- Septum
M--""*l~-Scptlal Neck
Rctrosiphonate Equatorial View
A.C:lllii?it--- Aperture
Internal Features =~--Central Siphunclc
External View
(a) ORTHOCERAS
AperturaJ View
(b) NAUTILUS
-----Goniatitic
Suture
r----Saddlc
Equatorial View Apcrtural View
Lobe
(c) GONIATITES
t----Apcrturc---,1
~-+--Suture Ccratitic
Ribs
AperturaJ View
Equatorial View
(d) CERA.TITES
FIG. 32 - 21 : CEPHALOPODA
tNTR I)
I NT, Ml N l'\l \EONTOl O ,>~
521
Subclass : Pulmonatu
Phys 1 (Fi\! . . ---0 ·)
Sh ·II asymm tric. thin/thick. L W; fusiform; onispirnl involute; spire smaller than body
whorl; spire sharp \\ ith a f •w wh rls flush ·d with smfncc; spiral angle acute; aperture
sinistral. elongated o al, ap rturnl fo ·c •11tirc (holostomatous); shell imperforated or
perforat •d; surface smooth.
Age : Jurassic- R "'cent
Group : Ptcropoda
Co1111/nria (Fig. 3_-2Qd)
Shell thin, chitinous, conical or pyramidal with four sides; each angle of pyramid with a
straight groove; each of lateral faces may have a median longitudinal groove; surface
smooth or ornamented with numerous transverse parallel ridges and sometimes with tine
longitudinal ridges; aperture rhombic, triangular or subcircular, often closed by incurved
triangular lobes.
Age : Ordovician-Jurassic
Te111aculites (Fig. 32-20e)
Shell thick, solid, elongated conical; straight or slightly curved, apical part with septa
and basal part often with a vascular enlargement; surface with prominent transverse and
parallel rings and with longitudinal striae.
Age : Ordivician-Devonian
Class : Cephalopoda
I. Symmetry; 2. Size; 3. Shape; 4. Periphery; 5. Coiling; 6. Umbilicus; 7. Septum;
--·----
8. Siphuncle; 9. Suture; IO. Aperture; 11. Surface sculpture.
Subclass : Orthoceratoidea
· Orthoceras (Fig. 32-21 a)
Shell radially symmetrical, small elongated, conical, circular in outline; uncoiled; septa
concave upward or hori:rnntal; suture lines straight, aperture circular in outline; siphuncle
central; septa) neck retrosiphonate surface usually smooth.
Age : Ordovician
Subclass : Nautiloidca
Nautilus (Fig. 32-21 b)
Shel! b!laterally symmetrical, thick/~l~in, more or less globose, periphery rounded; coiling ,,.
planisp1ral, almost conv~lute; umbilicus very small; septa curved backward; siphuncle
central; septal neck proJected backward (retrosiphonate): suture nautilitic (lobes and I
sanddl~s rounded); aperture simple subcircular in outline with an external median
truncation (hyponomic sinus); surface smooth or with Very fine radial striae. .,
Age : Triassic- Recent
'...
PALA EONTOLOGY
51~
Suture Ammonitic
Suture,- - - ~ ~
Ammonitic
"""IIICi;~=;g;;lii_
Je:--peripheral Apcrtural View
Equatorial View Ropy Keel
(a) AMALTHEUS
~~~~-Costae
Bifurcating
Near Periphery
Costac Bifurcating Near Umbilicus Equatorial View
Apertural View
(b) PERISPHINCTES
Apcrtural View
Equatorial View
(c) MACROC EPHALITES
Very Coarse Rib Bifurcating At Periphery
Rectangular--
Aperture
Tubercle
Ribs Bifurcating,...-::.......11
AtPeripheny ~~~,
Ammonitic Suture
I' Apcrtural View
Equatorial Vicw
(d) ACANTHOCERAS
JNTRODUCTION TO MICROPALAEONTOLOGY
525
Subclass : Ammonoidea
Goniatites (Fig. 32-2Ic)
Shell bilaterally symmetrical, thick, globose, margin rounded, planispiral, involute to

nearly convolute; umbilicus small; septa curved backward, septa! neck projected in both
direction; suture with angular lobes and saddles (goniatitic); aperture subcircular; surface
usually smooth or with fine radial straie.
Age : Carboniferous
Ceratites (Fig. 32-2 ld)
Shell bilateral symmetrical, thick, discoidal; external margin rounded or actuely rounded;
planispiral, involute; umbilicus moderately large and shallow; body chamber small; suture
with rounded saddles -and crenulated lobes (ceratitic suture); surface with coarse ribs often
bifurc~ting, rarely tuberculated near the periphery.
Age : Triassic
Amaltheus (Fig. 32-22a)
Shell bilateralJy symmetrical, thin, discoidal, much flattened, periphery angular with a
ropy keel; planispiral, involute, umbilicus shallow and small; aperture subelliptical,
compressed; suture ammonitic with both lobes and saddles much divided; surface with
radial ribs bifurcating near umbilicus, sometimes bearing spines.
Age : Jurassic
Perisphinctes (Fig. 32-22b)
Shell bilaterally symmetrical, thick discoidal with rounded periphery; planispiral, advolute;
umbilicus shallow and wide; suture ammonitic; aperture subelliptical; surface ornamented
with radial costae each bifurcating near the periphery.
Age : Jurassic
Macrocephalites (Fig. 32-22c)
Shell bilaterally symmetrical, thick, globose or inflated; planispiral, nearly convolute;
rounded margin; umbilicus small but deep; suture ammonitic, aperture, semicircular, often
with two lateral projections; surface with radiating ribs, bifurcating near the umbilicus.
Age : Jurassic
Acanthoceras (Fig. 32-22d)
Shell bilaterally symmetrical, usually large, thick, discoidal; peripheral margin flattened;
planispiral, involute; umbilicus large and deep; suture ammonitic; surface ornamental with
very coarse radial ribs which bifurcate near periphery bearing a tubercle at each point of
bifurcation.
PALAEONTOLOGY
526
----Aperture
Radial Striations
Bifurcating
Equatorial View
(a) SCAPHITES Side View
(b) B.4.CULITES
(c) TURRILITES
li---i~-Soft Phragmoconc
(Usually Not Preserved)
Section A cross
Guard
Ciuard
(d) BELEMNITES
'Cr
tNTRODUCTION TO MICROPALAEONT,
., ..... "°_' • \ • ' ' .
-
~ J
·)I·,
OLOGY
Scaphites (Fig. 32-23a) 527
Shell bilaterally symmetric I h'
. I b a ' t tck, boat-sh d . .
mvo ute, ut the last whorl un ·1 d ape ' margin rounded· pl· .. . , I ...
shallow; suture ammonitic· s cot._1 e and recurved in the form of a hoot11sp1br~I.' m.1t1ally
' ur ace ornamented . h . . ' um I icus large
an d a Iso near the periphery rows f b wit nbs, bifurcated near the umb ·1· .
• o tu ercles and/or s i . , . . . 1 1cus
Age : Upper Cretaceous ,. p nes ttt po111ts of bifurcation .
Baculites (Fig. 32-23b)
Shell bilaterally symmetrical stra· ht .

which coiled planispirally (~o t:g
I co~1cal or_ flattened; uncoiled, except the apical part
with fine growth lines. s y ost m fosstl); suture ammonitic, surface smooth or

Tttrrilites (Fig. 32-23c)
S~~ll asymm~trical, turreted, conispirally coiled, usually sinistral; evolute (whorls all
visible and I~ contact) suture ammonitic, aperture rounded, left-handed; surface
ornamented with transverse ribs/tubercles.
Subclass : Coeloida
Belemnites (Fig. 32-23d)
Shell radially symmetrical, conical having three major parts, the largest and the most
posterior part is the guard (usually found as fossils) within which fitted phragmocone
from which projected anterialy a tounge-shaped extension (pro-ostracum).
Guard of a belemmite shell is massive, bullet shaped, parallel sided, tapering posteriorly
to a point; indented at its anterior end by a conical cavity alveolus and within alveolus
fitting it exactly is the phragmocone which is a conical, thin-walled structure, projected
outside the guard and alveolus; the shell is septate, septa concave upward, siphuncle along
the ventral margin.
32.9 PHYLUM : ECHINODERMATA

Class : Echinoidea
I. Symmetry; 2. Shape; 3. Thickness and relative dimensions; 4. Margin; 5. Apical/
Oculogenital ring; 6. Periproct (shape and position); 7. Peristome (shape and position); i
8. Ambulacral areas (numbers, arrangement, relative length, shape, relative position with
interambs, nature of plates and poriferous zone); 9. Interambs; I 0. Surface sculpture.
,
Palae(Gco)WP-67
; . --
PALAF,'ON70 LOG Y
528
Anterior
AboraJ Side
Primary Tubercle--,,,~-.
Amb-~ff"rl!l;i~
Scrobicular---,,IA. \lc~I--- Deprcsscd
Granules Porifcrous Zone
Oral Side
Side View Amb (Magnified)
Aboral View
(a) CIDARIS
Anterior
Amb Of Oral Side
With ompound Plates
Aboral Side
And Biserial Pores
.,
AboraJ Amb With
Side View Simple Plate And
Uniserial Pore
Ambs (Magnified)
(b) ECHINOCONUS
Anterior
Porifcrous Zone With
Inner Circular And
Outer Slit-Like Pores
ood Grooves
Side View
lntaamb Amb (Magnified)
(c) CLYPEASTER
FIG. 32 - 24 : ECHINOIDEA
529
Group : Regularia
Order : Cidaroida
Cidaris (Fig. 32-24a)
Test radially symmetrical, spheroidal the summit and th b
W T- . . l d . l ' e ase equally flattened· thick·
L= > , apica . isc arge, mouth and anus circular, central diametrical) o ' osite'.
ambulacra
. five,. radially . arranged , all equal , simple , fl exuous. or
' stra1g
· ht, flyushed
pp with
· . '
mterambulacra, plates simple, pores uniseral, circular, pore pairs in each amb within a
d~pressed zone separated by rows of granules; interambs wide with larger plates each
with a large perforated tubercle surrounded by a ring of small scrobicular granules.
Group : Irregularia
Order : Hulectypoida
Echinoconus (Fig. 32-24b)
Test bilaterally symmetrical, thick, L = W = T, hemispherical, aboral surface highly
inflated with narrowing summit, base flat (giving the test nearly a conical shape), outline
pentagonal to oval; peristome central, periproct marginal, both circular; oculogenital ring
nearly central, small, with four genitals; ambulacra nearly radially arranged, equal sized,
flushed with surface, simple, straight; plates compound, each with three equal demiplates;
pores uniserial near the centre of aboral side to biserial to triserial on the oral side;
interambs with broad plates; surface ornamented with numerous small granules.
Age : Cretaceous
Order : Clypeasteroida
Clypeaster (Fig. 32-24c)
Test bilaterally symmetrical, thin, L = W > T; discoidal, subcircular to pentagonal in
outline; anterior margin rounded, posterior truncated, aboral summit slightly inflated, oral
side flat or slightly concave near peristome; periproct subcircular, inframarginal, peristome
circular, central, within a depression; oculogenital ring central; ambs five, bilaterally
arranged, equal sized, flushed with surface or slightly elevated, petaloid with simple plates;
poriferous zone uniserial, pore-pair with inner circular and outer slit-like; surface with
numerous small granules; oral side with five distinct food-grooves corresponding to five
ambs.
Age : Eocene-Recent
Scutella (Fig. 32-25a)
Test bilaterally symmetrical; thin, L = W > T; discoidal, posterior wider, aboral side
slightly elevated oral side flat; oculogenital ring small, central; periproct inframarginal,
circular; peristo~e central, circular; ambulacra bilaterally arranged, all equal, petaloid,
flushed with the surface; pores uniserial, pore-pair with outer slit-like and inner circular
pores; oral side with branching food-grooves radiating from peristome; surface with
numerous small granules.
Age : Oligocene-Miocene
530 PALAEONTOLocy
Anterior
Aboral Side
Side View
~--.---InncrCucw•
Outer Slit
Like Pores
Oral View
Amb (Magnified)
(a) SCUTELLA
Anterior
Aboral Side
c:::~:=~ Oral Side

Side View
Aboral View
(b) ECH/NOLAMPAS Amb (Magnified)

Anterior
BourrcJq_
Phy~odc J
Floscella Aboral Side
·\Mouth
Oral Side
Pores UniseriaJ Inner

Circular Outer Slit-Like
Aboral View
(c) SLYGMATOPYGUS
Amb <Masnified>
,,vrRoDUCT!ON TU MICROPALAEONTOLOGY
531
order : Cassiduloida
Echino/mnpas (Fig. 32-25b)
Test bilaterally symmetrical; thick; L > w > T· hem· . . . .
side inflated, oral side with a central dep .· ' ,spherical, oval m outline, aboral
. . . ress1on or nearly flat oft t db .
oculogemtal nng shifted slightly toward . . • en runcate ehmd;
tranversely elliptical, inframarginal· an1bulsacarnteb~1lor; mllouth central, polygonal; anus,
• c a 1 atera y arran d fl h d ·
elevated; anterior unpaired one smallest and post' . . I ge ' us e or slightly
· . . . enor pair argest, ambs subpetaloid with
two rows of s1mp1e pates
1 beanng unisenal circular conJugate
· . pores; .mteramb three times
.
larger than am b ; time granules throughout the surface· a poo I d I d fl ·
mouth. • r Y eve ope osce 11 a around
Age : Eocene-Recent
Stygmatopygus (Fig. 32-25c)
~est ?ilaterally sy~metrical; thick; L > W > T; hemispherical, outline nearly oval, aboral
side inflated, o~al side ~at or slightly concave; margin rounded; oculogenital ring slightly
toward postenor; penstome central, polygonal; periproct bottle or wedge-shaped,
supramarginal; ambs bilaterally arranged, sub.petaloid, flushed with surface, anterior
unpaired one largest and posterior lateral-pair smallest; plates simple, uniserial pore-pair
with outer silt-like and inner circular pores; surface with numerous small granules; a
flower-like floscella surrounding the peristome or mouth.
Age : Cretaceous
Order : Spatangoida
Micraster (Fig. 32-26a)
Test bilaterally symmetrical, thick; L > W > T; heart-shaped, truncated behind, dorsal
inflated, ventral flat or slightly convex; oculogenital disc eccentric, shifted towards
anterior, posterior genital absent; periproct marginal, circular; peristome anteriorly shifted,
circular; ambs bilaterally arranged, depressed, subpetaloid to petaloid, anterior pair largest
anteriorly sloping, anterior unpaired smallest but within a deep groove; plate simple; pore
uniserial, circular often conjugate, surface ornamented with numerous granules; on the
oral side posterior interamb bulges out forming a labrum; a s.ubanal/anal fasicole below/
surrounding the anus.
Age : Cretaceous-Ecocene
Hemiaster (Fig. 32-26b)
Test bilaterally symmetrical, thick : L > W > T; heart shaped, posteriorly truncated, but
highest behind; oculogenital ring posteriorly shifted, withoui posterior genital; periproct
marginal/supramarginal, longitudinally oval; peristome cresentic, anteriorly shifted; ambs
petaloid, depressed, bilaterally arranged, anterior one largest sloping anteriorly, posterior
l~terals smallest; plates simples with circular nonconjugate pores in unpaired amb and
silt-like pores in lateral ambs; posterior interamb on oral side bulges out beneath the mouth
forming a labrum; a peripetalous and a lateral fasciole present; surface ornamented with
small granules.
Age : Cretaceous- Recent
.~-i .
Pl\lA EONTOLOGy
532
Brey11ia (Fig. 32-26d) .
. . . 1 h' k· L > w > T; heart:-shapcd, truncated behind; aboral
Test bilaterally syn~dmetrl1ca ··tt t~~t.' oculogenital ring slightly posterior without posterior
s1··de 1'ntlated. ' oral s1 e a. mos ' '
. ·circuhr· .
periproct marginal, subCll'CU
. Iar; am bs b'llateralty
g enital· penstome antenor, semi , . ' 1 . . I
' fl h d ·th surface anterior one smallest s oping anterior y, posterior
arranged nearly us e WI ' . · · · h · I ·
.• ' • · 1 • unpaired amb simple, ind1stmct wit c1rcu ar nonconJugate
pair large~t; llate~ s1mf ~~id with circular conjugate pores placed within depressions
por~s; p~ire) ~m s pbe :bout three times larger than amb; on aboral side, all interambs
(penpochum ; mteram , . . b .
· ne bear three to four rows of large primary tu ere 1es; rest of the
except the posterior o . d ·.. .
surface with fine granules, a Jabrum below the mouth, wh1~h exten s postenally m the
form of a median ridge or carina; a peripetalous and an fasc1?le endopetalous present on
the aboral side and also present a subanal and a lateral fasc1ole.
Age : Oligocene-Recent
Schizaster (Fig. 32-26c)
Test bilaterally symmetrical, heart-shaped; thick; L > W > T; narrow behind; aboral side
inflated; oral side inflated towards posterior and gradually slopping anteriorly making
posterior side highest; oculogenital ring posteriorly shifted without posterior genital;
peristome near anterior margin, semicircular with a labrum behind, periproct marginal
or nearly inframarginal; Ambs bilaterally arranged, depressed, petaloid, anterior unpaired
amb largest sloping anteriorily into a deep narrow groove; posterior laterals smallest; plates
simple with circular nonconjugate pores in unpaired amb and slit-like pores in paired
ambs; numerous small granules on the surface; posterior interamb on oral side forming
plastron with different types of tubercles; a subanal and a lateral faciole present.
Age : Eocene-Recent
Class : Blastoidea
Pentremite~ (Fig. 32-27c)
Test radially _sy_mmetrical, .fl~w~r bud-like, aboral or undersurface of calyx with three
basal~ (two s1m1lar oth~r d1ss1m1lar) and five large radials forming the main parts of the
ca.lyx, upper surface. with five ambulacrals radially arranged, petaloid; each piercing the
. midway of _each rad~al; above the radials and in between two adjacent ambulacral areas
~cur deltmds, five m nui:riber and triangular in outline; mouths at the summit of calyx
m t~e centre and around it are five other openings called spiracles, of which the larger
on~ mcludes the anus; each ambulacral area has a median food groove and on either side
o_f it ~cur two types of plates, elongated lancet plates towards inner part and smaller·
sized s1deplat~s on outer side; each side-plate is.,perforated by a row of marginal pores.
Age : Carboniferous
Class : Crinoidea
Pentacrinus (Fig. 32-27b)
~alyx small, bowl-shaped, consisting of s 11 . . d
hke spines over the stem· arms (b h. . I ma mfrabasals, basals and radials, proJecte
' rac ia s) long, branched, uniserial; smaller branches.
ucTION TO MICROPALAEONTOLOGY
t1fROD s:n
I Anterior
Granules
Uniscrial Conjugate Pore
Aboral Side Circular
lntenunb
c.:~
,\111crior Oral Side
Side View
Aboral View Oral View

f 'MlCRASTER Amb ( Magnified)
Anterior
Amb
Slit-Like
Side View Pore
Aboral View
Anterior (b) HEM/ASTER Unpaired Amb Lateral Amb
Circular Pore Non Conjugate

Aboral Side
Slit-Like Pore
/:) . Oral Side

,\1111:nor
Side View
Aboral View Oral View Unpaired Amb Lateral Amb

(cl SCHlZASTER
Antcri11r
Circular Conjugate
Pore In
Pcripodiwn
SubanaJ
Fasciole
Oral View
Unpaired Amb ·
(d) BREYNU.
.. ~ .
