Table of Contents
Preface xiii
Prologue 1
Chapter 1 Introduction 3
1.1 The classical theory of natural selection 3
1.1.1 Limitations to the classical theory 4
1.1.2 Historical and non-directional evolution 7
1.2 The proposed theory of natural selection 8
1.2.1 Integrating the two theories 9
1.2.2 Deterministic and directional evolution 11
1.2.3 Extraterrestrial life 14
1.3 Evolutionary population dynamics 15
1.4 The structure of the book 15
Part I Traditional theoretical ecology 17
Chapter 2 Malthusian increase 19
Chapter 3 Density regulation 23
3.1 Resource regulation 25
3.2 Interference regulation 26
3.3 Continuous logistic growth 27
3.4 Discrete logistic growth 28
3.5 Delayed density dependence 33
Chapter 4 Predator-prey 37
4.1 The lotka-volterra equations 39
4.2 Predator caused extinction 40
4.3 Adding interference competition 41
Chapter 5 Food chains 43
5.1 Exploitative versus interference competition 44
Chapter 6 Inter-specific competition 47
6.1 Exploitation: competitive exclusion 48
6.2 Intra-specific interference: competitive coexistence 49
6.3 Intra- and inter-specific interference: Hutchinson's rule 50
6.4 Appendix 52
Part II Evolution by natural selection 55
Chapter 7 Basic relations 57
7.1 Age-structured demography 58
7.2 Physiological constraints 62
7.2.1 The physicist and the evolutionist 63
7.2.2 Evolutionary constraints 64
7.3 A few ecological constraints 66
Chapter 8 Fitness and selection 69
8.1 Selection at different levels 70
8.2 Selection in the classical theory 72
8.3 Selection by density dependent competitive interactions 73
Chapter 9 Historical versus deterministic evolution 77
9.1 Lamarck and darwin 79
9.2 Historicity versus determinism 80
9.2.1 A mathematical distinction 81
9.2.2 Dimensionality of theoretical optima 83
9.3 Integrating the two theories 84
9.4 Equilibria at different levels 85
Part III Evolution of basic traits 89
Chapter 10 Body mass 91
10.1 The classical theory and no body mass 92
10.2 Selection by density dependent competitive interactions 95
10.2.1 The cost of competitive interactions 96
10.2.2 Density dependent bias in resource access 97
10.3 Competitive interactions and a large body mass 99
10.3.1 Density independent interference 100
10.3.2 Density dependent interference 100
10.3.3 Evolution of interference 103
10.4 Some predicted patterns 104
10.4.1 Body mass balanced against mortality 105
10.4.2 Bergmann's rule 106
10.4.3 The island rule 106
Chapter 11 Population limitation 109
11.1 The classical theory and no limit 110
11.2 Competitive interactions and a nature in balance 111
11.2.1 The size of resource quanta 113
11.2.2 Genetic variation 114
11.2.3 Metabolic rate 115
11.2.4 Rate of production in the resource 116
Chapter 12 Reproduction 117
12.1 The classical theory and unlimited reproduction 119
12.2 Competitive interactions and balanced reproduction 121
12.2.1 The evolution of Lack's optimum 122
12.2.2 Metabolic rate, resource quanta and production 123
12.2.3 Reproduction balanced against mortality 125
Chapter 13 Body mass allometries 129
13.1 Foraging self-inhibition 131
13.2 Intra-population interference 132
13.3 The allometric deduction 134
13.4 Empirical evidence 136
13.5 Appendix 139
13.5.1 The foraging optimum 139
13.5.2 The solution to five allometric equations 140
13.5.3 Additional allometries 141
Part IV The evolutionary steady state 143
Chapter 14 Exponential increase in body mass 145
14.1 Exponential increase in resource consumption 146
14.2 Exponential increase in body mass 147
14.3 Body mass allometries at steady state 149
14.3.1 Within-species allometry between reproduction and body mass 151
14.4 Evolutionary constraints 153
14.4.1 A lower constraint on body mass 153
14.4.2 An upper constraint on body mass 154
14.4.3 An upper constraint on the exploitation efficiency 155
