Palaeoecology: Ecosystems, environments and evolution

424 Pages·1997·15.899 MB·English

by  Patrick J. Brenchley, David A. T. Harper (auth.)

Palaeoecology
Ecosystems, environments and evolution
Patrick J. Brenchley
Department of Earth Sciences
University of Liverpool
UK
and
David A.T . Harper
University College Galway
Ireland
CHAPMAN &. HALL
London· Weinheim . New York· Tokyo· Melbourne' Madras
Published by Cilapmao & HaD, an impriat of 1bolDSOn Science,
2-6 Bonndary Row, London SE18HN, UK
Thomson Science, 2-6 Boundary Row, London SE1 8HN, UK
Thomson Science, 115 Fifth Avenue, New York, NY 10003, USA
Thomson Science, Suite 750, 400 Market Street, Philadelphia, PA 19106,
USA
Thomson Science, Pappelallee 3, 69469 Weinheim, Germany
Fust edition 1998
© 1998 P.I. Brenchley and D.A.T. Harper <!!I
Thomson Science is a division of International Thomson Publishing I(f)P
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ISBN-13: 978-0-412-43450-1 e-ISBN-13: 978-1-4684-14lO-3
DOl: lO.l007/978-1-4684-1410-3
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To our families, who have in their
various ways enriched the habitats in
which this book was written.
Contents
Preface xiii
Figure and table acknowledgements XIV-XXV
1 Investigating the history of the biosphere 1
1.1 Introduction 1
1.2 Definitions and principles 4
1.3 Marine and terrestrial environments 5
1.4 Life modes and trophic strategies 7
1.5 Geological framework 8
1.6 Global change 9
1.6.1 The atmosphere 9
1.6.2 The hydrosphere 13
1.6.3 The lithosphere 14
1.6.4 Biological change 15
1.7 Geophysiology 15
1.8 Strategy 20
1.9 Summary points 20
1.10 Further reading 21
1.11 References 21
2 Environmental controls on biotic distribution 24
2.1 The structure of the biosphere. 24
2.2 Divisions of the marine environment 26
2.3 Limiting factors Oft the distribution of organisms 26
2.4 Light 27
2.4.1 Palaeoecology 29
Box 2.1 A bathymetric analysis of Carboniferous
carbonate mud mounds using algae 30
2.5 Nutrients 29
2.5.1 Palaeoecology 34
2.6 Oxygen 34
2.6.1 Palaeoecology 38
Box 2.2 The Posidonienschiefer - a controversial
Jurassic black shale 40
Box 2.3 Carbon cycling and carbon isotopes 42
2.7 Temperature 43
2.7.1 Palaeoecology 45
Box 2.4 Oxygen-stable isotopes 48
I viii I I Contents
2.8 Salinity 50
2.8.1 Palaeoecology 52
Box 2.5 Palaeosalinities in Jurassic rocks of the
Great Estuaric group 53
2.9 Substrate composition 54
2.9.1 Palaeoecology 55
2.10 Substrate mobility, sedimentation rates and turbidity 57
2.10.1 Palaeoecology 59
2.11 Depth 59
2.12 The effects of limiting factors in different
environments 60
2.13 Summary points 62
2.14 Further reading 63
2.15 References 63
3 Taphonomy 66
3.1 The study of fossil preservation 66
3.2 Preservation potential amongst biological
communities 66
3.3 The fidelity of fossil assemblages 67
3.4 Post-mortem preservation and loss 69
3.5 Destruction by physical, biological and chemical
processes on the sediment surface 72
3.6 Preservation and destruction of shells below the
sediment surface 73
3.6.1 The shallow diagenetic environment 74
3.6.2 Preservation in carbonate concretions 74
3.6.3 Pyritic preservation 76
3.6.4 Preservation in phosphate 76
3.6.5 Preservation in silica 78
3.7 Preservation of soft tissue 79
3.8 Fossil LagersUitten - the exceptional preservation
of fossils 80
3.9 Types of shell concentrations and time-averaging 84
3.10 Information from shell concentrations in the
marine environment 87
3.11 The taphonomy of vertebrates 92
3.12 The preservation of plants 96
3.13 Summary points 98
3.14 Further reading 100
3.15 References 100
4 Adaptive morphology 103
4.1 Introduction 103
4.2 Terminology 104
4.3 Influential factors 105
4.4 Growth strategies 107
4.4.1 Shape description 107
4.5 Investigative methods 108
I
Contents ~
4.5.1 Analogues and homologues 110
4.5.2 Paradigm approach 110
4.5.3 Experimental palaeoautecology 112
4.5.4 Computer simulation 114
4.6 Adaptation and preadaptation 116
4.6.1 Functional thresholds 117
4.6.2 Size as an adaptation 118
4.6.3 Heterochrony 118
4.6.4 Vestigial structures 119
4.7 Morphology and environment 120
4.7.1 Pre-Vendian biota 121
4.7.2 Vendian evolutionary biota 121
4.7.3 Tommotian evolutionary biota 123
4.7.4 Cambrian evolutionary biota 124
Box 4.1 Growth strategies and function of
archaeocyathans 126
Box 4.2 Cambrian trilobites from the Alum Shale 127
4.7.5 Palaeozoic evolutionary biota 128
Box 4.3 Life strategies of rugose corals 129
Box 4.4 Brachiopod life modes 130