534 PALA EONTOLOGY
(a) ENCRINUS
- - - - - Axillare
(b) PENTACRINUS
Ambulacra
Side View Top View
Baal View
(c) PENTREMITES
/
FIG. 32 - 27 : CRINOIDEA AND BLAS'FOJDEA
...
.
/NIW 1/ >tltT/ > 1 \fl '"' I 'Al .1W )N'/'0/,0 (; Y
(pinnuk s '-'''"' • ,1ff ,111 llw san,~ sid · 111' lh · 11111 i11 hran ·h ; s ·comlary b ran<.: h s hea r !'. mall
pinnuks: st,·1t1 hin ' . p ·ntagn1111I in 1111tli11 · : ·ac h part wilh nu m ber o f pl ate , .
Age : .lu rnssi ·
~,, ·ri1111.,· (Fig. 3 - 7a)
ul yx s:n1 sar-s hapl·d. infrnha sal small on · ·ulc<.1 within ste m, ba sal la rger, he xago na l,
rndial s p ·ntago nal: two fi x ·d hrac hial s in ·ac h ray, th e up per be in g auxili a ry; a rm
bifm ·:Hing; th• brnnc h ·s unis•rial at b ·ginnin, alte rnatin g at middl e and bi seri a l at the
•nd with minut • pinnuks; stem lo ng with a s mall canal.
A,• : Triussi ·
32.10 DES RIPTION OF PLANTS FOSSILS

Schc1w· of d •scriptio n
a) Leaf : imple/compound
(i) Simple : attachment, shape, apex, margin, base and venation.
(ii) Compound : type (pinnate/palmate)~ nature of rachis, shape and attachment of leaflets,
base, margin, apex and venation of leaflets.
(b Stem : Shape, outer ornamentation, sectional view, nodes/internodes, phyllotaxy (if
associated with leat).
(c) Flower : Shape, sepals, petals, reproductive features.
(d) Fruit : shape. petiole, surface sculpture.
A. Gymnospermae
Cycadofilicales (Ptcridospermae)
Glossopteris (Fig. 32-29c)
Leaf-form; simple, shape variable, commonly lanceolate with acute or acut l d d

ape . · · b · 1 e y roun e
. x, margm ~~tire, ase petto ate (mostly broken in fossil); a prominent mid-vein (of
wnh few stra1t1ons) from base to apex. secondary or lateral v . . ten
m· d . • ems smuous emerge from
1 -vem at an acute angles, branched or bifurcated re eatedl ··'
forming reticulate meshes which are elongated and tip y meetmg one another
unicostate venation). mer towards apex, (reticulate-
Age : Permian
Gangamopteris (Fig. 32-29d)
~af-.form;. simple, variable shapes (lanceolnte ov . .

~argm ent1r~ and apex obtuse or acutely round . d~te, _nc1culur, spntulute etc.) base sessile,
f
rom base, diverging towards margin in a curve~ ve1;1s numerous subparallely e merging
~~a.tedly. at a very acute angle meeting each oth:~1-t 1ap~d manner; each vein bifurcates
. ·- m1~-vem, ~nch of veins meeting at medit ormmg very fine elongated meshes·
assuming a mad ve· h' h . m pun of the l • f t· . ,'
- m w ac as obviously not . en o ten formmg a despression
Ar! • . P . contmuous upt I
f>al · ·c , . ~rmtan · o t le npe x (pseudomid-vein)
ac Gco)WP.68 .
,··~
P/\ LAEONTOLOG y
536
--- Mid-Vein
Lateral Veins
Leaflets Joined Racilis

Along Base Leaf - - - - Leaflet
(a) ALETHOPTERIS
.___ _ _ _ Rach is Of 2nd Order
~t(' IJII'----- Rachis Of I st Order

----Apex Rounded
Leaf Margin Entire
Base Sessile
th::.~-Vcnation
}
Leaflets Enlarged
(b) PECOPTERIS
....-----Leaflet
Globular/Orbicular
~::lf::~-:-::Single Vein
Bifurcating At
Apex
Rachis Of 2nd Order
Leaflets Enlarged
- - - Leaflet falcah:
(c) GLEICHENITES
-----Margin Entire
Stem Apex Acute
--'"""fl"""'"m'/A,.,.___ Mid- Vein

......._-.111.!~~~--Lateral Vein
.....,.'fH•l!::..----Base Sessile
Mid-Vein
A Leaflet
(d) CLADOPHLEBJS
Leaf
Fertile Lcat1ets
(e) MARATTIOPSIS
FIG. 32 - 28 : PLANT FOSSILS
•· . .,
cnus TV MICl,Ul~~L-\£()1\ TOJ.nr,y 5.17

/,\ JROV U
\.....-----,-:Two Leaves
Opposite At Node
Margin Entire
Apex Acute
:a::;..._~n---Basc Sessile
111---::i~~~--Stem Striated And 1 ~ - - Stem Articulated
Articulated And Striated
ar-- Apex Acutely lntcmodc

Rounded
Stem
Shoot
Shoot Venation Parallel (b) SPHENOPHYLLUM
Leaf
(a) SCHIZONEURA
I
~
llllf',6,%~1--Subparallel Veins
Bifurcating Producing
Elongated Meshes
Mid-Rib
Leaf
(d) GANGAMOPTERIS
- - - - - - Rectangular Transverse
Segments
(c) GLOSSOPTERIS
' .
Stem
(e) VERTEBRARU
FIG. 32 - 29: PLANT FOSSILS
538 PALAEONTOLOGY
-----Annular Rings
Truncated Apex
Margin Entire
Node
Veins ParaJJel
Divergent
Base Sessile
Leaf
(a) NOEGGERATHIOPSIS
( Cordaites)
--------R~rusStriated
...___ Apex Acute
Margin Entire
Base Sessile
'-----Venation Parallel
Leaflets Showing Multicostatc
Venation
Apex Truncated
Leaf
(c) PTILOPHYLLUM Lateral Vein
b,,,1111~-===J-----Rachis
Margin Enti
6+----~Leaflet
\
H-----Venation Parallel
Mid-Vei
----Margin Entire
Leaf
(d) PTEROPHYLLUM
(e) TAENIOPTERIS
FIG. 32 - 30 : PLANT FOSSILS
,. i ..
INTRODUCTION TO MICROPALAEONTOLOGy
Vertebraria (Fig. 32-29e)

r ir- 519
Stem-form; impression, rectangul
·d f d" ar elongated · ·
s1 es o a me ian furrow or .d m outline segment d
n ge with ' e a 1ternatel
I separated from the adjacent o

Age : Permian
b some longitudinal striat" ..
ne y a transverse rid f
ge or urrow.
Y on two
ions, each segment
Cordaitales
. Cord{l_ites(Noeggerathiopsis (Fig. 32 _30a)
Leaf-for~; simple, broad, spatulate; truncated

base sessile; numerous veins em . f or broadly rounded apex, margin smooth
. ergmg rom base and d" d · ·
manner with occasional bifurc f ,verge towar s apex m a subparallel
veins often intercept the m . a l?n at very acute angles towards the apex; short cross-
am vems.
Age : Permian
Dadoxylon (Fig. 32-30b)
Stem-form· cylindrical ofte I t II . . .
fi ' . . • n a era Y compressed, elltpt1cal m section· outer surface with
;ery me longi~udmal s~riations, trace of outer bark and elongated pr~tuberance (nodes);
central su_bcircular pith surrounded by several concentric growth lines and radially
arranged stnps of vascular bundles.
,•
Age : Permian
Cycadales
\
Ptilophyl/um (Fig. 32-30c)
Leaf-form; compound, unipinnate; rachis depressed/elevated, often striated; leflets falcate/

sickle-shaped, slightly alternately arranged on rachis at acute angles by their whole base;
base sessile, apex acute margin entire or dentate; numerous veins emerging form the base
of each leaflet running in a parallel to subparaJJel manner and again converging at apex
•'.
(multicostate, parallel, convergent venation). ,.•I
Age : Jurassic--Cretaceous - ii
I,
Pterophyllum/Nilssonia (Fig. 32-30d)
Leaf-form unipinnately compound, rachis wide, longitudinally striated also with fine
transverse striation along its two margins; leaflets often of unequal length; rectangular,
opposite or slightly alternate at right angles on rachis, apex truncated margin entire, base
sessile attached to rachis; numerous veins emerge from base of a leaflet run upto apex
in a parallel manner (multicostate-paraJJel).
,,
Age : Jurassic--Cretaceous
1
Otozamites (Fig. 32-31 a) •
Leaf-form; unipinnately compound, rachis narrow, often depressed; leaflets small elliptical,
slightly alternate' on rachis, lying at acute angles, with rounded base making rachis margin ,J
'--'
A .A L .ps.
. I Q a ¢j. $) \ ,4 ;seµ ao :a ·Ci\ ,!(; .
( '
PALA EONTnr n,,\'

540 ,-""---Leaflet Rdangular
Apex Rounded
7---'!ln,;;;..""'"\-- Leaflet Elliptical
~_..--Margin Entire
i;;;::::;.,..--B~e Sessile Notched
Leaf
Leaflets Showing Parallel

Leaflet
Leaf Divergent Venation
(b) DlCTl'OZAMITES
(a) OTOZA HITES
~~.,....,. .illllUI~--Rhombic Leafbases

Spirally Arranged
I(
Stem
(c) BUCKLAND/A
+.!i,-..--Vcnation ~A-- Leaf Acicular

Subparallel Spirally Arranged
Bifurcating
Apex Acute
Venation Parallel
Multicostate
1 t------ Petiole ~'l.\J~- Ste, .. With Rhombic Leal

BMC lmbricatcly Arranged
(d) RH1PIDOPS1S ......--stem
(c) ELATOCLADUS
FIG: 32 31 : PLANT FOSSILS
- ucr10N TO M!CROPALAEONTOLOGY
1NTROD 541
omewhat irregular; leaf margin entire and apex rounded· .
s . . d . . . ' verns, numerous from base
diverging towar s margm, occasionally bifurcating at acute angles.
Dictyozamites (Fig. 32-31 b)
L~af-form; unipinnately compound, rachis depressed, narrow; leaflets narrow rectangular,

shghtly alt~rnately arranged, ~ttached on rachis by upper part of the base, margin entire,
base constricted; nume~o~s vems emerging form base parallely, diverging and bifurcating
repeatedly near apex g1vmg a mesh structure with polygonal elongated meshes.
Taeniopteris (Fig. 32-30e)
Leaf-form; simple, spatulate or rectangular, base petiolate, margin entire, apex truncated
or broadly rounded; a strong mid-vein from base to apex with a few longitudinal striations;
lateral veins parallely emerge at right angles to mid-vein and run towards the margin
with occasional bifurcation near the margin.
Age : Permian-Jurassic
Bucklandia (Fig. 32-31 c)
Stem-form; cylindrical, slightly flattened with persistant elongated rhombic leaf-cushons
(bases), spirally arranged on the stem; each leaf-base convex with a depression at upper
part, individually showing imbricate arrangement; transverse sections show thick upper
bark.
Con iferalcs
Elatocladus (Fig. 32-31 e)
Shoot-form; stem narrow, branched with rhombic leaf-bases, imbricately arranged, bearing
needle-like to narrow falcate-shaped leaf, spirally arranged on stem by their decurrent
bases; leaf shows pointed apex, entire margin and sessile base; venation parallel,
multicostate, convergent.
Ginkgoales
Rhipidopsis (Fig. 32-31 d)
Leaf-form; compound, palmate; 4 leaflets at the tip of the short rachis (striated); each
leaflet spatulate with broad truncated apex, entire, margin sessile narrow base : veins
emerging form base diverging towards apex with repeated bifurcation at very acute angles.
Age : Permian I
,,(
.... . . , ,I.
':-. ·,·l 'J
.i... -
.. • .. •
, ._ . . . ;.
-'l' . -~ ~ •
. ~ .. . 1
Ll -
PALAEONTOLOGY
542
B. Pteridophyta
Equisetales
Schizoneura (Fig. 32-29a) · d ·
·d
arrow or w1 e, ar I t·culated into nodes and mterno es, striated by
Shoot-form; stem n t· us across the nodes; two leaves at each node
. . I .d and grooves con muo '
long1tudma n ges 1 t to spatulate with acute to acutely rounded apex
. ged. each leaf 1anceo a e '
opposite 1Y arran ' . . t d base· many veins emerge from base run towards
. · and sessile but constnc e ,
entire margm d ge at apex (multicostate-parallel-convergent).
apex in a subparallel manner an conver
Age : Permian
Sphenophyllales
Sphenophyllum (Fig. 32-29b)
Shoot-form; stem narrow, art t·culated into nodes and internodes, striated
• longitudinally
by few ridges and furrows; nodes swollen with six leaves arranged m a ~horl, each leaf
spatulate with truncated or rounded apex, entire margin and narr~w constncted ?ase; t~o
veins, emerge from base of each leaf, diverge towards apex with repeated bifurcation
giving a subparallel pattern.
Age : Permian
Filicales
Alethopteris (Fig. 32-28a)
Leaf-form : unipinnately compound, leaflets alternate, at acute angles with rachis, size
of leaflets decreasing towards apical part; rachis narrow but deep; each leaflet elongated,
tongue-shaped with rounded apex, entire margin and broad decurrent base; a strong mid-
veins form base to apex; lateral veins at acute angles with mid-vein, slightly alternate,
bifurcating at the middle.
Age : Permian
Pecopteris (Fig. 32-28b)
Leaf-form; bipinnately compound; rachis of first order narrow and deep, rachis of second
order alternating, each at an acute angle with primary rachis, leaflets small, oppositely
attached at right angles to rachis of second order; each leaflet elliptical, with rounded
a~x, entire margin and broad sessile base; a prominent mid-vein form base .to apex, lateral
vems at an acute angles to mid-vein.
Age : Permian-Cretaceous
Gleichenites (Fig. 32-28c)
Leaf-form.; bipinnately compound; primary rachis narrow, depressed, rachis of second

order at nght. angles to rac~is of first order, oppositely arranged, leaflets small slightly
alternate on secondary rachis; each leaflet orbicular, with rounded apex, entire margin ~
}
INTRODUCTION TO MICROPALAEONTOLOGY ·-1.·
and broad sessile base by whi h . . '·'
c it 1s attached 543 r
base o f eac h Ieaflet which bifu to rachis; one vein · i
rcates near the apex emerge from middle of
Age : Jurassic-Cretaceous one or two times.
Cladophlebis (Fig. 32-28d)
Leaf-form; bipinnately compound . .
two longitudinal ridges and furr • pnmary rach1s (rachis of first order) stout with /
ows,
deep. at actue angles and alternatel secondary rach' ( h' ' one
. is rac is of second order) narrow
. h I y arranged on pnm h' ,
at ng t ang es to secondary rach. . h . ary rac is; leaflets slightly alternate
· is. eac leaflet sickle-sh d f . '
enttre or dentate margin and b d . ape or alcate with acute apex
prominent mid-vein from baa troa sessile base by which it is attached to rachis· ~
se o apex· se d · '
running towards margin each w'th 1 a smg . 'I c?n ar~ veins at acute angles to mid-vein
e b1furcat1on.
Age : Permian-Cretaceous
Marattiopsis (Fig. 32-28e)
Leaf-form· unipinnately d· h'
• . . compoun , rac 1s narrow but deep; leaflet slightly alternate at
acute angles. with rach1s, decreasing in size towards the apical part; each leaflet elongated
sp.atula~e, with rounded apex, entire margin and broad base by which it is attached; a
mid-:em from base to apex; lateral veins at acute angles with mid-vein running towards
margm, lower leaflets fertile showing small elliptical sporangia parallel to lateral veins.
32.11 MOLAR TOOTH OF SOME MAMMALS

Hipparion (Fig. 32,.32a)
Molar tooth elongated, prismatic, squarish in outline; root smaller than crown, hypsodont;
crown surface flat sloping inward, lingual side slightly convex; both inner and outer side
with three longitudinal ridges and two corresponding furrows; upper surface of the crown J.
f .
with three cones : an el1iptical protocone in front, a metacone on outer side and a paracone
on inner side; Each cone with enamel border which shows complicated folding, cement r
r
occurs within the valleys; protocone isolated. ...:...:
.~
Age : Pliocene-Pleistocene
Equus (Fig. 32-32b) '1 .
Similar to Hipparion, except the tooth slightly more elongated with more sloping upper
surface; protocone joined with the outer cusp. t
. •'.--~ '
Age : Upper Pliocene-Recent
Rhinoceros (Fig. 32-32c)
Tooth tabular, squarish in outline; labial side of crown flat; lingual side slightly convex;
upper surface with an outer ectolph, deeply notched at inner side, oblique to ectoloph
occur two smaller cross lophs at the end of outer side. .J
Age : Upper Pliocene-Recent ~

c•.'' .
Palae(Geo)WP-69
-
t\. . . . . .
" '.
L
PALAEONTOLocy
544
Metac0ne
Sloping Upper
Surface
OfCrown
,
, I
Lingual Side View
(a) HIPPARJON
Paracone
Front View f Inner Side)

(b) EQUUS
Inner Side View

(c) RHINOCERAS
FIG. 32 - 32 : MOLAR TOOTH OF MAMMALS
ucrJON TO MICROPALAEONTOLOGY
1NTROD 545
I. .,...r----,.-- Large Cones Fonning
Transverse Ridges
Smaller Connules
-----Root
(a) TRILOPHODON
Abraded Top Of Connules-----

Valleys
With Enamel Border
---Transverse Ridges Made
Up OfConnules
(b) STEGODON
Transverse Ridges-----\
Bordered By Enamel
Upper Surface Undulating
Upper Surface Of Crown
Outer Side View

(c) ELEPHAS
FIG. 32 - 33 : MOLAR TOOTH OF MAMMALS
~·· .
"~ .'
546 PALAEONTOLocy
Stegodon (Fig. 32-33b)

Molar tooth large, elongated rectangular in outline, flat sided,_ root larger than crown·
upper surface of the crown composed of 7 to 8 transverse ridges and correspondin~
furrows; each ridge composed of 5/7 connules, the upper surface of each becomes flat
cross-crested; lophodont. '
Age : Pliocene-Pleistocene
Elephas (Fig. 32-33c)
Molar tooth large, elongated, elliptical in outline, root smaller than crown; crown surface
smooth; upper surface of crown composed of flattened lamellae oc~urring transversely
side by side; the outer margin of each lamella composed of enamel which encloses dentine
materials inside, cement fills the space between the successive lamellae.
Age : Pleistocene-Recent
Trilophodon (Fig. 32-33a)
Molar tooth moderately large, elliptical in outline, upper surface of crown composed of
three and a half distinct transverse ridges; each ridge with two large cones with rounded
apex and a few smaller connules in between larger cones, ridges separated by v-shaped
valleys; lophodont.
Age : Miocene-Pliocene
BIBLIOGRAPHY
Ager,. D. W. (1963). Principles

. of Palae oeco. 1ogy. McGraw-Hills Book
Aguirre, E. ( 1969). Evolutwnarv. lzistorv. ~f e1ep 1zants. Science t 64 t 3Co., INC.
Alexander, R. McN.
. (1975). The Chordate · C am b ndge
. . Press
Univ. ' • 66-76.
· and the fossil· r;
Allen, K. & Bnggs, D. E. G. (eds.)(1989) . Evo l ut,on d B th
Alvarez, W. ( 1990). An extraterrestrial
. . 4 ), 44
. ·ri1c. Amer. , 263(ecor;
impact · S c1ent1 . _52
e . aven.
.
Andrews, H. N. (1961) . Studies m Palaeobot any.. John w·1ley & Sons.
Press.
Archibald, J .D. ( 1996). Dinosaur extinction and ti
,e en d °if an Era. what f ossi/s say. Columbia Univ.