14.5 Evolution as a deterministically unfolding process 156
Chapter 15 Exponential increase in metabolic rate 159
15.1 Scaling time with metabolism 160
15.2 Metabolic rate and lifespan of horses 57 million years ago 162
Chapter 16 Dwarfing and extinction 165
16.1 Dwarfing 166
16.2 Allometric disorder 168
16.3 Deterministic back-folding of biological systems 169
16.4 Why did mammals persist when dinosaurs became extinct? 170
Part V Evolution of derived traits 173
Chapter 17 Senescence and soma 175
17.1 On soma 176
17.2 On senescence 178
17.3 Evolution of senescence and soma 179
17.3.1 Trade-off between self-repair and senescence 179
17.3.2 Classical theory and unclear prediction 180
17.3.3 Competitive interactions and a clear transition 181
Chapter 18 Group size 183
18.1 Cost of grouping 185
18.2 Evolution of group size 186
Chapter 19 Fisherian sex ratios 191
19.1 One male per female 192
19.2 Investment sex ratios 194
19.3 Sex ratios in eusocial species 195
19.4 Local mating and female biased sex ratios 197
19.5 Four-fold cost of sex and limits to fisherian sex ratios 198
19.5.1 Two-fold cost of males 199
19.5.2 Two-fold cost of meiosis 202
Chapter 20 Males and sex ratios 205
20.1 Cost of males 207
20.2 Evolution of males 208
20.3 Evolution of sex ratios 209
20.4 Evolution of local mating 212
20.5 Evolution of male and female size 214
20.6 Male characters and sexual versus non-sexual selection 217
Chapter 21 Sexual reproduction and ploidy level 223
21.1 Sexual reproduction 225
21.2 Evolution of sexual inheritance 226
21.2.1 Evolution of diploid and haplodiploid genomes 227
21.3 Sex in sessile organisms 228
Chapter 22 Eusociality 231
22.1 Evolution of eusociality and worker caste 232
22.2 Evolution of kin selection and offspring workers 236
22.3 Sex ratios in eusocial species 238
22.3.1 Fisherian sex ratio with variation in worker sex ratio 239
22.3.2 Evolution of sex ratios in the worker caste 240
Worker sex ratio in diploids 242
Worker sex ratio in haplodiploids 242
22.3.3 Evolution of sex ratios in the sexual caste 243
Selection by two-fold cost of sexual males 244
Selection by pair formation 246
22.4 Diploid and haplodiploid eusocial species 248
22.4.1 Fisherian sex ratio with variation in ploidy level 249
22.4.2 Evolution of ploidy level 251
Two-fold cost males and a haplodiploid genome 252
Pair formation and a diploid genome 253
Part VI Evolutionary population dynamics 257
Chapter 23 Fundamental theorem replaces Malthusian law 259
23.1 Fundamental theorem leads to hyper-exponential increase 260
Chapter 24 Single species cycles 263
24.1 Logistic equation with density dependent selection 265
24.2 Population cycle driven by a cyclic population equilibrium 267
24.3 Cyclic phenotypes 269
24.4 The sex ratio cycle 270
24.5 Forest insects 271
24.6 Population cycle allometry 272
24.7 Implications of neutral stability 273
24.8 Extreme perturbations 274
24.9 Appendix 275
24.9.1 population equation with selection 275
Evolutionary changes in the growth rate 275
The evolutionary equilibrium 277
Stability and dynamic behaviour 278
24.9.2 Population equation with selection on the sex ratio 280
24.9.3 Parameter estimation 282
Epilogue 283
Chapter 25 Summary 285
25.1 Traditional theoretical ecology 286
25.1.1 Food chains 286
25.1.2 Competitive coexistence 287
25.2 Evolution of basic traits 287
25.2.1 Body mass 288
25.2.2 Population limitation 289
25.2.3 Reproduction 290
25.2.4 Body mass allometries 291
25.3 Evolutionary steady state 292
25.4 Evolution of derived traits 294
25.4.1 Senescence and soma 294
25.4.2 Males and sex ratios 295
25.4.3 Sexual reproduction and ploidy level 298
25.4.4 Eusocial colonies 300
25.5 Evolutionary population dynamics 302
25.6 Conclusion 303
References 305
Author index 323
Subject index 329
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