Malthusian Relativity ι** = 1 / ψ
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A General Theory of Evolution
By selection by density dependent competitive interactions

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