Archibald,vertebrate G
J.D . & Bryant i s· .
A
L J (1990) . o:«.
IJJerentw
· I Crawceous/Tertillry exti11crio11s of non-marine
s. eo · oc. mer., sp. pap~ 247, 549-62.
Arnold,. C . A. (1947). Introduction to Palaeobotany. McGraw-Hills Book Co., INC.
Awasllu, N. (1974). Neogene angiospermous wood. In: Surange, K. R. et. al (eds.). Aspects and Apprisal
of Tnd1a11 Palaeobotany. B. S. I ., Lucknow.
Bailey, J. & Seddon; T. (1994). Prehistoric world. Oxford Univ. Press.
Bakker, R. l . ( 1972). A11atomiC<1l and ecological evidence of emlothenuy i11 dinosaurs. Nature. 238 81 ·
85. •
Bakker, R. J. (1975). Dinosaur renaissance. Scientific. Amer., 232(4), 58-78.
l\aksi, S. K. ( l 972). Fossil [,sh remains from coastal Gondwana Rag/,abapuram M udstone, W. Godavari
dist., A. P., India, Proc. Ind. Nat. Sci. Acad., 381, pt. A(l & 2).
Bak~i. S. K. ( 1976). Pu raj i,•a••idya. Wesl Bengal Book Board. ·
Balakrishnan. M. S. (1974). Fossil Charophyta and Rhodophyta. In: Surange, K. R. et. al (eds). Aspecu
and Apprisal of Indian Palaeobotany. B . S. l., Lucknow.
Bandy, O.L. ( 1964). General corre lotion of foraminifera/ structure with euvi mnment. In : App11K1ches w
Palaeoecology. J. Imbrie & N.D. Newell (eds.) 75-90.
Bandyapadhyay, S. (1999). Gondwana vertebrates of India. pJNSA 65, A(J), 285-313.
Banerjee, D. M. (1970). Aravalli stromatolites from Udaipur, Rajast/um. I. Geo!. Soc. Ind .• 124, 349-55.
Barghoom, E. S.( l 97 l ). The o/de.<t fossils . Scienti fie. Amer.• 224, 30-42.
Barnes, R. D. (1974). Invertebrate Z,mlogy. 3rd ed.,' W. B. Saunders & Co.
Barrington, E. I. w. (l ). Biology of Hemichorcft1ta and p,owcfwrdata. Freeman & Co.
I
965
Bates, R. H. & &st, B. A. (! ). The structure of 0 ,rracod carapace. Lethaia 6. 177-94.
972
Bayer V (1977) C dS t Neues fahrb Geol Palacont., 155. I 65-2 I 5.
• · . ep1aa 1opo - ep en. · ·
Bccrbower, J. R . (1960). Search for the past. Prentice Hall.
Benton, M. I. (1986). The hi.,torv of life on the earth . Kinglisher, London.
\ Benton, M. J.(ed.). ). Phyl,;geny aiul c1a.,sijict1tit>n "! ,e,rc,pt>d.,. 2 vols., Syst. Assoc. sp. vol. 35B ..
0988
Clarendon Press.
Benton, M. I. ( 1997). Vertebrate Palaeontology· Blackwell Science Ud.
(i)
--- - ------
PALAEONTOLOGY
(ii) T ( 1997). Basic Palaeo11tology. Addi son Wesley Longman.

Benton. M. J. & Harper. D. A. . . . ;F.e1·al Res I 95-1 I 8. 3, 187- 195.
,,. H.( 1971).(1973). J.o/Fora mmv, . . ,
Be rgcr. W. . C . rf ·, Press
. . p R ( 1978). Sponges. Univ. a I orma .
8 t!rgqu1st, · · Cl d Press
, ·11 N. J. ( 1965). Th e origin of vertebrates. aren on . .
Bt:rn · · Bl · .1,· ~ London
. h A ( 1992) Human evolutum. ac~1e, .
B1hboroug , · · b 'd u · Press
. 7 Elements of Palaeo11tology. Cam n ge mv. . .
Black, R. ( I9 0). R ., 11 A J (eds. ). ( 1987). Fossil Invertebrates . Blackwell Sc1ent.
Boardman, R. S., Cht!ctham, A. H. & owe ' . .
Publ.
B0 II'1 H M ( J966) Zo11at1011 . . tace. ous to· Pliocene marine se d unents
o1,I' c ,e
' b ase d on p l a11 ktome
.
· fi · · ifi
'
orm1111 era.· Bol . Jnfor· Assoc · Yenz · de Geol. Min. Y. Petrol. 9( I). . . .
.
Bonaparte, J. F. ( I 976). P,sanosau, .us mertii . Casamiquela and origin of ormtluschta. lour. Palaeont.,
50, 808-20.
Bose, M. N. (1966). Plant fossil remains from Rajmahal and Jabbalpur Series in The Upper Gondwana
of India. Symp. Florist. Stratigr. Gondwanaland, B. S. I., Lucknow.
Bown, T. M. & Krause, D. w. (J979). Origin of tribosphenic molar and metathe.rian a,~ eu.theria11 dental
formulae In : Lillegroven,. z. et. al (eds.). Mesozoic mammals. Umv. California Press, 172-
81.
Brace, C. L. ( 1967). Stages of human evolution, human and culture, Prentice Hall.
Brady, H. B. (1884). Report on the foraminifera. Rept. Vox. Challengers, Zoo!., 9, 814pp.
Brasier, M. D. ( 1975). Morphology and habitats of living benthonic foraminifera from Caribbean
carbonate environments. Revta & sp. Micropalaeont., 7, 567-578.
Brasier, M. D. (1980). Mic,vfossils. Gorge Allen and Unwin.
Briggs, D.E.G., Clarkson, E.N.K. & Aldrige, R.J. (1983). The conodollf animal. Lethaia, 16,1-14.
Brown, C. A. ( 1960). Palynological technique. Baton Rounge LA, C . A. Brown.
Brown, J. H. & Gibson, A. C. (1983). Biogeography. Louis & Mosby Co.
Brusca, G. J. (1975). General pattern of invertebrate development. Mad Riv. Press.
Buettner Janusch, J. (1966). Origin of Man. John Wiley & Sons.
Butzer, K. (1977). Human evolution, llominoids of the Miocene. Science, 197, 224-46.
Butzer, K. (1978). Environment, culture and human evolution. Amer. Scient., 65, 572-84,
Camp, C. L. & Hanna, G . D. (1973). Methods of Palaeontology. Univ. California Press.
Carpenter, K. et al. (eds). (1994). Dinosaur eggs and babies. Cambridge Univ. Press.
Carroll, R. L. (1964 ). Earliest reptiles. Jour. Linnearn Soc. Zool., 45, 61-83.
Carroll, R. L. ( 1969). Vertebrate palaeolltology and evolutio11. W. H. Freeman.
Chang, K. (1977). Chinese Palaeoanthropology. Ann. Rev. Anthr. 6, 137-59.
Chandra, S. (J 974). Fem an.d Fem-lik~ plants from Lower Gondwana. In : Surange, K. R. et. al (eds.)
Aspects and Appr,sal of lnd1a11 Palaeobotany. B. S. I., Lucknow.
Chandr~, S. (1974). Glossopteris and allied genera-Marphological studies. Ibid., 126-143.
ChatterJee,SS. (1991 ). Cranial anatomy and relationship of a Triassic bird ft~m Texas. Phil. Trans. Roy.
oc. sr. B, ]32, 227-346.
Ch~tterJee, S. ( 1995). The Triassic bird Protoavis., Archaeopteryx 15- ~ I
n
Chtppl k G w ' · • _., ·
on ar, . . (1944). Algae in the Cretaceous of Narmada valley, Sci·. Cult.,
r- ·-
10, 130-31.
.. '
,, .
BfBLfOGRAPHY
Chitaley, S. D. ( I97~ ). Palaeogene cmRiosperms (ex., . iii

and Appn.ml of Indian Palaobotocm . ;ept111g wood). In : Surange, K. R. , ( )
D. L. et. al ( 1981 ). Conodonta In . R b~ . S. I. Lucknow. et. al. (eds). Aspects
Clark ' . · · o inson R A
part W, Geo I. Soc. Amer. and Kan .. . U . · · (ed.). Treatise 011 Inv, , 1
. .. sas niv. Press. ere Jrate Palaeontology.
Clark. R. B. ( 1964 ). Dyna nu cs of metawa11 e l .
vo ut1011 · The · . ;? . ..
Press. . . , ong111 of coelom and segments. Clarendon
Clarkson, E. N. K. (1986). In.vertebrate Palaeollt01
. .• · ogy a11d evolution All. & U . ·-
Cloud, P. E. ( 1968). Significance of Gunflint (P , . b .· . · en nwin Publ. Ltd.
recam uan) nucroflora Science 2(148 ~
Cloud, P. E. ( 1968). Premetazoa11 evolution and . . · • ), 27-35.
Centennial papers, Yale Univ. Press. ongm of the metawa. Yale Univ. Peabody Museum
Coates, M. J. & Clark, J. A. (1990). Polydactyly in earl' ·t k . . .
ies · 11ow11 tetrapod limbs. Nature J47 66-69
Colbert, E. H. ( 1951 ). The dinosaur book. 2nd ed. McGraw-Hills Book Co. '· ' ·
Colbert, E. H. ( 1961 ). Dinosaurs, their discovery and their world. E. P. Dutton & Co.
Colbert, E. H. (1965). The age of Reptile. W. W. Norton.
Colbert, E. H. (1973). Wandering lands and animals. E. P. Dutton. ,.._
Colbert, E. H. ( 1982). Evolution of vertebrates, 4th ed. John Willey & Sons. .•
Colbert, E. H. ( 1989). The Triassic dinosaur Coelophysis. Museum Northern Arizona, Bull. 57, 1-160.
Courtillot, V. (1990). A volcanic eruption. Scientific Amer., 263(4), 89-107.
Cox, C. B., Healey, L. N., & Moore P. D. C. (1973). Biogeography : An ecological and evolutionary
approach. Blackwe,11.
Crain, I. K. (1971). Possible direct cause relation between geomagnetic reversal and biological exti11ctio11.
Bull. Geol. Soc. Amer., 82, 2603-2606.
Crimes. T. P. & Harper, J. c. (eds.). (1970). Trace fossils. 1-2, Geo!. Jour., sp. issue. 9, Seel House
Press.
Cushing, E. J. & Wright, A. E. (eds.). (1967). Quaternary Palaeoecology. Yale Univ. Press.
., .
Cus hman, J . A . ( 1948)
. C' • :I'. a the,·r classiifi1cation and economic use. 4th ed. Cambridge Mass.
. rorammt.,e~ , ..,. .
Harvard Univ. Press.
Cushman. J. A. (1959). Foraminifera. Harvard ·uni"v. Press.
Darrah, W. C. (1960). Principles of · Palaeobotany.
· Rona Id Press Co · •
. h if I d'a
O 1 World Press, Kolkata.
Dasgupta, A (2010). Phanerozaic strat1grap Y n •
Day, H. Michael (1969). Fossil Man. Hamlyn Publ. Co. I ( d ) . A peers and Apprisal of Indian ..
Deb, S. (1974). Mesozaic pteridophytes In : Suran ge K · R· et · a c s. . . s
Palaeobotany. B. S. I., Lucknow. . · H 1t Rinehart & Winston.
D l · of Fossil p1ants. o
eleroryas, T. (1963). Morphology and evo uttoll . .-. 220 47-50.
D t Nice Scicnuhc Amer., ,
e Lumley, H. (1969). A palaeotithic camp a . · . human remains from Arago cave, southeastem
De Lumley, H. & De Lumley, M. ( 1973). Pre-Neandlerthall1 162-68. . -
s
,-..' ~
· t Anthropo ogy. ' B
D·
France. Year book of Phys1ca
h
enton, E. J. (1974). On buoyancy and t e 1Je
[,I'. oif modem an
.d fiouil cepha/opods. Proc. Roy. oc .•
· ~
185, 273-99. . '] d •d Columbia Univ. Press.
Doh 2 h · · Oif species. -1 r c ·•
ansky Th. ( 1950). Genetics and orzgm • . - J82 32-41.
D0 b2 h 1 · . Sc1enuhc Amer., '
ansky Th. ( 1951 ). Genetic basis of evo utwn. . h Id Publ Co
O
Dods E d Product Rem • •
on, · 0. (1960). Evolution-Progress an ·
.£ es .tac ]>?· 4 0
PALAJiON'fOLOGy
(iv) ., · J'Jmce.,·.,·e.,· and ,vlden ·e. Bclhcvcn.

( 1989) Ma.,·s t!Xf 1" ,m1'
Donovan, S. K. (ed. .). · . ,,. M"i·ozoic-Cenozoic c:limate clw11J.:e.1·. Pro<.:. North Amer.
Dorf. E. ( 1970). Palaeo b ot,~111c · al ev1dem·e
. OJ "· •
. c0 nvcn11on I. 323-47. . .
Palucont. . • L ( 1983 ). Updatinf( the theory of Mutatw11, Amer. S<.:1cnt. 73,
Drake, J. W, Glickman, B. W. & Ripley, . .
62-30.
4) Mik,v11eologie, 2 vols. London.
Ehrenberg. C . ( 185 · o .
E' h. D L. (1976). Geological time. Prentice Hall. . .
'.c er, .
Eimerl. S. & lrvcn, ·
• D ( 1972) The Primates. Timc-Lifc-Internallonal.
·
Eld ed '. N. ( 1997). Fossils. Princeton Univ. Press. . .
r gc, . . f 1 ( 1980) Phy/o"enetic: pattern and evoltllionary proc:e.,·ses. Columhia Univ.
Eldredge, N. & Crucra l, . · o
Press.
Eldredge, N. & Stanley s. M. ( 1984 ). living fossils. Springer-Verlag. . . .
Elliott. D. K. ( 1987). A reassessmelll of Astraspis desiderata, the oldest North Amenca ve,1ebrate. Science,
237, 190-2.
Erdtman. G. ( 1972). Pollen morphology and plant taxonomy : Angiosperm. Harper Publ. Co.
Ericson, D. B., & Wollin, G. ( 1968). Pleistocene climate and chronology of deep sea sediments. Science,
162, 1227-34.
Erwin, D. H. (1993). The great Palaeozoic crisis. Columbia Univ. Press.
Fastovsky, D. E. & Weishampcl, D. B. (1996). The evolution and extinction of dinosaurs. Cambridge
Univ. Press.
Feduccia A. (1995). Explosive evolution in Tertiary birds and mammals. Science, 267, 637-8.
Frerich, W. E., Heiman, M. E. et. al., ( 1972). latitudinal variation of plankto11ic test porosity. J. Foram.
Res. 2, 6- I 3.
Frey, R. W. (ed.). ( 1975). The study of trace fossils. Springer-Verlag.
Funnell, B. M. ( I967). Forami11ifera & Radio/aria as depth i11dicators in the marine environment. Mar.
Geol. 5. 333-347.
Funnell, B. M. & Riedel, W.R. (eds.) (1971). The Micropalaeontology of oceans. Cambridge Univ. Press.
Galloway, J. J. ( 1930). A ma11ual of foraminifera. Principia Press.
Gallon, P. M. ( 1978). Fabrosauridae, The basal family of Omithschian dinosaurs. Palaeontologische
Zeitschrift, 52, 138-59.
Garstang, W. (1922). The theory of recapitulation : a critical statement of biogenetic law. J. Linn. Soc.
Lond., 35, 81.
Garstang, W. (195 I). larval forms and other zoological verses. Blackwell.
Gee, H. ( 1996). Origin of vertebrates. Chapman & Hall.
Gelvin, B. ~.980). Morphometric affinities of Giga11topithecus. Amer. jour. Phys. Anthropology, 53, 541·
6
Geological Survey of India Sp. Publ ( 1988).
Glaessner, M. F. (1944 ). Principles of Micropalaeomology. John Wiley & Sons.
Glaessner, M. F. (1962). Percambrian fossils. Biol. Rev.37, 467-94.
Glacssner, ;5~9!2~~e, M. ( 1966). late Precambrian fossils from Ediacara, South Australia. Palaeo11t.,
Gotham, ~ : Sahni, B. (1937). Fossil pla,w; from Po Series of Spiti. Rec. Geol. Surv. Ind. 72(2) .t95·
/
HIBLIOGRAPlll' (v)
i(lUld. S. J. ( 1977). 11to~et1 ) and PJ,y/ogeny. Cambridge Univ. Press.

Gowdu. S. ( 1974). Preca111/Jrit111 a11d Cambrian l'la11t.\·. In : Surangc, K. R. et. al. (eds.). Aspects and
Appris 11 <l Indian Palaeobotany. B. S. I., Lucknow.
Gowdn. S & Srinivns. T. N. ( 1969). Microfossils form Arc/wean cqmplex of Mysore. J . Geol. Soc. Ind .,
10(2). 201-8 .
Grnnt, V. ( 1963). TJ,e Ori~in of adaptation. Columbia Univ. Press.
Hullnm. A. ( •d.) { 1977). Patterns of evolution. Elsevier Puhl. Co.
Hurpcr C. W. ( 1975). OriE,:i11 of species i11 geologic time . Science, 190, 47-48.
Hnynes, G. ( 1991 ). Mammorl,s, Mastodon ts and Elephants. Cambridge Univ. Press.
Hnynes. J. ( 1965). Symbiosis, wall structure and habitats in forami11ifera. Cush. Fond. Foramin ., Res.
16. 40-43.
Hedley, R. H. & Adams, C. G. ( 1974) ( 1976). Foraminifera. Vol-1-2. Lond. Acad. Press.
Henning. W. ( 1966). Phylogenetic sy.ftematic:s. Univ. Illinois Press.
Heywood. V. H. & Mencil, J .(cds). ( 1964). Phenetic and Phylogenetic: classification, Syst. Assoc., Publ. 6.
Hoq. B. U. & Boersma, A. (eds.). ( 1978). l111roduction to marine Micropalaeontology. Elsevier Pub I-.
Co.
Horner, J.R. (1984). Tire nesting bel,aviour of dinosaurs. Scientific. Amer. 250(4), 92-99. ·
Horner. J.R & Dobb. E( 1997). Dinosaur Lives, Harper Collins.
House, M. R. (ed.) ( 1979). The Origin of major invertebrate groups. Syst. Assoc .• sp. vol. 12, Acad.
Press.
Hyman, C. H. ( 1959). The Invertebrates. McGraw-Hills Book Co.
Hynes. J. ( 1965). Symbiosis, wall structure and habitats in Forami11ifera. Cush. Found. ·Foram. Res. 16,
40-43.
Jain, K. P. ( 1974). Fossil Dinoflagellates, Acritarchs, Tasinanitid... and Namwplcmkton.s. In : Su range,
K. R. et. al. (eds.). Aspects and Apprisal of l11dia11 Palaeobotany. B. S. I., :Lucknow.
Jain. S. L. ( 1973). New specimens of lower Jurassic holostea11 fishes from India. Palaeont., 16, 149-77.
Jain, S. L., Kully, T. S., Roychowdhury, T. K., Chatterjee, S. ( 1975). The Sauropod dinosaur from Lower
Jurassic Kora Formation of India. Proc. Roy. Soc. Lond., 188, 221-228.
Jain. S. L., Robinson, P. L. & Roychowdhury. T. K. ( 1964). A new vertebrate fauna from Triassic of the
Deccan, India. Q. Jour. Geol. Soc. Lond., 120( I). 115-24.
Jarvik, F. ( 1996). Tire Devonian tetrapod lcthyostega. Fossil & Srata, 40, 1-213.