Box 4.5 Function of bryozoan colonies 133
Box 4.6 Palaeozoic bivalve life modes 134
Box 4.7 Trilobite ecomorphs 135
Box 4.8 Life modes and radiations of the graptolites 136
4.7.6 Modern evolutionary biota 137
Box 4.9 Modern bivalve life modes 139
Box 4.10 Gastropod morphology and adaptation 140
Box 4.11 Echinoid life strategies 141
Box 4.12 Cephalopod life styles 142
4.8 Summary points 142
4.9 Further reading 143
4.10 References 143
5 Trace fossils 148
5.1 Introduction 148
5.2 Preservation and taxonomy of trace fossils 149
5.3 Operational categories 152
5.4 Trace fossils and the behaviour of the producer 152
Box 5.1 Ethological classification of trace fossils 154
5.5 Trace fossils and palaeoenvironments 153
5.6 Marine and marginal marine ichnofacies 153
5.6.1 Focus on marine ichnofacies 155
Box 5.2 Computer simulation of deep-sea trails 160
5.7 Biological activity and ichnofabrics 159
5.8 Frozen profiles 162
5.9 Bioerosion 163
Box 5.3 Bioeroders 165
5.10 Terrestrial ichnofacies 165
I
~ Contents
5.10.1 Invertebrate traces 167
5.10.2 Vertebrate tracks 167
Box 5.4 Dinosaur trackways 171
5.10.3 Animal-plant interactions 170
5.11 Evolution of trace fossils 170
5.12 Summary points 175
5.13 Further reading 176
5.14 References 176
6 Fossils as environmental indicators 179
6.1 Evidence used in environmental analysis 179
6.2 Discriminating between environments on clastic
shelves 180
6.2.1 Biofacies distribution 180
6.2.2 Trace fossils 184
6.2.3 Diversity, biomass and size gradients 186
6.2.4 Taphonomic state 188
6.2.5 Types of shell concentration 189
6.3 Fossils as bathymetric indicators for marine shelf
sediments 191
6.4 Environmental indicators in deep marine sediments 196
6.5 Carbonate environments 197
6.5.1 Carbonate ramps 200
6.5.2 Rimmed shelves 201
6.5.3 Epeiric basins 202
6.6 Oxygen deficient environments 203
6.7 Environments with high and low salinity 207
6.8 Firmness of substrate 209
6.9 Shell concentrations, sedimentation rate and
sequence stratigraphy 211
6.10 Summary points 214
6.11 Further reading 215
6.12 References 215
7 Populations and communities 218
7.1 Population structure and dynamics 218
7.2 Types of popUlations 219
7.2.1 Size-frequency analyses 219
7.2.2 Age of fossil specimens 221
7.2.3 Survivorship curves 222
Box 7.1 Population analysis of a sample of the
fossil brachiopod Dielasma from the
Permian of northeast England 225
7.3 Variation in populations 223
7.4 Spatial distribution 224
7.5 Opportunist and eqUilibrium species 225
7.6 Life strategies and trade-offs 226
7.7 Long-term survivorship pattern 227
7.8 Community structure 227
10
Contents
7.9 Palaeo communities 230
7.10 Numerical analysis of community data 232
Box 7.2 Ecological indices 235
7.11 Community organization 240
7.11.1 Trophic structure 240
Box 7.3 Reconstruction of a Jurassic tropic structare 245
7.11.2 Tiering 251
7.11.3 Coevolution 252
7.11.4 Community succession 253
7.12 Species diversity 256
7.12.1 Diversity patterns on different scales 256
7.12.2 Diversity trend in marine habitats 257
7.12.3 Measuring diversity in palaeontological
samples 260
7.13 Environmental distribution of Phanerozoic
communities 262
7.14 Community replacement 263
7.15 Summary points 265
7.16 Further reading 266
7.17 References 267
8 Palaeobiogeography 271
8.1 Introduction 271
8.2 Modern biogeography 272
8.3 Some concepts and definitions 274
8.4 Controls on biogeography 275
8.4.1 Dispersal biogeography 276
Box 8.1 The role of larvae 277
8.4.2 Vicariance biogeography 279
Box 8.2 Distance and similarity coefficients 280
Box 8.3 Cladistics and biogeography 282
8.5 Recognition of past biogeographic provinces 285
Box 8.4 Faunal provinces through time 285
8.6 Palaeoclimatology 286
8.7 Plate movements 289
Box 8.5 North American Cordillera 290
8.8 Biogeography and evolution 293
Box 8.6 Changing diversity, immigration and
extinction 297
8.9 Biogeography and extinction 298
8.10 Summary points 298
8.11 Further reading 299
8.12 References 300
9 Evolutionary palaeoecology of the marine biosphere 303
9.1 The early history of life 303
9.2 Diversification events in Earth history 304
9.2.1 The origin of life and the earliest Procaryota 304
9.2.2 Appearance of the Eucaryota 305
I II
xii Contents
9.2.3 Appearance of the Metazoa 306
9.2.4 Broad patterns of metazoan diversification 306
9.2.5 The Ediacara fauna 307
9.2.6 The early Cambrian evolutionary explosion 308
9.2.7 Diversification of the three great
evolutionary faunas 311
Box 9.1 measuring changing diversity 316