Jenkins. D. G. ( 1965). Pla11ktonic foraminifera and Tertiary interco11tine11tal correlations. Micropal. 11 ;
265-277.
Jenkins. F. A. Jr. & Crompton. A. W. ( 1979). Triconodo11ta. In : Lillegraven, A. et. al (eds.). Mesozoic
mammals. Univ. California Press. 74-90.
Johanson, D. C. & White, T. D. ( 1979). A systematic assessment of early Aji'ica11 hominids. Science,
203, 321-30.
John, B. S. ( 1977). The ice age : Past and Present. William Collins & Sons.
Jones, D. J. (1956). Introduction to microfossils. Harper & Brothers.
Jordan, E. L. & Verma P. S. ( 1981 ). Chordata zoology and elements of animal physiology. Chand &, Co.
Ltd.
Juker, T. H. ( I 966). Molecules a11d evolution. Columbia Univ. Press.
Pulal!(Gl!o)WP-70
-
,. ~ \ ·,
·.;. '•} :'
_ .. -· . ' t.. ' l .....
•
PALAEONTOLOGY
(vi) Kuchneotherium preceursoris. Zool. Jour. Linn.
Kermack, D. M. et aI. ( 1968) · The Welsh pantothere
Soc., . 47, 407-23. . . C B s Publ.
Kathal P. K. ( 1998). Microfossils and their appltcatwns. . . 1. Acad Press.
• . · · ofmammas. ·
Kemp, T. S. (1982). Mammal-like replttes .and origm
64) 0 · · 0,,. Fe
1 Reinhold
Keosian, J. ( 19 · rigm 1 1' • . • ven J A 9 79 ). Eutheria. In : Lellegraven, J.A. et al.
Kielan-Jaworowska, Z., Bown, T. M. & Lillegr~ '. ·Pr· O
(ed ) M · mammals Univ Cahforma ess. .
s. . esozotc . Ii e and evolution of graptolithina. Proc. Geol. Soc.
Kirk, N. ( 1969). Some thoughts on ecology, mode of i/1
Land., 1659, 273-92. d 128 127 3.,
· · ,,. f't s Jour Geol. Soc. Lon ., , - .1.
Kirk, N. ( 1972). More thoughts on automobtllty 01 g rap to I e · ·
.
Kitt, D. B. (1974 ). Palaeontology and evo luttonary th eo ry · Evolution, 28, 458-72.
KJein, R. (1969). Man and culture in Late Pleistocene. Chandler Publ. Co.
Knowlton, F. H. (1957). Plants of the past. Princeton Univ. Press.
Kormondy, E. J. (1991). Concept of Ecology. 3rd ed. Prentice Hall.
Krishnan, M. S. ( 1968). Geology of India and Burma. Higginbothams. . .
Krumbein, w. c.
& Garrel R. M. (1952). Origin and classification of chemical sediments m terms of
,JI and oxidation-reduction potential. four. Geol., 60, 1-33.
Kummel, B. & Raup. o. M. (eds.). (1965). Handbook of palaeontological techniques, Freeman & Co.
Kutty, T. S. (1972). Permian reptilian fauna from India. Nature, 237, 462-463.
Lakhanpal, R. M. (1974). Physical condition of bulian Tertiary in the light of palaeobotanical evidences.
In: Surange, K. R. et al. (eds.). Aspects and Apprisal of Indian Palaeobotany, B.S.I., Lucknow,
516-24.
Lakhanpal, R. M. (1974). Floristic evidences of stratigraphical subdivisions of Indian Tertiary. In :
Surange, K. R. et al. (eds). Aspects and Apprisal of Indian Palaeobotany, B.S.I., Lucknow,
496-501.
Laporte, L. F. (1968). Anceient environments. Prentice Hall.
Larwood, G. P. & Neilson, C. (1981 ). Recent and fossil Bryozoans. Olsen & Olsen.
Leaky, L. S. B. (1967). An Early Miocene member of hominidae. Nature, 213, 155-63.
Leaky, L. (1967). Olduvai Gorge. vol. 1, Cambridge Univ. Press.
Leaky, M. G. et al. ( 1995). New 4 million year old hominid from Kanapoi and Allia Bay, Kenya, Nature,
376, 565- 71.
Leblich, J. A. R. & Tappan, H. (1964). Protista and Sarcodina. In Tretease on Invertebrate palaeontology.
Part. C.R.C. Moore (ed.) (2 vols) Geol. Soc. Amer. & Kansas Univ Press.
Lehman, V. (1981). The Ammonites, their life and world. Cambridge Univ. Press.
Linsley, R. M. (1977). Some laws of gastropod shell fonns. Palaeobiology. 3, 196-206.
Linsley, R. M. (1978). Shell forms and evolution of gastropods. Amer. Scient., 66, 432-41.
Lucus, S. G. & Luo, Z. (1993). Adelobasipleus from Upper Triassic of West Texas : the oldest mammal.
Jour. Vert. Palaeont., 13, 309-34. ·
Lull R. S. ( 1945). Organic evolution. Macmillan & Co.
Mahabale, T. S. _0974). Cenozoic pteridophyta. In : Surange, K. R. et. al. (eds). As ects and Apprisal
of Indian Palaeobotany B. S. /., Lucknow. 72-8. p
Maheswari. H. K. (1974). Palaeozoic lycopsida and sph~nopsida. Ibid., 51- 1.
6
/ '
''
a a 4
..
._,.,,. .
BIBLIOGRAPHY
Maity, P. K. ( 1974 ). Pre-Gondwana Land l .

P ams. Ibid., 47-50 (vii)
Manton, S. M. ( 1973). Arthropod phylog ·
e11y-a modern synthe\·is Jo
Manton, S. M. ()977). The Arthropoda· hab't fi . . . ur. Geol. Soc. Lond. 171 111
. ' I s, unctt011al mor hot . ' -30.
Martin, P. S. & Klem, R. G (eds.). (198 4) Q 1 . P ogy and evolution, Oxford Univ p
· ua ernary extinct' . rcss.
Press. ions : a prehistoric revolution u · A .
. n1v. nzona
Martin, R. D. ( 1990). Primate origin and evol t.
.. u wn, a phylogenetic approach Cha man
Mathew, W. D. ( 1929). Crtt1cal observations of s· /'k . . · P & Hall.
60. iwa I mammals. Bull. Amer. Mus. Nat. Hist., 56, 437-
Mathew, W. H. ( 1962). Fossils : an imroduction to pre h · . /;I'. B
. - is1one 1.Je. arnes & Noble Book.
Maynard Smith. J. (1975). The theory of evolution Ha d h p .
. . rmon swort , engum.
Maynard Smith, J. (ed.). (1982). Evolution now- a century a•.,,ter D · M - .
. , 'J' arwm. acm1 11 ian & Co.
McAleaster, A. L. (1968). The history of life. Prentice Hall.
McGlassor, R. H. (1959). Foraminiferal biofacies around Santa Catalina Island Ca/il'omia. Micropal
5. 217-240. · ' 'J' .
Meglitch, P. A. & Schram F. R. (1991 ). Invertebrate Zoology. 3rd ed. Oxford Univ. Press. •
'i
Meyen, S. V. (1987). Fundamentals of Palaeontology. Chapman & Hall.
Misra, R. C. (1974). Origin and distribution of plant life. In : Surange, K. R. et. al (eds.). Aspects and
Apprisals of Indian Palaeobotany. B.S.I., Lucknow, 359-68.
Montagu, A. (1965). Homo habilis. Science 149, 918.
Moody, P. A. (1970). Introduction to evolution, 3rd ed., Harper Brothers.
Moody, R. ( 1977). The fossil world. Hamlyn Publ. Gr. Ltd.
Moody, R. (1986). Fossils. Macmillan.
Moore, R.C. (ed.). (1956). Treatise on Invertebrate Palaeontology. Part F Coelentera. Geol. Soc. Amer.
& Univ. Kansas Press.
Moore R.C. (ed.). (]957). Treatise on Invertebrate Palaeontology. Part L. Mollusca 4 (Ammonoids). Geol.
Soc. America & Univ. Kansas Press.
Moore, R. C. (ed.) (1955). Pt. P, Arthropoda 2, Ibid.
Moore, R. C. (ed.) (1959). Pt. 0, Arthropoda I, Ibid.
Moore, R. C. (ed.) (1961). Pt. Q, Arthropoda 3, Ibid.
Moore, R. C. (ed.) (1964). Pt. C, (Protista I-2). Ibid. 'd
Moore, R. C. (ed.) (1964). Pt. K, Mollusca 3 (Nautt
·101'd )
s· lb, ·
Moore, R. C. (ed.) (1965). Pt. H, Brahiopoda. Ibid.
Moore, R. c. (ed.) (1966). Pt. s, Echinodermata 1. Ibid.
Moore, R. c. (ed.) ( 1969). Pt. I, Gastropoda. Ibid.
Moore, R. c. (ed.) 0 9 69 ). Pt. N, Gastropoda. Ibid.
Moore, R. C. (ed.) (1975). Pt. R, Arthropoda. Ibid. b te nataeontology. Part T. Echinodermata
.
Moore, R. C. & Teichert (eds.). · on Jnverte ra n
(l 978). Treatise
•
2. Ibid. . 52 . Invertebrate Fossils. McGraw-Hills Books Co. ;
Moore, R. C., Lalicker, C G., & Fisher, A. G. (t 9 )
Morton, J. E, (1969). Molluscs. Hutchinson, London. I a unch1'11s Syst. Assoc. 3, 61-80.
N' · ·rregu ar se ·
ichols, D. (1959). More of life and taxonomy Ill '
].
!:
PALAEONTOLOGY
(viii)
N" h I D (1974). Echinoderms. Hutchinson, London. .
ic O s, · . d
Nichols D ( 1975). Uniqueness of echmo erms. x o
o i rd Univ Press.
. .
• · C T ( 1985 ) Palaeontology _ A11. Jntroductwn . Pergamon Press.
Nield, E. W. & Tucker, V. . . . ..
63). History of fishes. Hill & Wang.
Novman, J. R. (19 - .
1965). Evolution of life. Weidenfield & Nicolson .
OJ sen, E. C . ( . . J ) 20-24
'B . E (1984). What was the Acheulean hand axe. Nat. Htst., ( une ' .
O nen, . . . p
Padian, K. (ed.) ( 1986). The beginning of the age of Di11osaurs. Cambridge Umv. ress.
Pal, A. K. ( 1971 ). Algal flora of Bagh group of Madhya Pradesh Sem. Palaeo. Palyn. Ind . Stratigr.,
Calcutta. abst. 35.
Phleger, F. B. (1960). Ecology and distribution· of recent forami11ifera . Baltimore : John Hopkins Press.
Poirier, F. E. (1990). Understanding human evolution. 2nd. ed., Prentice Hall.
Postuma, J. A. (1971). Manual of planktonic foraminifera. Amsterdam, Elsevier. ·
Prakash, U. (1974). Palaeogene angiosperms. In : Surange, K. R. et. al. (eds.). Aspects and Apprisal oj
Indian Palaeo~otany. B.S.I., Lucknow, 306-20.
Prothero, D. R. & Schoch, M. (eds.). (1989). Evolution of Perissodactyls. Oxford Univ. Press.
Puri, G. S. (1948). Preliminary note on the Pleistocene flora of Karewa. Q. Jour. Geo!. Min. Met. Soc.
Ind., 20, 61-66.
Ramsay, A.· J. S. (ed.) (1977). Oceanic micropalaeontology. 2 vols. Academic Press.
Rao, R. & Pia, J. (1936). Fossil algae from uppermost Cretaceous beds (Niniyur group) of Trichinopo/y
dist., South India. Mem. Geo!. Surv. India., Palaeont. Indica, n. s., 21, 1-47.
Raup. D. M. (1966). Geometric analysis of shell coiling : Several problems. Jour. Palaeont., 40, I 178-
90.
Raup. D. M. & Stanley, S. M. (1971, 1985). Principles of Palaeontology. 1st and 2nd ed. ·Freeman ·&
Co.
Ray, A. (2008). Fossils in Earth Sciences. p. 429, Prentic Hall ·
Ray, S. (2001 ). Small Permian dicynodonts from India. Pal. Res., 5(3) 177-91:
Ray. Y. (1986). Neanderthals: a new look at an old face. Jour. Hum. Evol., 15, 151-64.
Ritchie, A. & Gi_Iber-Tomlinson, J.(1977). First Ordovician vertebrates from the southern hemisphere.
Alchermga. I, 351-68. .
Robinson, R. A. .<ed). (1983). Treatise on Invertebrate Palaeontology. Pt. G. (Bryozoa). Geol. Soc. Amer.
& Univ. Kansas Press. · ·
Robinson, R. A. & Teichert C (eds) (1979) TJ • · · .
Fi 'I" . .' · ·· · reatlse on Invertebrate Palqeontology Pt. A (Introduction,
oss, izatwn, Bwgeography and Biostratigraphy). Geol. Soc. Amer. & . Univ. Kansas Press.
Romer, A. S. (1966). Vertebrate Palaeontology. Chicago Univ. Press.
Romer, A. S. ( I 967) Ma· t · .
. f}Or s eps ,n vertebrate evolution. Science, 158 1629-37
Romer A S & p . · ' .· ·
' . . arsons, T. S. ( 1970). The vertebrate body 4th ed W B S d
Rose K o (19 82 · · · . ·• · · an ers.
Rose' K . I) . ). Skeleton. of D~acodexis, Oldest known· artiodactyi. Science, 216.' 621-23.
. . (1995), The earlier pnmates Evol. Anthr., 3 159-73 .
Roychoudhury T K { 1965 ' ··
. .• · · ). A new metaposaurid am h ·b · fi . · if
. Central India. Phil. Trans. Roy Soc., (B) P ., ,a rom Upper Tnassic Maleri forma11011 °
Rudwick M J S , 25 0 , 152.
' ·. · · 0~64). Inference offi · · ·
27-40. . un~t,on from struct~re in fossils. Brit. Jour. Philos. Sci., ·15B,
"'tr;
· $Q.sa
otBLfOGRAPHY
dwick, M. J. S. ( 1970). living and fossil b ·h . (ix)
RLI • rac topods, Hutchin .
ni A. (200 I). Dmosaurs of India. Nat B k T son.
Sa h • . · oo rust, India.
"hni, B. ( 1932). A petrified Williamsonia rrom R . I .
S" .,, a1maia/ f1t/l\· 1 /' p
Sahni, B. (1936). Karewas of Kashmir. Curr. Sci ., (5). 10-16. · • lllta. laeontologia Indica. 20.
Sahni ' B. (·1953 ). Notes on possible psi/ophvre .

· remams from Spiti NW f-1' I
Saidova, Kh . M. ( 1967). Sediment stratigraphy a d . • tma aya., Palacobot., 2, 1-3.
1
forami11ifera during Quatema,y Prag -~ . ~a aoeoge.ography of the Pacific ocean by benthonic
. tess Ill ceangraphy, 4, 143-157
Saluja, S. K., Rehman, K. & Rawat, M. s. ( 197 I) M ' . . ·
Congr., Pt III, 318. · tcmplanktons form Bhimas. Proc . 58th Ind . Sci.
Saluja, S. K. & Arora. C. M. ( 1971 ). Fossil palyno . I fi .

Palyn. 1 I' 65-83. mo,p 1s orm V111dhya11 of Rajastha11. Rev. Palaeobot.
Savage, R. J. & Long, M. R. ( 1986). Mammal evolution . Brit . Mus . Na t . H'.

1:,t. , Lon d on .
Schafer, C. T. &
. Pelletier . · I st Int · sympostum
. ' B. R.. ( 1977) . · on bentho111c· foram111ifera
. . of co11ti11e11tal
margm. Sp. Publ. Manl. Sediments. No. I. (2 vols).
Schafer, W. ( I 972). Ecology and Palaeoecology of marine environment. Chicago Univ. Press.
Scoh, D. S. & F. S. Medioli ( 1978). Vertical zonation of marsh foraminifera as accurate indicators oj
former sea level. Nature 272. 528-531.
Scrutton, C. T. (1965). Periodicity in Devonian coral growth. Palaeont., 7, 552-8.
Sereno, P. C. & Novas, F. E., (1992). Complete skull and skeleton of an early dinosaur. Sci., 258. I 137-
40.
Sereno, P. C. et. al. ( 1993). Primitive dinosaur skeletom from Argentina anct::'early evolution of Dinosauria.
Nature, 361, 64-6·6.
Shaw, A. B. (1964 ). Time in stratigraphy. Mc Graw-Hills Book Co.
Shukla, A. C & Mishra, S. P. ( J 975). Essential of Palaeolllology. Vikash Pub I. House.
Shenk, E. T. & Mc Master, J. H. (1956). Procedure in Taxonomy. Standford Univ. Press.
Shreeve, J. ( 1996). The Neanderthal enigma : solving the mystery of modem human origins, Viking.
Shrock, R. R. & Twenhofel, w. H. ( 1953). Principles of Invertebrate Palaeontology. McGraw-Hills Book
Co.
Simons, E. L. ( 1954 ). Dawn ape of the Fa yum, Nat. Hist., 93, 18-20.
Simons, E. L. ( 1967). The earliest ape. Scient. Amer., 217( 12), 28-35.
Simons, E. L. & Chopra, S. R. L. ( 1969). Gigantopithecus (Pongidae -Hominoidea) : a new species fmm
north India. Post ilia, 138, 1-18.
Simpson, G. G. (1953). The major features of evolution. Columbia Univ. Press.
Simpson, G. G. ( 1961 ). Horses. Anchor Garden City.
Simpson, _G. G. ( 1967). The meaning of evolution. 2nd ed., Yale Univ. Press. . . .
Sliter w v· bathymetric distribution of benrho111c forammifera. J.
' · . & Bakes, R. A. ( 1972). Cretaceous .
Foramin. Res. 2. J 67-183.
s .
muh, A. (1984). Echi11oid Palaeobiology. Allen & Unwin.
Sneath, P.H.A. & .Sokal, R.P. ( 1973). Numerical Taxonomy. Freeman & Co. . . . .. .
Sokal R p · /· . . 'pies and practice of Srat1st1cs III bwlog,cal
' · · & Rohlf, F. J. ( 1969). Biometry : t 1e prmct
research. Freeman & Co.
__._ --=-- - -
PALAEONTOLOGy
(x) . h E, m Bhima Series of Goulburga dist .. Mysore

· · asa T. N. & Gowda, S · S · ( I 970). · Fossil
. .
Acntarc 1 ro
p
. .
Symp. Purana Form. Penmsu 1ar 1nd1a, Univ.
Snmv ' . d h . logic significance. roe.
state /11d1a an t etr geo
Saga~ (M. P.), India. . W dhyan Formation of India.· Proc . Palaeopalynot
. R M ( 1971 ). Microorganic remams from m .
Srivastava, · ·
Sem., Calcutta. F· h bits in the bivalia (mollusca). Geol. Soc. Amer. ,
S M ( 1970). Relation of shell form to i1e a
Stan Iay, . ·
125
Mem., · . d lution of byssally attached bivalved mollusks. Jour.
Stanley, S. M. ( 1972). Functw11al morpho 1ogy an evo
Palaeont. , 46, 165-212.
79). Macroevolution : Pattern and process. Freeman & Co.
Stan Iey, S. M . ( 19 .
. G
S te bb ms, . . L (1966) Process o~ organic evolution. Prentice Hall.
. 'J • B II A
.
Stelh, I. G. & Creath, . . w B (1964) p;
. o raminVeral
'J'
ratios and regional envtromnents. u . mer. Ass.
Petrol. Geol., 48, 1810-1827.
Stewart, W. N & Rot h we II , G . W(l993) . Palaeobotany and Evolution of Plants. (2nd ed.), Cambridge
Univ. Press.
Surange, K. R. (1966). Distribution of Glossopers flora form Lower Gondwana, India. In : Symp. Florist.
Stratrg., Gondwanaland.
Swinnerton, H. H. (1950). Outline of Palaeontology. 3rd ed., Edward Arnold & Co.
Swinton, W.E. ( 1965). Fossil birds. Brit. Mus. Nat. Hist. , London.
Sylvester-Bradly P. C. (ed.). The species concept in Palaeontology. Syst Assoc ., Publ. 2.
Szalay F. S. Novcek, M. J., & Mckenna, M. C. (1993). Mammal phylogeny . 2 vols, Springer-Verlag.
Tassy, P. & Shoshani, J. (1988). The Tethytheria : Elephants and their relatives In : Benton, M. J. (ed).
The phylogeny and classification of tetrapods. Syst. Assoc., sp. vol. 35B, Clarendon Press.
283-315.
Tiechert, C. (ed.). (1981). Treatise on Invertebrate Palaeolltology. Pt. F. (Coelenterata) . Geol. Soc. Amer.
& Univ. Kansas Press.
Tiechert, C. & Yochelson, E. L. (ed.). (1967). Essays in Palaeomology and Stratigraphy. sp. pub. 2, Univ.
Kansas Press.
Thompson D'A.W. (1942). On growth and form. Cambridge Univ. Press.
Uchio, T. (1960). Ecology of living benthonic foraminifera from San Diego area. Spec. Publ. Cush. Found.
Foram. Res. 5.
Valdiya, K. S. (J 969). Stromatolites of the Lesser Himalayan carbonate formations and Vindhya11s. Jour.
Geol. Soc. Ind., 10(1 ). 1-25.
Van Valen, L. (1965). Tree shrews, primates and fossils, Evolution, 19, 137-57.
Van Valen, L. & Sloan, R. L. ( 1965). The earliest primates. Science. 150, 743-45.
Venkatachala, B. S. & Maheswani, H. K. (eds.). The palaeobotanists. B . S. I., Lucknow.
Vermij, C. J. (J 983). Traces and trends of predation with special reference to bivalved animals. Palaeonl.,
26, 455-66.
Vishnu Mittre ( 1974) Qu t . . .
.· a ernary vegetatwn m northern regw11. In : Surange K R t l ( d ) Aspects
. and Apprisal of Indian Palaeobotany. B. S. I., Lucknow, 6 _ _' · · e ·a· e s. ·
57 64
V1shanathaia M N & A th R
, M· swa anarayana ao A.N. (1967). Algal stromatolites from Cuddapah Formation
near utssukota, Annanthapur dist., (A.P.) . /11d., Ind. Min. 8( 1-2) 62-65
Waechter, J. (1977). Prehistoric Man . Octopus Book Ltd. , .
--
••
oGRAPHY
BIBI)
(xi)
Wagner, C. W. ( 1964). Manual of larger forami11ifera . The Hague
,. A., Falk., 0., Smith, R. & Pickfood, M. ( 1983 ) "''h ·
wat ..er, .I . . . I e skull of Proco I ,(, .
I
and crama capacity. Nature, 305, 525_27. nsu aJnca11as; reconstruciion
p & Martin, A. W. (l 978). 011 the buoyancy oif th N .
Ward ' . . . e pearIy aut,Lus. lour. Exp. Zool., 205, 5-12
Washburn, S. L. (ed.). ( 1963). Classificatro,1 and human evolution. Aldine Puhl. .
Weishampel, D. B., Osmolska, H, Dodson, P. ( 1990). The Dinosauria. Univ. of California Press
Wells. J. W. ( 1957). Coral reef Geol. Soc. Amer. Mem., 67(2), 609. 31 . ·
Wells, J. W. (1963). Coral growth and geochronology Nature, 197,.948 _50 _
White, T. 0., Suwa, G. & Asfaw, B. (1994). Australopithecus ramidus, a 11ew species of early hominid
from Aramic, Ethiopia. Nature, 37 l, 306-12; 375, 88.
Whittaker, R.H. (1969). Concept of kingdom of organisms. Science. 163. 150-160.
Wolpoff, M. (1980). Palaeoanthropology. Alfred, A. Knopt Inc.
Woods, B. (1992). Origin and evolution of the genus Homo. Nature, 355 783-90.
woods; B.• Martin, L. & Andrews, P. (eds.). ( 1986). Major topics in primate a11d human evolution.
Cambridge Univ. Press.
Woods, H. (1946). Palaeontology. Cambridge Univ. Press.
Young, J. z. (1981 ). The life of vertebrates. Clarendon Press.
INDEX
PART I : PRINCIPLES OF PALAEONTOLOGY Bed 38
Ahiotic factors Benthic 31, 61
63-64 Berm 59
Accretion 25
Adaptation Binomial nomenclature 15-16
81-82
Adaptive radiation/divergence Biocoenosis 10. 57
85, 91, 92
Adaptive zones Biogenetic law/recapitulation 78
92
Addition Biologic associations
26
Age commensalism 62
38
Age of ammonites mutual ism 62
50
Age of amphibias parasitism 62
50, 77
Age of angiosperms Biogeography 30
49, 77
Age of birds Biogeographic/faunal province 31
51
Age of brachiopods Biome 34
49
Age of ferns Biostratigraphic zonation 51
77
Age of fishes 50, 77 Biostratigraphic units/zones 51-53
Age of gymnosperms 49, 77 acme zone 53
Age of invertebrates 77 assemblage zone 53
Age of mammals 51, 77 concurrent range zone 53
Age of reptiles 51, 77 peak zone 53
Age of trilobites 49 interval zone 53
Aggregated/Patchy distribution 69 taxon range zone 52
Ahermatypic corals 63 teil zone 53, 54
Alleles 79-80 Biostratigraphy 38
dominant 80 Biostratinomy 9
multiple 80 Biotic factors 64-65
recessive 80 Bioturbation 9
Allometric growth 29, 88 Blastocoel 22
Allopatric population 83 Body chamber 27
Allometry 29 Body plan 22-23
Amber 5 amerous 23
Anagenesis 91 metamerous 23
Analogy 67 pseudometamerous 23
Analogous structure 67, 78 oligomerous 23
Angara land 55 Breeding materials 7
Animalia 17 Burgess Shale Formation 49
Annual rings 55 Burrows and borings ·7
Archaeopteryx 51, 55
Asylum 41 Calcium Carbonate Compensation
Autecological study 65-69 Depth (CCCD) 3. 34
Autotrophs Calyx 26
64
Carbon compounds 45
Uarricrs· lo d'1spersal 6
32 Carbonization
l'alal·(UcnJ\\ I' (i)

I
iL _ _
_.,,.
t
PALAEONTOLOGY
(ii)
7 Distillized fossils 6
Cast fossils Divergent evolution 87-88
Catastrophist concept
40
83 Dollo's law 90
Character displacement 7
45-46 Domichnia
Chemical trace of life 70
80 Dominant species
Chromosome mutation 10
14 Drifted assemblage
Chronospecies
Chronostratigraphic units 38
38 Ecdysis 26
Chronostratigraphy 92
12-13 Ecological displacement
Cladism 60
13 Ecologic niche
Cladogram 92
12-21 Ecologic replacement
Classification of organisms
12-13 Ecology (d) 57
Phylogenetic
13-14 terminologies 60-62
taxonomic
13 Ediacara fauna 47
typomorphic/phenetic
22-23 Endemic 31
Cocelom
acoelomate 22 Environments 59-60
coelomate 23 marine 59
pseudocoelomate 22 non-marine 59
Columnals 26 mixed (Transitional) 59-60
Companion species 70 Eon 38
Conodonts Epiplankton/Pseudoplankton 31
Consumers Epoch 38
primary 64 Equus 56
secondary 64 Era 38
tertiary 64 Eury geographic 31
Convergent evolution 87-88 Euryhaline 64
Cope's/Daperet's law 89-90 Evolutionary convergence 78, 85
Coprolites 7 Exotic assemblage 10
Cocquina 2 Explosive evolution 91-92
Coriolis force 30 Extermination 41
Correlation by fossils IO, 53 External mould 6
Corridors 32 Extinction 41
Cosmopolitan 31
Coty losaurs 50 Faecal pallets 7
Cotype/Syntype 17 Faunal asylum 41
Cryptogam 17-18 Feeding traces 7
Cubichnia 7 Fig Tree Chert Formation 46
Cyanophytic algae 45 Filter bridge 32
Flagellate chambers 29
·Darwinism/Darwin's theory 73-74 Flagella 43
Death assemblage IO Fodinichnia 7
Density (Species) 69 Food ctiain (Food web) 65
Derived characters 12-13 Food habits 65
Diagenetic study 9 Food pyramid 65
Dimorphic species 14 Foot prints 7
Disjunct eurygeographic 31 Form genera 14
Directional selection 82
Dispersal
Form species 14
30-32 Formation 38
INDEX
(iii)
Fortuitous species 70 Growth of animals (Contd.)
Fossils : conditions of preservation 2-3 an isometric
definition 27
I isometric
imperfection 27
7, 9 mixed 26-27
kinds 1-2 moulting/ecdysis 26
modes of preservation 3-7 Gullet 43
taphonomic alterations 9-10 Gunflint Chert Flora 46
use 10-11
Fossil assemblage 10 Habitat 57
Fossil native Heredity 41
Fossil petrificata 1 Hermatypic/reef corals 36, 63
Fugichnia 7 Heterochronous fauna 41, 67
Functional morphology 67 Heterotrophs 64
Fungi 17 Heterozygous 79
Holotype 17
Gametes Homeomorphism 14, 67
heterogametes 80 Homologous structures 67, 78
isogametes 80 Homology 67, 78
Gastrol iths 7 Homonyms 16
Gene 74-75, 79 Homoplastic 67
dominant 75-76 Homozygous 79
recessive 75-76 Hybrid generation 75
Gene mutation 80 Hypertely 89-90
Gene pool 80 Hyracotherium 56
Generic name 15
Generitype 16 lchnofossils 7
Genetic drift 81 lchthyostega 55
Genetic equilibrium 80 Immigrants 31
Geographic barrier 31 Imprint fossils 6
Geographic environment Inadaptive orthogenesis 90
marine 59 lndegeneous assemblage 10
non-marine 59 Index/guide fossils 53
mixed 59-60 Indigenous 31
special 60 Indirect fossils 6-7
Geographic isolation 78, 82 In situ fossils 10
Geological time 38 Internal moulds 7
time units 38 Irreversibility 90
Gondwanaland 55 lsochemical alterations 6
Gondwana supercontinent 32 Isochronous 67
Grades of animal 22-23 Iterative evolution 92
diploblastic 22
primitive multicellular 22 Labyrinthodonts 50
triploblastic 22-23 Lamarkism 72-73
unicellular .• 22 Law of faunal succession 39, 40
Gradualistic evolution 87 Law of palingenesis 78
Group 16
38 Law of priority
Growth of animals 25-29 Law of superposition 39
accretion 25 Lebenspurrens 7
addition 17
26 Lectotype
PALAEONTOLOGY
~\I)
IO Organic evolution 71-94
Lifo assemhlage
61-62 basic concept 72
Life habits
Limiting factors 63 course 89-91
Lineage 87-88 evidences 76-79
Lithostratigraphy 38 factors 80-82
Lithostratigraphic/Rock units 38 ideas 71-72
Living fossils 88-89 modern views 79-80
pattern 87-88
Macroevolution 85 periodicity 91-94
Magnetic pole reversal 93 processes 82-87
Mammal-like reptiles 51, 77 theories 72-74
Mass extinction 91, 92-94 Organic hard part 5
Matrix 70 Orthogenesis 89
Megaevolution 87 Overprod.u ction 73-74
Megafossils I
Mendel ism 74-75 Paedomorph 91
Meroplanktons 31, 32 Paedomorphosis 90-91
Metazoa 22 Palaeobiogeography 30
Microevolution/Speciation 82-83 Palaeoecology (def.) 57
Microfossils 1-2 Palichnological data 68
Missing links 11, 77 Palingenesis 78
Monrea 17 Palynology I
Morphogenus 87
Panchronism 88-89
Morphological convergence 67
Parallel evolution 88
Morphospecies 14, 87
Mould fossil Parazoa 22
6
Moulting Paratypes 17
26
Multispecies ecology Pascichnia 7
70
Mutant Patchy distribution 69
79
Mutation 79, .80, 83 Permineralized fossils 6
Mutation theory Petrified fossil -6
79
Mutual ism 64 Petrified replacement fossils 6
Phanerogamia 17-20
Natural selection 41, 74, 81, 82 Phenetic classification 13
Nektons 32 Phyletic evolution 82
Nemesis 93 Phylogenetic classification 12, 13, 88
Neo-Darwinism 79 Phylogenetic evolution 72, 88
Neoteny 90-91 Phylogenetic tree 13
Neotype Phylogeny
17 72
Nip Planktons
59 31
Nomenclature Plantae
12 17
Non-adaptive characters Planula
81 45
Non-adaptive genes Platyceratid gastropods
82 65
Norm of reaction Plastotype
27 17
Numerical taxonomy
13 Plesiotype 17
Point mutation 82
Ontogeny
Oon's cloud 25, 72 Polyphyletic origin 51
93 Polyploidy 83. 91
Optically active/inactive
Optimum temperature 46 Polymorphic species 14
63 Pre-adaptation 82
/VDEX
J8
.e
45
74
15
16
74
32
. -- ~..,,
83
65, 70
16
Quanrum speciation
alrerations 9-10
Random distribution 69 nm) 9
Rate of growth 2 - 9 Tu n Range 2.r.lne 52
an isometric 7-29 Ta., n my 12
isometric - 9 Jax n mi categoies 13-14
Rate of evolution 88 llleory of cataclysm 71
bradytelic &8-89 lbeol)' of germplasrn 73
horotelic 88 lbeory of natural selection/organic 40 41. 73-74
tachytalic 88 Theory of spontaneous origin of life 71
Recombination 80-81 Thanatocoenosis 10
Recryst.all i zed fossils 6 lime rock unit 38
Recurrent fauna 41 Trace fossils 7
Reducers/decomposers/Trans formers 65 Trade wind 30
Regressive/Retrogressive evolution 90 T' pe 16-17
Remain assemblage 10 corypdsyntypc: 17
Repacernent fossil 6 generitypc: 16
Reproductive isolation 83 holotypc: 17
Reptile-like mammals 77 lectOlype 17
Resin 5 neotype . 17
Rules of scaling 29 paratypc 17
plasiotypc: 17
Secondary/hypotype 16 plastotype 17
Sex chromosomes 76 primary/proterotypc: 16, 17
Sexual dimorphism 76 secondary/hypotype 16, 17
Sewal wright's effect 81 topotypc: 17
Simplification 90 Typomorphic classification 13
Species 14-17
concept 14 Umbo 25
naming 14-15 Uniform distribution 69
type specimen 16-17
Species diversity 34 Variation 41, 74
Species frequency 70 Vestigial/Rudimentary structures 78
Specifidtrivial name 15
Stabilizing selection 82 Williston·s law 90
Stage 39
Stem reptile 50 Zone 38
Stenographic 31 Zone fossils 53
Stenohaline 64 Zooxanthellae 36, 64
\ ~
y
p RT II : 11'" ), I ~
AulOZOOecla
hell 126 151
Au tozooid
151 A 'lCU Laria
145 AxiaJ Jobe
IO I
Axial segment.s/furrov. s
183 Ax iaJ structure
Am i 217 219
Am l~ I a.reas/Ambs 219, 223, 236
Amm ni ticonc Bactriticone
192 Baculicone
Ammonitte suture 181 , 193
Amoebocytes/Amoeboid cell Barriet" reef
97
Am phidctic Basals
151
Am phithyrid Basal chambet-
126
Anal p ramid BasaJ plate disc l -~
235
Anal lube Beak 125, I'
235
Anapbycus Beak ridge Lo
180
Ancestrula Belemnoid I ,
137
An isoma arian Biramous appendages
155 - J
An n I iphonatc
An n lu layer
18 1
Blastoids
Body whorl
-~
16
175
An mphalu Border 199
169
Antennae Brachia 1-
198
Anlef Brachials 2]5. 236
137
Antho7..oa Brachia] canal Ll l
103
Aperture BrachiaVDorsal val I! 1-
171, 179
Apertural diameter Brachidium 120, 130. 13I
143
Apen_uraJ pine
237
Brac hi phore 1:
Apex Brachio poda Shell 1.0- 1:
10 3, 164, 175
ApicaJ/SpiraVPteuraJ angle cl ifi a1i n l ~I
171
Apical system dimen ion and oriental.i on 1 29
217
Aplacophoran
140, 145 ecology 1.t'