9.2.8 Diversity independent models of
Phanerozoic diversity 317
9.2.9 The origins of diversity 317
9.3 Extinction 321
9.3.1 Patterns of extinction 321
9.3.2 Analysing patterns of extinction 322
9.3.3 Causes of extinction 326
9.3.4 The causes and ecology of some of the
major extinctions 335
9.3.5 Recovery from mass extinctions 346
9.3.6 The effect of major extinctions on evolution 347
9.3.7 Radiation, stasis and extinction 348
9.4 Summary points 349
9.5 Further reading 351
9.6 References 351
10 Fossil terrestrial ecosystems 357
10.1 Introduction 357
10.2 Initial adaptations and the early terrestrial record 358
10.2.1 Plants 359
10.2.2 Animals 361
10.3 Conquest of the Air 365
10.4 Trends 367
10.5 Terrestrial ecosystems. through time 370
10.5.1 Palaeozoic ecosystems 371
Box 10.1 Rhynie biota 374
Box 10.2 East Kirkton Lake 375
Box 10.3 Nyrany Lake 377
10.5.2 Mesozoic ecosystems 376
Box 10.4 Otter assemblage 379
Box 10.5 Cotswold assemblage 380
Box 10.6 Morrison assemblage 383
Box 10.7 Hell Creek assemblages 385
10.5.3 Cenozoic ecosystems 384
Box 10.8 East African Savanna 390
10.6 Mass extinctions 390
10.7 The Modern ecosystem 391
10.8 Summary points 392
10.9 Further reading 393
10.10 References 393
Index 397
Preface
Palaeoecology uses the fossil record to reconstruct the life habits of past organ
isms, their association in communities and their relationship to the environ
ments in which they lived. Traditionally, studies have concentrated on case
histories, using an array of fossil assemblages to reconstruct past ecosystems.
However, in recent years the focus of palaeoecology has been broadening as
it has become more fully appreciated that the biosphere is intimately linked
to the history of the planet as a whole. Progressively, there have been more
studies of the evolution of the ecological world. now comprising a sub-disci
pline of evolutionary palaeoecology. Other studies have focussed on the inter
play between the physical and biological world to show that, as in a Gaian
system, influences may work both ways; climate may influence the prevailing
ecosystems, but they in their turn may modify the climate systems. These are
the big themes with which we introduce the book. In the following sections
we have covered the traditional fields of palaeoecology, but have also explored
the ways in which palaeoecology can contribute to a broader understanding
of the evolution of the Earth. We have described how basic ecological infor
mation comes from applying knowledge of living organisms to the study of
past ecosystems and to the reconstruction of the past ecological world, at
population, community and biogeographic levels. We have also shown how
preservation affects the record on which such reconstructions are based. Such
palaeoecology knowledge is the basis for a better understanding of past envi
ronments, but it is also the basis for a broader appreciation of the role of
ecological systems in the evolution of our planet. In the later parts of the
book the nature of ecological evolution is explored in both marine and terres
trial environments with emphasis on changes in diversity, periods of radiation
and times of extinction and how these might relate to climate change, pertur
bations of the ocean, bolide impact, but also to the momentum of organic
systems themselves.
We are grateful to numerous colleagues and friends who have contributed
to this book with advice, discussion and support; in particular Jim Marshall,
Charlie Underwood, Euan Clarkson and Mike Benton, who were generous
with both their time and help. Special thanks go to Mike Benton, Euan
Clarkson, Andy Jeram, Paddy Orr, Alan Owen, Chris Paul, Ron Pickerill,
John Hudson, Derek Briggs, Andrew Smith, Paul Wignall, Jim Marshall and
Charlie Underwood who read chapters and offered good advice.
We would like to thank Dianne Edwards, Ed Jarzembowski, Andy Jeram,
Martin Lockley, Ron Pickerill, Euan Clarkson, John Hudson and Richard
Fortey for providing photographs for inclusion in the book.
The book arose from initial discussions with Simon Tull and we were
encouraged to continue writing by Ruth Cripwell and Ian Francis who has
seen the book through to completion. To these we offer our thanks.
Lastly, but very importantly we would like to acknowledge the work of
Kay Lancaster who has prepared all the figures for publication and so has
given her special stamp to the finished work. We have greatly appreciated
her skill, advice and perseverance.