Apodeme geological hi tory 135- Ll6
203
Appendifer homeomorphi m 135
Aptycus 203 13
morphology 12.S- 127. 129- ~
180 J.
Archaeocyathid shell structure
Archetypal 117 1 35
stratigraphic importance
210 179
Brevicone
INDEX (vii)
Bristle 137 Ceratobactriticonc 192

Brosh 125 Ceriantipatharia 112
Bryoroa 137-139 Chaetognath 211
ecology 137, 139 Chain colony 107
general features 137 Chalky layer 181
geological history 139 Cheek 199
morphology 137 fixigena/fixed cheek 199
Budding 113 librigena/frce cheek 199
Buttress 155 Cheilostomata 139
Byssal notch 151, 157 Chela 198
Byssal thread 161 Chilidial plates 126
Bys.sate forms 161 Chilidium 126
Choanocytes/Collar eel Is 97
Cadicone 179 Chondrophore 155
Caecum/Caecae 120, 125 Cnidaria JOI, 103
Calcareous sponges 99 general character IOI
Callus 172 subdivisions 101-103
Calyx/Cal ice I05, 235, 236 Coelenterata 101
Camera/Chamber 180 Coenosteum 119
Cameral fluid 186, 187, 189 Collar cells 97
Carapace 198 Colonial/Compound coral 103, 105, 107
Cardelles 137 Columella 111, 169
CardinaVHinge area 151 Columellar lip 171
Cardinal (Lateral) extremity 126 Columnals 235
Cardinal/Hinge plate 129 Connecting rings 181
Cardinal process 130 Connecting suture 201
Cardinal/Hinge teeth 151 Common canal 237
CardinaJia 129 Conchioline 148
Carina/Keel and sulcus 183 Conispiral (Trochospiral) 169, 179
Caudal spine 205 Conjugate pores 221
Cephalon/Head shield 199 Connecting rings 181
Cephalic spine 205 Conularids 119
Cephalic suture 201 Convolute 169, 180
Cephalopoda (Shell) 175-194 Coral Anthozoa 103-117
causes of extinction (Ammonoids) 193 classification 112
classification 183, 185-186 ecology I 1S-116
coiling (Shell) 179, 180 evolution I 13, 115
general features 175 geological history I 16
geological history 194 reefs 115, 116
evolution (Ammonoid) 189-193 reproduction and growth 113
microstructure (Shell) 175 skeletal morphology 103-112
morphology (Shell) 175-186 stratigraphic use 116-117
morphological specialization (Shell) 189 Corallite 103, 105
naming 175 Corallum 103
ornamentation (Shell) 183 Core cells 125
shape (Shell) 179 Corona 219
stratigraphic use 194 Coronal plates 219
Cephalothorax
Ccratitic suture
198 Coxa 203
181, 193 Cranidium 199
PALAEONTOLOGY
(viii)
216, 236 Echinoid 216-236
'Crinoids
237 classification 228-229
general morphology
239 ecology 229. 231 , 233
geological history
237 functional morphology 226-228
Crossing canal
235 morphology 216-226
Crown
Crural base 129 origin evolution and geological
Crural lamellae 130 history 233
Crural process 130 orientation and dimensions 223
· 130 ornamental features 221-222
Cruralium
Crus/Crura 129 shape/symmetry 216, 217
Cruziana 208 stratigraphic importance 235
·cryptomphalus 169 Ectoderm 101
Cubichnia 208 Endocones 181
Cryptostoma 139 Endobyssate 161
Cup 117 Endocyclic/Regularia 216, 228
Cyclostoma 139 Endoderm IOI
Cyrenoid 153 Endopodite 198,203
Cyrtocone 179 Endopunctate she II 120
Cyrtochoanitic 181 Epibyssate 161
Cystid 137 Epidermis 211
Epirostrum 183
Deductor muscle scar 130-131 Epitheca 105
pelthyrial cavity 129 Epithelial cells/Pinacocytes 97
Delthyrium 126 Epithyrid . 126
Deltidial plates 126 · Equiangular/Logarithmic spire 144
.Deltidium 126 Evolute 169, 180
Deltoids 236 Evolution of ammonoidea
Dendri.tic suture 181 ancestry 189, 191
Dental plates/teeth t'29 phylogeny 191
Dental sockets 129 phenotypic trends 191
Dextral 169 Exocycl ic/lrregularia 216, 228
Diaphragm 137 Exhalant siphon 173
Di!=h<;>graptid fauna 239 Exopodite 198, 203
Dicyclic 235 Exuvia 207
Dimayarian 155 Eye 199
Dip/sutural angle 171
Dipleurula 211 Facial suture 199,201
Diplograptid fauna 239 Fasciculated colony 107
Dissepiment pl, 117 Fasciole 222. 228
Dissepimentarium 111 Fibre fascicles 109
·Dollo's Jaw 192 Flagellate chambers 97
Oorsum 180 Floscellae 222
Double Crossing Canal 237 Foramen 126
Doublure 201 Fossula 109
Fringing reef 115
Echinodermata 211-216 Fusella 237
classifications 216
151
general characters 211,216 Gape
INDEX (ix)
Gastropoda (Shell) I <,4 . 74 I lydrozoa IOI
classi Ii cation 72 1lyperntrophic 169
coiling (Shell) Hypnnome/funncl 175
composition (Shell) 64 Hypononiic Ninus 180
ecology 172- 73 Hypostome 201
evolution & geological history 74 Hypostomal suture 201
general features 64 Hypostracum 148
morphology (Shell) I 64- 72 Hypothyrid 126
morphological analysis (Shell form) 171- 74
ornamentation (Shell) 72 Impcrforatcd shell 169
shape (Shell) 67 Impunctatc shell 120
Gcnal angle 99 Inarticulata 120
Genal spine 199, 205 Indian fossils 241-246
Generating curve 141 Palaeozoic . 241 -243
Genital plates/pores 217 Mesozoic 243-245
Gilatinosa 100 Cenozoic 245-246
Gill branch 203 Inductura 172
Glabella 199 Incurrent canals 97
Goniatitic suture 181, 193 Inhalant siphon 172, 173
Graptolites 237-240 Inner lip 171
biological affinity 239 lntcrambulacral areas/Interambs 219
ecology 239 lntcrarea/Hinge area 125
general morphology 237 Interarea angle 126
geological history & evolution 239 Internal ligaments/Resilium 153
Guard/Rostrum 183 Internodals 235
Gutter 17) lnterporiferous zone 221
Gullet I03 lntervallum 117
Gyrocone 179 Involute 169, 180
Gyroceraticone 192 Isochronous homeomorphism 135
lsomayarian 155
Hemiomphalus )69 Isopygus 205
Hermatypic coral 115 Iterative evolution 191
Heterocorall ia 113
Heterochronous homeomorphism 135 Jugum 130
Heteromorphs 192
Heteropygus 205 Labrum 222
Hcterostroph ic 169 Lamellibranchia 148
Heterozooid 137 Lancet plate 236
Hexacoral la/Scleracti nia I 07, 11 I , I 13 Lateral lobe 198
Hinge axis 125 Lateral teeth 153
Hinge line 125, 151 Left valve 148
Hinge plate 129, 151 Leucon/Rhagon 97
Hinge teeth 129 Ligament 151
Holaspis 207· Ligamcntal pit/groove 151
Holdfast Line of Commissure 127, 151
117, 235
Holochoanitic Living/Body chamber 175, 180
181
Holostomatous Living fossil 136
Hornylaycr
171
Lituiticone 179
181
Hydrorhi1.ac 181
119 Lobe
Palac:(Geo)Wp_ 72
-
PALAEONTOLOGY
L ·ulae 117 Nautilitic suture 181, 193
Loculi 117 Neck (Gastropod) 171
L phophore/Bm ·hia 120, 130 Neck furrow 199
Lu inoid 153 Nimatocyst/Cnidoblast 101
lumen 235 Nodals 235
Nonstrophic shell 125
Macropygus 205 Notothyrium 126
Madreporite 219, 226 Nuda 100
Mamdon 119
Mandibks Tril bites) 198 Occipital segment/furrow 199
Mantle 140 Occipital node 199
Mantle cavity 140 Octocoralla 112
Marginal furrow 199 Ocular plates/pores 217
Marginal notch/Lunule 222 Oculo-genital ring/system 217
Marginal spines 205 Operculum 137, 17i
Massive colony 105, 107 Opisthocline 148
Maxillae 198 Opisthodetic 151
Median fold/ridge 127 Opisthogyral 148
Median septum 129, 237 Orifice 137
Median sulcus 127 Orthoceratitic suture 181, 186
Median suture 201 Orthocline 148
Medusa IOI Orthochoanitic 181
Meraspis 207 Orthocone 179
Mesenchyme 100 Orthogyral 148
Mesenteries IOI, 103 Orthostrophic 169
Mesogloea 101 Osculum 97
Mesozooecial cavity 137 Osphradia 172
Mesothyrid 126 Ostia 97
Metameres/somites 198 Ostracum 144, 148, 175
Metamorphosis 198 Outer lip 171
Micropygus 203 Outer ramus/feleopodite 203
Metatheca 237 Ovicell 137
Mimocone 192 Oxycone 179
Mollusca 140-147
fundamental organization 140 Palaeontological clock 117
effect of predation 147 Palaeontological relay 191
growth and shape (Shell) 143-144 Pali/Palus 111
origin and phylogeny 144-145 Palintrope 126
subdivisions 140, 143 Pillars 119
Monocyclic 235 Pallial line 155
Monograptid fauna 239 Pallial sinus 155, 157
Monomayarian 155 Pallial markings )31
Monoplacophorans 145 Palpebral lobe 201
Moulting/Ecdysis 198 Paragaster 97, 117
Multispiral 167 Parazoans 97
Mural pores
105 Pariet/septum 117
Muscle Scar 171
155 Parietaly lip
Myophore 167
130, 155 Paucispiral
Pedicellariae 222
Nacreous layer 120
175 Pedicle/Peduncle
I DEX .xi)
f\.•d id c orx: ning Prolotheca 103. 237
f\,"ll id • \.'.ntral \'al vc 126 Protocond 1 143, 175. I XO
J\,·ll" YI h Bi alvia (S hdl ) 120 Pscudupunclalc shell 125
·lassiticati n 148- 163
Pscudoscpla 112
d ·ntition/tccth I ath.:rn 158- 159 Pum:lae 120
Jimcnsi n and orientati on (Shell 153 Pygidium 203, 205
~ol gy ) 157
159, 16 1-162
g~ncral k;1turcs 148 Racial senesce nce 192
geological history 162 Radials 235, 236
morph log} hell ) 148-158 Radial canal 226
148 Radula · 140. 164
naming
155 Ramp and shelf 171
rnamentation (Shell)
Reclined 237
stratigraphic value 162
Rejuvenescence ti 3
Pt:llicle 181
187
183 Relative Strength Index
Pen 208
Perl rated shell 169 Repichnia
153
120, 144, 148, 164 Resilium
J>eriostr3CUITI 155
221 Resilifer 181
Peripodium
217 Retrosiphonate 237
Periproct/Anus
171, 217 Rhabdosomc 239
Peristome Rhabdopleura
223 97
Perignathic girdle Rhagon
126 226
Permesothyrid Ring canal
237 148
Pendent 169 Right valve
Phaneromphalus 201. 23 1
175, 180, 183 Rostrum 201
Phragmocone 231 Rostral suture 161
Pillar 169, 179 Rudistids 126
Planispiral 223 Rugae 208
Plastron 179 Rusophycus
Platycone 198 181
Pleurit.e 201 Saddle 237
Pleuron 203 Scandent 198
Pleural lobes/furrow 203 Sclerite 100
Podomere IOI Sclerocytes/Spongocytes 222
Polyp 191, 195
Scrobicular granules IOI
Polyphletic 140, 145 Scyphozoa 137
Polyplacophorans 101 Sea mosses 120
Polypoid 137 Secondary layer 171
Polyzoa 137 Sclenizone/slit hand 137
Poster 221 Setue/Scrn 103. 105. 107. 137. 180. 183
Poriferous zone 97 Septa (Septum)
Porocytes 120 Septa of Coral 109
Primary layer confluent 109
183
Pro-ostraeum cntosepta 109
181
Prosiphonate cxoscpta 109
148
Prosocline cxsert septa 109
148
Prosogyral insert septa 107
97
Prosopyles mnjor septn 107
207
Protaspis minor septa
208
Protichnites
PALAEONTOLOGY
(xii)
Strophic shell 125
Septa of Coral (Contd.) 107 Suture 180
primary/protosepta Sycon 97
I07
sccondary/mctasepta Synapticulae 109, 117
109
microstructure
109
Scptal surface ornamentations Tabcllae 112
187. 189 103, 112, 117, 119
Septal fluting 181 Tabulae
Septal neck Tabularium 112
181
Septal suture Tabulata 107, 112
I 37, 203
Sc tac/bristle Taleolae 125
140, 148
Shell Telson 205
171
Shoulder (Ga~tropod) Teeth and sockets 148, 151
237
Sicula Teeth pattern (Pelecypoda) 153
mctasicula 237
desmodont 153
prosicula 237
edentulus/palaeconcha 153
Silicious sponges 99
dysodont 153
Sinistral
169
heterodont 153
171
Sinus 153
171 isodont
Siphonal notch/canal 153
140 pachydont
Siphonal system 153
171 schizodont
Siphonostomatous 153
181, 183, 186-187 taxodont
Siphuncle 235
181 Tegmen
Siphuncle tube 203
164, 171 Telopodite
Slit band/Selenizone 107, I 12, 113
103 Tetracoralla/Rugosa
Solitary/Simple coral 198
Sphaerocone 179 Tergite
Theca I03, I 05, 237
Spicules 97, 100
112 Thecarium 105
Spider's web 198, 20 I, 205
130 Thorax
Spiralia 203
171 Thoracic segments
Spiral suture 164
167 Torsion (Gastropod)
Spire JOO, 109
Spired echinoid 231 Trabeculae
Trepostoma 137
Spiracles 236
129 Trilobites 198-210
Spondylium 205, 207
97, 100 classification
Spongin 207. 209
97 ecdysis and ontogeny
Spongocoel 208
97-100 ecology
Sponge/Porifera 210
classification 100 geological history
general characters 198
geological history 100
morphological features · 199
morphology 97
ornamental features 205
skeletons 100
97 stratigraphic use 208, 210
systematic position 208
Squamose/Imbricate structure 127 trace fossils
164
Stalk/Stem 235 Trochophore
226
Sternite 198 Tube feet
179
Stipes 237 Turrilicone
144
Stolon 137 Turbellarian flatworm
Stone canal 226 169. 180
Stony bryozoa 137 Umbilicus 180
Stromatoporoid 119 Umbilical plug
/'Vi)£ .
(xiii)
Umhi ·al should~r 180 Apatosaurus 333
l mbo 125, 148 Apes 353. 355
Arandraspis 293
, ·atn• attarhm~nt 130- 13 1, 155 Archaic Homo sapiens 371
Varix 172 Archaegosa u rus 303
\t'nlt'r 180 Archaeohippus 342
V1;.~:irul l.f tissepiment II 1 Archaeopteryx 291, 311
\ ibrtK'UIU 137 Ardipithecus ramidus 364
\'irgt·lla/N~ma 237 315
Armadillos
Armoured fishes 293
Wat~r vascular system 211, 226 Artiodactyls 319, 320, 381
Whorl 167. 180 Astragalus 269
Wh" rl height 180 Astrapotherids 319
Wh rl pmfil • 167 Astraspis 293
Wh )r\ translation 144 Australopithecine stage 362. 363-367
fossil record 363-364
Zoantharia 112 fossil sites 365, 367
Zoaria 137 lifeways 364-366
ZNlrium 137 morphology 366
2(.)(XC'iun Cystid 137 Australopithecus 323. 359, 362, 364
z . 137, 237 A. afare11sis 359, 364
A. africa11as 364
PA.Rf III : VERTEBRATE F~ILS A. a11ame11sis 364
A. boisei 364
A :-anchodians (Spiny sharks) 295 A. robustus 361, 364
Acamh Mega 303
A~lian culture 370, 372 Barapasaurus 328
Actin p<erygii (Ray-fin fish) 297 Baurusuchus 308
.{ rl basileus 313 Beluchitherium 319
Aex_\'pl pirhecus 353, 362 Aves (Birds) 252, 254-255, 259, 263, 269,280,
Age of mammoth 348 290-291, 311
Agnacha.s 293 Biso11 320
A asaurws 309, 328, 333 Bony fishes 298
•..\mbt- d " 348 Bony girdles 269
Amb{)I)(XJS 319 Bos 320
298 Bovids 320, 382
Amni e ~gs 289 Brachiosaurus 311, :us
Ampttibia 252, 259, 263, 287, 300-304 Bradidonts 297
dassification 300 Bramatherium 381
Amphicyon 317 Bra11chiosaurus 303
Ampltipirh~nu 362 Brontosaurus 311, 333
~ 305-306 Brontotherids 319
342 Bro11totherium 319
327, 334 Bronze age 375
379 Bucapra 320
382 Buettnaria 303
320
320 Calcaneum 269
323 Came/us 320
PALAEONTOLOGY
(xiv) or, luvallois t • ·hniqu ·

372
.ot'II ·nl stf'II ·tun.:s (M11111111 tils)/hnrn 278-279
Camptosnurid 319
CamptosauruJ 'oryplwdo,i 291 , 305, 308
Canids '01ylosu11r
315, 317
Cannon bone.: Cr ·odonts
Capra ro-Mag nnn 111 1111
:n3
286. 297. 299, 303
Captorhinomorph 291 n ,ssopt cry gia ns
313
Carinatac '.\ 17. '.\81 ·m.\'filo rt i a
Carnivoras ryptodircs
306
'.\24, '.\29- '.n I. '.\'.\'.\ 293
Carnosaur ydostonwta
Catastrophist ymhospo,idyl 11s 306
extra-tc.:rrcstrial ,node
:n6 317
'.\'.\6 ynodic:tis
vulcanological model Cy11n~11atli11s 307
'.\74
Cave arts
32'.\, 353
Ceboids Dapedius 298
'.\53
Cebupirhecia Deccan intcrtrappcnn 376
263
Ccntrum articulation Deltat/1,1ricli 1.1111 317
acoeloas 263
263 Dcnwa formation 376
amphicoeleus 115
26'.\ Dcrmoptcrus
heterocoeleus 320
opisthocoelous 263 Desmostylin
Dharmararn formation 376
procoelous 263
293 Diacodexis 320
Cepha/opsis 305
Ceratodus 299 Diaclectes
327, 329, 333 Diadcctomorph 305
Ceratopsians
323, 353 Diapsids 308-311
Cercopithecids
320 Dinstcma 340
Cervus
Cetaceans 317 Dibelodon 348
Cetiosaurus 333 Dichobunoids 320
Chalicotheres 319 Dicy11odo,i 307
Cheirolopsis 298, 299 Digital posture (Mammals) 269-271
Chelonias 306 digitigrade 269
Chiropteras 315 plantigrade 269
Chondrichthyes 295 unguligradc 271
Chondrostean 297, 298 Digital tips (Tetrapods) 280
Chordata 251 claws 280
diagnostic feature of major groups 253 hoofs 280
subdivisions 251 nails 280
Cladognathus 293 nail groove 280
C/adosel/ache 295, 297 subunguis 280
Classical neanderthals 370 unguis 280
Climatius 295 Dimettvdo11 307
Coccosteus 295 Dinichthyes 295
Coelacanth 300 Dinocephalians 307
Coelacanthids 299 Dinosaurus 309, 324-337
Coelophysis 309, 327, 333 ancestry 127-328
Coelurosaurs 324, 329, 333 causes of extinction 334-337
Condylarths 317,340 classification 324-327
Condonts 293 111-332
control of body temperature
Copper age 375 egg fossils
332
L
INDEX (xv)
Dinosm1rn s (Co ntd.) £ 11,y ops .101
fo ss il s il ·s 32 8 E,y tli ros11ch11.\· .109
general charnclcrs 333 Euperkeria 317
gcolngical hi story 333-334 Eury apsi ds 306
lifcways 329-331 Eurymylu.\· 317
struc ture fo r defe nce 331 Eusuchians 308
inosaur park 379 Eustlie11optero11 287. 299
Dinot he res 347, 382 Evolution o f j aw 286
Di11otheri11111 347 Evolution of limb bon es 271
Diplcuru la 284 archiptcryg iam 271
Diploca11/11s 304 icthyoptery g iam 271
Diplodocus 311 , 333 che iropte rygiam 271
Dipno ids 297, 299
Dipterus 299 Fabrosaurus 327, 3'.B
Docodo nts 315 Feather (Birds) 280
Dryolestids 313 contour feathers 280
Dryopitheci ne 381 down feat he rs 280 .
Dryopitherns 323, 355, 362, 364 filoplume 280
Duck-billed dinosaurus 327, 331 rachis 280
Dugong 323 vane 280
Felid 3 17
Ecolagic replace ment 300 Fish (Pisces) 251, 253, 267-269, 277-279
Edaphosaurs 307 Fish fins 279
Edentate 315 acetabular region 279
Elasmosaurus 306 besipterygium - 279
Elephantids 347 ceratotrichia 279
Elephantoids 347 coracoid 279
Elephants 323, 345-349 isopubic bar 279
ancestral form 346 mesopterygium 279
major traits 345 protoptery gi um 279
phylogenetic history 347-349 pterygoniophores/somactids 279
trends of evolution 346 Fish scale 277-279
Elephas 323, 383 cosmine 277
Elephas maximus 345, 34$ cosmoid 277
Eleutheromis 29l cteni 279
Epihippus 341 ctenoid 279
Elongatoo/ithus 333, 379 cycloid 279
Emblomeres 303 dentine 277
Em bri thopods 320 ganoid 277
Endothermic animals 332 ganoin 277
Entclodonts 320 isopedine 277
Eogyrinus 303 leptoid 277
Eohippus 340, 341 placoid 279
Eoraptor 327 Fish tail 267
Eosuchian 308 diphyccrcal 267
Epihippus 341 heterocercal 267
Equidac/Equids 320, 338 homoccrcal
Equus 267
320,341,342,381,382 protocercal
267
(xvi) PALAEONTOLOGY
Fissipcds 317 Homo erect us 323. 359, 361 , 364, 367 , 369,
Fore limb 269 370. 371. 372
digits 269 Homo l,abilis 323, 359, 364. 367
humerus 269 H. /ieidelberge11sis 370
radius 269 H . 11ec11ulertlw/e11sis :no
ulna 269 H. sapie11s 323, 359, 369, 370, 371
H. sapiens 11ea11dert/w/e11sis :no
Ga/ago 353 H. sapie11s mdesie11sis 371
Gangamopteris hed 376 H. sapie11s sapie11s 370, 374
Giga11topithecus 355. 362, 363 H. sapie11s soloensis 371
G. bilaspurr11sis 363 Hoofed mammals 317, 340
Giga111osaurus 311 Horse 319, 338-345
Giraffa 320 adaptation of modern horse 338-340
Giraffakt:ryx :no. 38 J, 382 basic ancestral characters 340
Giraffids 320 phylogenetic history 341-342
Glorified reptiles 290 progressive trends of evolution 341
Glypodonts 315 Hybodonts 297
Gomphotheres (Trilophodonlids) 347 Hy bodus 297
Gomphotherium 347 Hydaspitherium :no, 381
Gondwanaland 379 Hyenodonts 317
Gondwana rock 376 Hylonomus 305
Gondwana vertebrates of India 376-379 Hypohippus 342
Gorgosaurus 333 Hyracoids 320
Gradualistic model 336 Hyracotherium 319, 340, 341
Ground sloth 315
Ice age man 369,371
Habiline stage 367 lclhyosaurs 317
Ha/itherium 323 lchthyosaurus 306
Hardrosaur 333 /chthyostega 287, 303
Hardrosaurus 328, 334 Ichthyostegids 287, 303
Hemicyo11 317 Ictidosaur 308
Herrarasaurus 327 lguanodon 328, 331, 334
Hesperocyon 317 /ndractos 317
Hesperomis 311 lndraloris 353
HnJJerosuchus 309,327 Insectivoras 315, 323
Heterodontosaurus 327, 333 Intercentrum 303
Hind limb 269
digits 269 Janess 297
femur 269 Jamoytius 293
fibula 269 lawless vertebrates (Agnathas) 293, 294
tibia 269 Jaw suspension 258, 259
Hippario11 342, 381, 382 amphistrylic 258
Hipparion fauna 342 autodiastylic 258
Hippidium 342 nutostylic 258
Hippopotamus 320, 383, 384 craniostylic 258
Holostei 297 hyostylic 258
Homeothcrmy 332 slreptost y Iic 259
Hominids 323, 382
Hominoidea (Hominoids) 323, 361 Key11iapi.fliec11s 355, 362, 364
L
1NDEX
(xvii)
K ta forinntion 376 Man (evolution)
0 'd 313 J5 I-J75
Ku..:hncoitu:r'. aocestry
Kue/11 ,eothe,., ""' 313 :\55
kins of
35:\-355
stages of evolution 361 -375
Lahyrinthodonts 286,287,290,300,303
systematic position 351-353
Lagomorphs 315, 317 trends of evolution 355-361
Lambeosa11ms 333 Marsupialia
315
Lameta beds 376 Mastodon americanas 348
Laplatosa11111s 328, 379 Mastodon ts
347
Latimeria 300 Median tin (Fish) 251
Lemurs 353 Megaloolithu.\· 333, 379
Lemuria 379 Megalosaurus 328, 333
upidosi~ll 299 Megatlterium 315
Lepisosteus 298 Megaz.ostrodo11 313
Lepospondyls 300, 303 Megaladapis 353
Leptolepis 298 Melanosoums 333
Limb 251 Merychippus 342, 381
Limb hones (Tetrapods) 269 Merycopotamus 320
fore limb 269 Mesopitheccus 323
hind limb 269 Mesoltippus 341
evolution 271 Mesosuchians 308
Linmopithecus 362 Mesosaurus 306
Linmoselis 305 Microdon 298
Lissamphibias 300, 304 Miohippus 341-342
Litopteras 319 Miomastodon 348
l)ving fossil 300 Missing link 291 , 364
Lophophorate 284 Mobile arts 374
Loris Moeritheres 323, 346
353
Lorises Moeritherium 323, 346
353
Loxodonta Monkeys 353
323
Loxodo11ta aifricanas Monoclonius 334
345, 348
Lower/Early palaeolithic culture Monotremes 315
370
Lucy Morganucodon 313
364
l ystrosaurus Morganucodonts 313
309, 379
Moropus 319
Macacus Mosc:hops 307
323 Mousterian culture 372
Macaque
353 Myxi11e 293
Macropoma
300
Macrotherium
319 Neanderthal cemetery 372
Maiasauras
333 Neanderthal man 370-~ \372
Maleri formation
378 Negative allometry ~43
Mammalia (Mammals) 252-253, 254, 262, 273. Neoceratodus 299
275, 279, 280, 345 Neolithic culture 375
classification 252-253, 312-313 New world monkeys (Cehoils) 353
evolution Non-ruminant 320
292
teeth
273, 275 Notharctus 353
Mammal-like reptile Nothosaurus 306
Mamn1011,,,. 294, 306
348-349 Noloungulata 319
Palae(Gco)Wp. 71
PALAEONTOLOGY
353 Pelvic girdle

25 l, 269
Old world monkey
364, 367, 369 acetabulum 269
Olduv:ii gorge
367. 370 ilium 269
Oldowan tools
307 ischium 269
Ophiacodon
Ophiduperorr 303 pubis 269
Oreodont 320 tetra-radiate 269
Ornitholestes 333 triradiate 269
Omithomimus 33 l, 333 Pelycosaur 307
Ornilhopods 327, 333 Pennipeds 317
309, 333 Perissodacty ls 319, 338, 381
Ornithischians
Ornithosuchus 309, 324, 327 Petralocosaurus 308
Orohipp11s 341 Petromyza11 (Lamprey) 293
Onhogenesis 338 Phenacodus 319
Orycteropus (Aardvark) 319 Phiomia 347
Osteichthyes 297 Pholidonti~s 315
Osteolepis 287, 299 Phytosaurs 308
Ostracoderms 293 Pinnipeds 317
Ostrich dinosaurs 329 Pisanosaurus 327
Oxyaenids 317 Pithecanthropine stage 369-370
lifeways and culture 369
Pachycephalosaurus 329, 331, 334 major traits 369
PaJaeodonts 320 stratigraphic distribution 369
PaJaeolilhic culture 370, 372-374 Pithecanthropus erectus (Java man) 369
PaJaeomastodon 347 Placoderms 295
PaJaeoniscoida 297, 298 Placodonts 306
Palaeoniscus 298 Placodus 306
Palaeonodon ts 315 Plateosaurs 333
Palaeopithecus 323 Plateosaurus 333
Pa.Jaeoth yris 305 Platybeledon 348
Palatoquadrale 253 Plesiosaurus 306
Panduangia 362. Pleurocanth shark 297
Pangolin (Pholicodontid) 315 Pieurodire 306
Panchet formation 379 Pleurocentrum 303
Pantolambda 319 Pliohippus 342
Panthotherids 313 Pliomastodon 348
Papio (Baboon) 353 Polyphyletic 292
Parahippus 342 Pongids (Apes) 323
Parallel evolution 300 Positive allometry 343
Paramys 317 Power grip 364
Paranthropus 366 Pre-man stage 362-363
Parapithecus 323, 353, 362 Precision grip 364
Parasauroloph 334 Primates 351-352
Pavement teeth 297 adaptation for arboreal life 350-351
Pectoral girdle 251, 269 insectivora-primate transition 350
coracoid 269 Proboscideans 319, 381
glenoid cavity 269 Probosuchus 308
precoracoid 269 Proconsul 323, 355, 362
scapula 269 Procolopho11 306
suprascapula 269 Procolophonids 306
JtvDEX
(xix)
,oga11 ochelys 306 Ruminants
P sive neanderthals 370 320
progres
J>rolacerta 308 Salamanders
p,op/iopithecus 323, 353, 362 Sapient stage 304
J>rosauropods 327, 333 370-375
appearance of H. sapie11s sapie11s 372
prosimians 323 archaic H. sapie11s
prosaqualodo11 317 371
cultural development 374-375
Proroavis 311 fossil sites
pro1ocet11s 317 373
neanderthal man 371-372
proroceratops 328, 334
neandcrthals & H. sapiens 370-371
proropterus 299 Saurischians 309, 324-327, 333
Protosaurs 306, 309
Sauropods 309, 327. 328, :n I, 333
Protosiren 323 Sauropodomorpha 327
Protosuchians 308 Sea cows 320
Prera11odon 309 Sebesuchians 308
Pterosaurs (Flying reptiles) 290 Sebescus 308
Pyrotherids 319 Semionotus 298
Senmopithecus 353
Rabbits 317 Sercopterygii 297
Rachi tome 303 Serride11tim,s 347
Racial senility 337 Seymouria 292, 305
Ramapithecus 362 Shark 295, 297
Rat fish 297 Simians 325
Ratitae (Ratites) 291, 311 Sinanthropus peki11e11sis (Peking man) 369
Rays 295, 297 Si11ocodo11 313
Rechnisaurus 379 Sirenians 320
Redfieldia 298 Sivapithecus 323, 353, 362
Reptiles 252, 254, 259, 263, 287-290, 304-311 · Siwalik Group 376, 379
amniote egg 304 Siwalik hills 382
evolution 287-290 Siwalik mammals 379-386
subdivisions 252, 304-305 cause of disappearance 381-382
geological history 305-311 climatic conditions 382
Rhachitome 303 evolution and migration 379, 381
Rhamphorhynchus 309 surrounding hahitates 382
Rhinoceratids 319 Skates 295, 297
Rhinoceros 319 Skull (Vertebrate) 249, 256-261
Rhipidistians (Rhizodont) 299 auditory meatus 261
Rhizodonts 299 branchial arches 256
Rhynchocephalians 308 chondrocranium 256
:hynchosaurs 308 dermal bones 258
?bYncliotherium . 348 dermatocranium 256
R1 s (Tetrapods) 267 embryonic components 256
capitulurn 267 fcncstra ovalis 259
diapophysis 267 foramen magnum 256
parapophysis 267 gill arches 256
sternum 267 hyoid arch 256
tubcrculum 267 hypophysial fcncstra 256
Un .
R.n,1 cinate process 267 internasal septum 261
""cnts
315, 317 jaw suspension 258
(xx) PALAEONTOLOGY
Skull (Vertebrate) (Co11td.) Symmetrodonts 313

madibular arches 256 Sy naps ids 292, 306-308
neurocranium 256 Sync/10110/oph us 347, 348
olfactory capsule 256
optic capsule 256 Taeniodont 315
otic capsule 256 Tail 249
parachordals 256 tail fin 251
precordals 256 Tapirs 319
splanchnocranium 256 Tapirus 319
turhinals 261 Tarbosaurus 333
viseral arches 256 Tarsiers 353
sclerotic 256 Tec:to11i11s 353
skull of tetrapods 259-261 Teeth (Mammals) 271. 277
Smilodo11 317 brachydont 277
Soft hammer technique :no bunodont 277
Spalacotherium 313 hunolophodont 277
Sphe11acodo11 307 hunosclcnodont 277
Sphenacodonts 307 canine 273
Sp/Je11odo11 308 carnassial tooth 277
Spiny sharks 295 cement 277
Squamata 308 cheek teeth 275
Stahlekriid dicynodont 379 dental formula 273
Stegoceras 328, 334 dentine 277
Stegodon 348, 381 enamel 277
Stegodonts 348 incisors 273
Stegolophodontids 348 hypsodont 277
Stego/ophodon 348, 381 molars 273
Stegomastodon 348 premolars 273
Stegosaurs 327, 329 secodont 277
Stegosaurus 311, 328, 331, 334, 379 sclcnodonl 277
Stem-reptile 291 Teeth (Vertebrates) 271
Stcrcospondyls 303 acrodonts 271
Sternum (Tetrapods) 267 diphyodonts 271
breast bone 269 hcterodont 271
carina/sternal crest 269 homodont 271
cpisternum 267 labyrinthodont 273
manubrium 269 plcurodont 271
mcsosternum 267, 269 polyphyodont 271
metasternal process 269 thecodont 271
omosternum 267 Teleosteans (Teleostei) 297, 298
presternum 269 Terrible lizard 324
sternabrate 269 Tetrac/ae11odo11 340
sternal plate 267 Tetralophodon 347
thoracic busket 267 Thecodonts 309. 327
xiphisternum 267, 269 Theropods 333
xiphoid processes 269 Therapsids 307
xiphoid cartilage 269 Theriodonts
307
363
Suid 320 Theropithecus ge/ada
Sus 324
320, 381 Theropod dinosaurs
1rvDEX
(xxi)
fiki formation 376 Vertebrae (Comd.)
fillodonts 315 diapophyscs
fitanosaurus 328, 379 haemal arch 263
rrachodon 334 haemal spine
251. 263
320, 382 251. 263
'fragulids hypapophysis
320 neural arch
263
rragufus 249, 263
rree sloth 315 neural canal
304 263
Triadobatrachus neural process
328, 334 249
Triceratops neural spine 251, 263
'friconodonts 313, 382 parapophyses 263
Triloplwdo11 347,348 pygostyle 263
Trimerorachis 305 sacrum 263
Tritrylodo11 308 synsacrum 263
Tritrylodont 307-308 transverse process 263
Trituberculine molar 275, 313 urostyle 263
Trituberculine-sectorial zygapophyses 263
(Tribosphenicffrilobosphenic) 313, 319 Vertebral column 249
entoconid 315 Vertebrate 249
hypoconid 315
hypoconulid 315 Washburn's scheme 356
Mesoconid 315
metacone 315 Xenungulates 319
metconid 313
metaconule 313 Yerrapalli formation 376, 379
paracone 313 You11gia 308
paraconid 313
paraconule 313 Zalambdalestes 315
protocone 313
protoconid 313 PART IV : PLANT FOSSILS
talonid 313 Abies 415
trigon 313 Acer 412, 415, 430
trigonid 313 Acicularia 427, 428
Tuang man 364 Acritarchs 417-418
Tuatara 308 Actinostele 395
Tubulidontids 319 418
Adiantites
Tupaia (Ptiloceras) 323 Age of cycads 407
Turbosaurus ~28 Algal structures/Stromatol ites 401, 417
'Iylopods 320 413, 420
Angara flora
'Iyranosaurids 329 Angaroplois 413
Tyranosauras 309, 328, 332, 333 411-412
Angiospermae
Annual 392
Uintatherium 319 428
Arabicodium
Ungulates 410
319 Araucarites
~Pper palaeolithic culture 374 Ara11carioxylo11 410, 427
roctale 412
Ursus 304 Arb11tus
317 Archaeopteris 413
Archaeopteris flora 413
Vertebrae
249, 261, 263 Archaeopteris /atifolia 407
centrum Articulaticcae/Arthrophytu 405
249, 263
(>lxii)
PALAEONTOLOGY
Artisia 410 Cordaicarpus
Asteropl1yll 418 Cordaitales
410
A11stmliceras 409, 407
424 Cordaites/Noeggerathiopsis
Cortex
410
Baicalia 392, 395
417, 418 Crossotheca
Baiera 414 407
Cryptozoa,i
Balero 411,414 417
· Ctem,is
Barogwa11athi11 lo11gifolia 409, 414
403 Cycadeoidea
Benneltilales 409
407 Cycadophyton/Cycadales
Berbies 407
430 Cycas
Betula 409
415 Cyc/ostigma
Biennial 413
392 Cymose branching
8011ie11a 392
428
Brachyphyllum 410 Dacridium
Bract 415
399 Dadoxy/011 410
Buckla11dia 409 Dichotomous branching
Bunter floras 392
413 Dicroidium 424
Buriadia 410 Dicroidium flora 413
Dictyopteridium 419
Calamitales 405 Dictyozamites 409, 414
Calamites 405 Dipterocarpus 415, 428
Callipteris 413 Dissacate pollen 419
Callixylon 410, 413 Dissocladella 428
Calymites 405
Calymmatotheca 407 Ectoploic 395
Calypteris 407 Elatoe/adus 410
Cathysian flora 413 Endodermis 392, 395
Caytonia 407 Endoploic 395
Cenophytes (Cenozoic florras) · 412, 414-416 Epidermis 392, 395
Cephalotaxas 410 Equisetales 405
Chelo11iceras 424 Equisetites 405
Cheriolepis 410 Equisetum 405
Cladophlebis 414 Eur-American flora 413
Clethra 415 Eurydesma 420
Clypeina 428
Cocos 415, 428 Fermoria 417
Collenia 417, 418 Ficus 415
Columbiceras 424 Filicales 405
Columnaris 418 Flower 399
Compound leaf 399 Form genera 401
pinnate 399 Form species 401
palmate 399 Fruit 399
Complete flower 399 Furcula granulifera · 411
Coniferales 407
Co11iopteris 414 Ga11gamopte ris 407, 413
Conophyto,i 418 Gigantopteris 413
Conularia 420 Ginkgo 411, 414. 415
Cooksonia flora 412 Ginkgo biloba 410
Cordaianthus 410 Ginkgoales 407, 410
INDEX (xxiii)
Gi11kgoites 411 Leaf (Contd. )
Gi11kgopliyl/11m 411 margin 395
Gleic/1e11ites 414 phyllotaxy 399
Glossopteris 407, 411, 419, 420 simple leaf 399
GI) ptost rvbus 415 venation 395
Gondwana flora of India 418-426 Leaflet 399
Gyn111osolem 417 Leaf Scar 392
Gymnosolcmida 401 lebocedrus 415
Gymnospcrmac 407-411 lepidode11d ro11 403, 413
Lycopodium 401
Haplostele 395 lepidophyllum 403
Hartzia 411 lepidostrobus 403, 413
Helimeda 428 Libocedrus 415
Herbs 392 Lithocarpus 412
annual 392 Living fossil 410
biennial 392 Lobatommlaria 413
perennial 392 Lower Gondwana/Glossoptcris
Holarctic province 414 flora of India 413. 419-423
boreal zone 414-415 age 420
tethyan zone 414-415 botanical affinity 419-420
Ho/osporella 427, 428 components and stratigraphic
Homoxylon rajmahalensis 41 I, 424 distribution 420
Hyenia flora 412 geographic distribution 419-420
Hyeniales 405 origin 419
Hydrophycus 417 Lycopodium 403
Lycopsids 401. 403
Indigo/era 430 lygenostoma 407
lndo-Maleisan province 415 Lyginopteris 405. 407
lndophyton 417 lystrosaurus 424
lndopolia 428 lystrosaurus zo11e 424
/ndostrobus 410, 427
/soetes 403 Magnolia 409, 412, 415
Mammicus 418
Juniperus 430 Marattiopsis 414
Jurusa11ia 418 Matonidium indicum 426
Mesophytes/Mesozoic floras 413-414
Keteleeria 410 Mesoembryoxy/011 427
Keuper floras 413 Mesoxylon 410
Kussiella 401, 418 Minjaria 417
Modified stem 392
Labachia 410 Mohagaostrobus 427
Laplatosau rus 426
Larix 415 Navifussa 417
Lateral branching 392 Nelumbium 11 430
Leaf 395-401 Neomeris 427. 428
apex 395 Neotropic province 415
base 395 Neuropteris 407. 41:'
compound leaf 399 Nipa 411
lamina 395 Nilsso11ia 414
(oiv)
--
,,A I .A l·.'ONro I.OG y
NorRgerathio1,.vi.,· 4IO Prcl:11111hriun flora of lndiu
Non-vascular plan ls 400 Prrulu <' I u.,· 417-4 IX
Notal province 415-416 l'mduct,u fmm<t 42()
N ·mphat!a 42()
412 l'mlepidode11drrm
N pa 41~
415, 427 l'mtol, ye11iu
l'mtopte ridium 405
Orioporr-lla 405
428 Prolostclc
Osmundu 195
415 Ps e"doc t eII i.,·
Otozamitt!.'i 409
411,414 Psilophylalcs
401, 403
Psilophyto11
Pachyphyl/11m 401, 418
414 Psi/ophyto11 flora
Pachyptt!ris 413
414 P. pri11<:eps
Pagiophyllum 418
414 Psilopsids
Palaeophytcs/Palacozoic floras 412-413 401-403
Palat!oxy/011 Psilotalcs
413 Psygmophyllum 401
Palmoxylon 4IO, 413
411, 424, 428 Ptcridophytcs
Pararhesis
415 400
Paroria Pteridospcrm/Cycadofi Iicalcs/Sccd ferns
430 405, 407
Puoptuis Pterophyllum
413 Pteropsids 409, 414
Perennial
392 Ptilophyllum 405
Petiole 409, 414, 423
Petiolate 395 Ptilophyllum flora
395 Py11us 414, 423
Phaenicopsis
411 410
Phleboptuis
Phloem 414 Quercus
392, 395 415
Phyllocladus
Phylloglossus 415 Racemosc branching
Phyllotaxy 403 Rhacopteris 392
392 Racopteris flora 413
Phyllothua
405 R. cf circularis 415,418
Picea
Pinus 415 Rachis 418
Pith 415 Rajmahal flora 399
392, 395 R. i11equilatera
423, 424
Pityolepsis
410 R. ovata 418
Pityophyllum
414 Rhipidopsis 418
Pityostrobus
410, 427 Rhodea sp.
410, 413
Plectostele
395 Rhynia 418
Pleurotomaria
420 403
Podocedrus Rhyzome
415 392
Podocarpo11 Root system
415 391
Podozamites prop root
414 391
Populus root cap
415 391
Populus primaeva root hairs 391
Porsea 411 Rosa
415 412, 415
Post-Cambrian-Pre-Gondwana flora of India 418 Rugocystis 417
Post-Gondwana flora of India 426-430
Sabal 415
Deccan intertrappcan flora 427-428
Neogene flora 428-430 Sabalites 411
Quaternary flora of Kashmir 430 Salix 415
Upper-Cretaceous-Palaeocene flora 428 Sapidus 415
Scliizolepis 414
IN/JEX (xxv)
Sci, iWII <-'II I'll 405, 4 IJ 413

Tringia
Seed fern 414-415
405 Tropical province
Selagi11el/a 401 415
African zone
Sessile leaf 195 lndo-Muh;san zone 415
Shoot system '.WI Neolropic zone 415
re product ivc 191 True ferns 405
vegetative 391 Tuber 392
Shurbs 392 7imguJsia 401,417
Sigil/aria 403
Siphonostclc 395 Upper gondwana/Ptilophyllum/Rajmahal
Sorbus 412 flora of India 423-426
Spcrmatophytu 405 age 424-426
Sphe11obaiera 411 botanical affinity 424
Sphcnophy II ales 405 constitutents and distribution 424
Sphenophyllum 405 geographic occurrence 423-424
Sphenopsida 401 comparison with Glossopteris tlora 426
Sphenopteridium 413
Sphenopteridium furcillatum 418 Vascular plants 400
Sphenopteris 413 Venation 395-399
Spirifer 420 reticulate 395-399
Sporangia (Cones) 403 parallel 395-399
Stele 392 Vertebraria 407, 413, 419
Stem Viracarpon 415
bark 392 Voltzia floras 413
branching 392
tissue structure 392-395 Wealdon tlora 426
intcrnode 392 Weischselia reticulata 426
leaf scar 395 Williamsonia 409,414
node 392 Williamso11ia sewardia11a 409. 424
vascular bundles 392 Windwardia 411
Stigmania 403
Streptorynchus 420 Xylem 391, 392. 395
Stromatolites 401,417
Svalbardia flora 412 Zamites 414
Zosterophyllum flora 412
Tae11iopte ris 409
Takliostrobus 410, 427
417 PART V : MICROFOSSILS
Tasman ites
Taxas 410 Acodus 467
Terquemella 427 Acolpate pollen 475
Thallophytes 400 Acritarchs 470
Thimifeldia 407 Actipylea 461
Thuja 415 Adont 463
Ttlia
415 Aletc spore 475
Tttanosaurus 426 Allogromi11a 439, 452
Trachaea 400 Alternation of generation 439, 473
Trachaeophytes 400 Ammobacu/ires
392 458
Trees Ammodicaceans 459
Trigonocarpus 413 Ammodiscus
443
1~
PA LAEONTOLOGy
(xxvi)
458 Chamberlcts 445
Ammo11ia Chara 471
443
Ammovcrtdla C. coelata 471
463
Amphidont C. subglobossa 471
471
Am phi pleura C. malcomsoni 471
465
Anndida Charophyta 471
463
Antennae Chitinozoa 434
449
Aperture (Foraminifora) Chrysomonanidinids 469
475
Aporatc spore Chryosophyta 470
470
Archaeofavosi11a Coccoliths 470
469
Archeopyle Coccolithophorids 470
463, 465
Arthropod s/a Coccolithus 470
461
Assimilative layer Coccosphere 470
Astrorhiza
443 443-444
Coiled tests (Foraminifera)
convolute 444
Bathysiphon 443, 459
evolute 444
Biloculine 444, 458
involute 444
Biomorphs 470
milioline 444
Brachiopoda 465
planispiral 443
Bullate aperture 449
semiinvolute 444
trochospiral 443
Calcareous nannoplanktons 469
Colpi 475
Calcareous 475
test wall (Foraminifera) 439,443 Colporate pollen
443 Compylonies 471
glassy (Perforated)
443 Composition of testwall (Foraminifera) 439, 443
imperforated
443 agglutinated 439
porcellaneous
443 calcareous 439
pseudopunctate
451 proteinacaous 439
Calcarina
471 Concept of Iysocline 457
Calonies
461 Conchostrata 465
Calymma
452 Coniform 465
Canal & fissure
462, 463 Conodont s/a 465-467
Carapace
465 Conodont predator 467
Caudal furca
475 Cortex 471
Cavate
457 Cribrate 449
CCD/CCCD
461 Cribrohankenina 449
Central layer
471 Crown/Corona 471
Central nodule
471 Crustacea 462
Centric diatoms
463 Cycoccolithina 470
Cephalothorax
443 Cymbella 471
Chambers (Foraminifora)
443 Cypris 463
biserial
445 Cyst stage 469
coiled
445 Cytheridea 463
deuteroconch
444 Cyzicus
465
embryonic
nepionic/peri-embryonic 445
445 Denticles
465
protoconch
473
secondary /I ateral 445 Deploid generation
445
uncoiled 443 Deuteroconch
443 Diatoms 434, 470, 471
uniserial
. ? -
INDEX (xx vii)

Diatomaceous (Fuller's) earth 471 Frustulcs 470
Diatomin 470 Fusu/ina 443, 459
Digygopleura 463 Fusulinids 459
Dino flage llates 470
Dinophyceae 469 Gamctangium 473
Discolith 470 Gamelophytic plant 473, 475
Discorbis 451 Gamonl 439
Discocyclina 449, 451 Gamonl generation 439
Discocyclonids 445 Girdle 470
Distacodid conodonls 467 Globigerina 454. 458
G. pachydema 454
Echinoid Ossicles 434 Globigerina ooze 437, 458
Ecological study (Foraminifera) 453-459 Globigeri nids 459, 461
chemical environment 456-457 Globorotalia 451
contribution lo marine sediments 458-459 Globorotalids 459
food habit 457 Globotru ncana 458, 460
influence on shell-morphology 454 Globotruncanids 459
light 456 Globule (Anlheridium) 471
role of temperature 454 Growth stages (Foraminifera) 444
role of waler depth 454-455 embryonic 444
salient features 458 ephebic 444
salinity 456 geronlic 444
substrate 456 neanic 444
Eel-like animal 467 nepionic 444
Elphidium 458
Embryonic apparatus/chambers 443-445 Ha11tke11ina 451
ceulepidine 445 Haploid generation 473
isolepidine 445 Hastigeri11ella 451
nephrolepidinc 445 Heterohelix 451
polylepidine 445 Heterospore 475
Epitheca/Epivalvc 470 Homospore 475
Estheria 465 Hypotheca/Hypoval ve 470
E. ma11gliensis 465
Estheriids 465 lmperforate test 443
Eunice 465 In stars 463
Exine 475 In tine 475
Extra-capsular layer 461
Eye tubercle 463 Labiate aperture 449
Laeosphaeriada 470
Fibrous conodonts 467 Lagena 439
Foramen/foramina 437,449 Lagynacid 459
Foraminifera 437-461 Lamellar conodont 467
classification 452-453 Larger foraminifera 439
ecology 453-459 Lateral/Alar prolongation 444
general characters 437 Lateral/Secondary chambers 445
geological history 459 Left Valve 462
living foraminifcra 437, 439 leiotriletes 477
skeletal morphology 439-453 lepidocycli11a 445, 449, 45 1
use & application 459-461 Lcpidocyclinids 445
- r/ -~
,- /
~
\~
(xx viii) PALAEONTOLOGY

Lituolacca 459 Numrnulitids
459
Living foraminifcra 417-419 Nummulil.ic limesto ne 437
Living fossils 465
Lonchodina 467 Oogonium/Nu<.:ulc 471
Lonchodus 467 Ooze 458-459
Orbitoids 449, 459
Macropalaeontology 411 Ornamenlation (Foraminifcra) 451-452
Mandible 463 cancellate 451 -452
Marginal cord 444 costae 45]
Maxillae 463 flange 451
Median tooth 463 granules
Megafossils
451
434 keel 451
Megalospheric form 439 papillae
Megaspore 451
475 perforation 452
Merodont 463 pillars
Microfossils 451
434-478 pustule 451
acritarchs 470 septa! suture
advantage of study 451
434 Ostracods 434, 463, 465
application 477
chara 471 Palynology 434, 471
classification 436-437 Parasaccate - 477
coccoliths 469 Pectiniform conodonts 465
conodonts 465 Peneroplis 444
diatoms 470 Pennate diatoms 471
dinotlagellates 469 Perforate/glassy test
esthcriids 443
465 Peripylca
foraminifcra 437 461
Photic zone
limitation of study 459, 469
435 Phytoplanktons
ostracods 462, 465 457
Pillars
radiolaria
461 451
incised
scolecodonts
465 451
inflated
spore & pollen 471 451
Microgranular test wall
453
pseudo
residual
451 '
Micropalaeontology
433, 434, 471 451
textural
Microscopic arthropods
467 451
Microspheric form Planktonic foraminifera 453, 454, 457, 459
Microspore
439 Planktonic protozoans
475 461
Miogypsinids Planktic stage
445 469
Milioliids Pleurosigma
459 471
Miliolina Polar nodule
453, 456 470
Miospore Pollen
475 473
Monera Pollen tube
436 473
Monolete marks Polospore
475 475
Monoletc spore Polychaetes
475 465
Monophylca Porcelaneous test 443
461 Pores
475
Nannoplanktons Pre-pollen 473
Nodosaria 469, 470 Prioniodus
443, 478 467
Normal pore canal Protista 436
·-
,-,~· Nurnmulites 463
436, 447
Protoconch 445
11
Pscudochi ti nous/Protci naceous 439
... -=---- -
a ==
INDEX (xxix)
Pseudopodia 437, 461 Spore 473, 475
Punclosporitcs 477 Spore molher cell 473
Pyrrhophyta 469 Spore-pollen 471-477
advanlage of slutly 471
Quinqueloculie 444 ecology & tlis1rihu1ion 477
life cycle 473
Redial pore canal 463 morphology 475
Radiolaria 461-462 Spore lelrad 475
Radiolarian chert 462 Sporophylic plant 473
Ramiform condont 465 Slolon 449, 452
Raphe 470 Sulcus 463
Rhizammi11a 443 Symhiotic associalion 457. 469
Right valve 463
Rotalia 443, 453, 456 Tcctin 439
Rotaliinids 459 Tcctum 475
Rotaliinina 453,461 Terminal 100th 463
Test 437
Schizomicaphyta 436 Test wall (Foraminifera) 439, 445
Schizont 439 agglutinated 439
Schizont generation 439 calcareous 4W
Scolecodonts 434, 465, 467, 471 lamellar 440
Septa/Septum 447 non lamellar 440
Septa! suture/filaments 447 proteinaceous 439
basal 447 Tetrode/la 463
meandrine 447 Textularia 456
reticulate 447 Textulariids 459
sigmoidal 447 Textulariina 453
subreticulate 447 Tintinids (Ciliophora) 436
Sercedictyum 461 Tolypammi11a 459
Silicaflagellates 470 Tremalith 470
Skeletal morphology (Foraminifera) 439-453 Triloculina 443
aperture 449 Triloculine 444
arrangement of chambers 443 Tripylea 461
coiled test 443 Tube cell 471
composition-structure (Test wall) 439
function of test 447 Umbilicus 444
growth stages 444 Umbo 444
multilocular test 443 Use (Foraminifera) 459-461
ornamentations 451 hiostratigraphic 459
septa/suture 447 environmental 460
septa! suture 447
shape of tests 447 White matter 467
shape of chambers 447
unilocular test 443 Zooxa111helle 469
Smaller foraminifera 439
Somites 465 PART VI : STUDY OF Fossn.s
Spicules 434
Spiral suture 447 Aca11thoce ras 525
Sporangium 473 Alethopteris 542
PA LAEONTOLOcy
(xxx)
490 Farosites 495
Alveoliua F11sus
525 52)
Amalthtms
511
An:a Ga11gamopteris
507 535
Artlryris
490 Gleichenites 542
Assili11a
507 Globotnmcana 495
Atrypa
Glossopteris 535
Baculites 527 Glycimeris 511
Belenmites 527 Goniatites 525
Bellerop/1011 517 Gryplraea 515
Brey11ia 532
Bucklandia 541 Halysites 495
Hemiaster 531
Calceola 496 Hippario11 543
Calyme11e 501 Hippurites 517
Cardita 513
Cardium 513 Jsastrea 501
Ceratites 525
Ceritlrium 519 Lepidocycli11a 493
Cho11etes 505 Leptae11a 505
Cidaris 529
Cladophlebis 543 Maceration 484
Clypeaster 529 Macrocephalites 525
Collection of fossils 485, 487 Marattiopsis 543
Conularia 523 Micraster 531
Conus 521 Montlivaltia 501
Cordaites/Noeggerathiopsis 539 Murex 521
Cyclolites 501
Cypraea 519 Natica 519
Cyrena 513 Nautilus 523
Cystiphyllum 496 Nerita 519
Nucula 509
Dadoxy/011 539 Nummul ites 490
Description of fossil specimens 484-489
other characters 489 Ortlzis 505
shape 488 Orthoceras 523
SIZC 488 Ostrea 515
symmetry 487 Otozamites 539
Dictyozamites 541
Disaggregation 484 Paradoxides 503
Discocyclina 490 Pecopteris 542
Pecte11 515
Echinoconus 529 Penerop/is 495
Echi11olampas 531 Pentacri11us 532
Elatocladus
541 Pe11tamerus 509
Elephas
546 Pentremites 532
E11cri11us
Equus 535 Perisphinctes 525
Exogyra 543 Phacops 501
517 Physa 523
·1.
INDEX (xxxi)
Productus 503 Syri11gopora 496
Pterophyllum/Nilssv11 ia 539 Syri11gothyris 507
Ptilophyllum 539
Taeuiopteris 541
Redlichia 503 Te11taculites 523
Refinesquina 505 Terebratula 509
Rhinoceros 543 Textularia 493
Rhipidopsis 541 Trigonia 511
Rhy11cho11e/la 507 Triloculina 495
Rotalia 493 Trilophodo11 546
Trochus 519
Scapliites 527 Turbo 517
Schizaster 532 Turri lites 527
Schizoneura 542 Turritel/a 519
Scutel/a 529
Separation & Cleaning of fossils 483-484 Unio 511
Sphe11ophyllum 542
Spirifer 505 Venus 513
Spiriferina 507 Vertebraria 539
Spondylus 515 Voluta 521
Stegodon 546
Streptorhynchus 507 Wage11ophyllum/Lo11sdeleia 496
Strophome11a 505
Stygmatopygus 531 Zaphrentis 496

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An Introduction

•. . ) THE. WORLD PRESS PRIVATE LIMITED

PREFACE TO ·THE .SECOND EDITION

' , / , • ,', , : , \ I ' ,

Kolkata \ ; ~ ' :. ' .

Chapter 3 : Grade and Growth of Animals 22-29

Chapter 4 : Spatial Distribution of Organisms 30-37

4.4 Some Related Terms on 'Dispersal' 31

. ~: INVERTEBRATE FOSSILS 95-245

~apter 8 : Porifera (Sponges) 97-100

8.3 Classification JOO

PART III : VERTEBRATE Fossn.s 247-386

21.9 Bony Girdles and Limb Bones 269

23.2 Advent of Fishes 295

27 .3 Importance of Dinosaur Fossils of India 379

PART IV PLANT Fossn.s 387-430

PARTV : MICROFOSSILS 431-478

31.9.4 Diatoms 470

PART VI STUDY OF Fossn.s 479-546

Chapter 32 : Practical Works on Fossils 481-546

Table No. Contents Page No.

10.6 Internal morphology of Brachiopod shell 128

23. 11 A simple phylogenetic tree of Vertebrates showing evolution of different

31.13 Radioloria 462

A GENERAL ACCOUNT OF FOSSILS

1.1 FOSSILS-WHAT ARE THEY?

1.2 KIND OF FOSSILS

1.3 CONDITIONS OF PRESERVATION

1.3.1 Presence of hard skeletal matters · . .

1.3.2 Detachment from atmosphere

1.3.3 Sedimentation and permanent covering

1.3.S Effect of diage11etic processes

1.3.6 Less deformation and metamorphism of rocks

1.4 MODES OF PRESERVATION (Fig. 1-2)

p'= 7 p" = 7.8

A GENERAL ACCOUNT OF FOSSILS 5

1.4.2 Hard part fossils

(i) Unaltered hard parts

(ii) Altered hard parts .

In some cases, it is found that a mineral of the original skeleton be

1.4.3 Indirect fossil

1.5 CAUSES OF IMPERFECTION OF FOSSIL RECORD

(b) DISTILUZED FOSSIL

Dcfonned and loss of symmetry Articulated Disarticulated valves

Unusortcd shells of different siu

1.6 TAPHONOMJC~ALTERATION (Fig. 1-3)

1.6.1 Taphonomic changes found within a fossil (Fig. l-3a)

1.6.2 Changes foulld in fossil assemblage (Fig. 1-3b) ·

1.7 USE OF FOSSIL

1.7.1 Fossils ~sed in biost~atigrapliic classification of sedimentary rocks

1.7.2 Fossils used in correlation and age determin ti'

l o t eir fossils m older rocks may indicate a

1.7.4 Fossils used in reco11struction of palaeogeography

1.7.5 Fossils used as evidence of prehistoric life and organic evolution

1.7.6 Fossils used as economic tools

2.1 CLASSIFICATION AND NOMENCLATU~E ble types of animals and plants in

2.2 PHYLOGENETIC CLASSIFICATION . . .

2.3 PHENETIC (TYPOMORPHIC) CLASSIFICATION

2.4 TAXONOMIC CATEGORIES

2.5 CONCEPT OF A SPECIES · . . . .

2.6 NAMING A SPECIES : BIONOMIAL SYSTEM OF NOMENCL.\TURE

sapiens sapiens. . . · d'vidual variants

2 7 L AW OF PRIORITY-HOMONYMS AND SYNONYMS .

2.8 TYPE SPECIMENS