Natural Systems

Dr Markus Eichhorn, School of Biology, University of Nottingham, UK.

• Preface xv0.0.1 To students xvi0.0.2 To instructors xviiAcknowledgments xxiAbbreviations xxiii1 Introduction: Defining nature 11.1 How little we know 11.2 Pressing questions 21.3 The hierarchy of nature 31.4 Biodiversity 51.5 Myths to bust 71.6 Further reading 81.6.1 Recommended reading 8PART I SPECIES2 What is a species? 32.1 The big question 32.2 Species concepts 62.2.1 Nominalistic species concepts 62.2.2 Morphological species concepts 72.2.3 Biological species concepts 82.2.4 Phylogenetic species concepts 102.2.5 Genetic species concepts 112.3 Solving the riddle 152.4 Coda: Species richness 162.5 Conclusions 162.5.1 Recommended reading 172.5.2 Questions for the future 173 The history of life 213.1 The big question 213.2 Sources of evidence 213.2.1 The fossil record 213.2.2 Molecular evidence 223.3 A brief history of diversity 233.4 Uneven diversity 293.5 Conclusions 313.5.1 Recommended reading 323.5.2 Questions for the future 324 How many species are there? 354.1 The big question 354.2 How can we not know? 364.3 Discovery rates 374.4 Scaling 404.5 Sampling-based methods 414.6 Other organisms 474.7 Wrapping up 484.8 Conclusions 504.8.1 Recommended reading 504.8.2 Questions for the future 50PART II DIVERSITY5 Measuring diversity 575.1 The big question 575.2 Scales of diversity 585.3 Species richness 585.4 Believing in estimates 635.5 A SAD story 655.6 Diversity of species 675.7 Other measures of diversity 715.8 diversity 755.9 Case study: the Binatang project 765.10 Conclusions 805.10.1 Recommended reading 805.10.2 Questions for the future 806 Niches 856.1 The big question 856.2 Historical background 866.3 Back to basics 896.4 Birth and death rates 906.5 The ZNGI 956.6 Impact vectors 986.7 Supply points 986.8 Coexistence 986.9 The evidence 1066.10 Implications 1126.11 Conclusions 1166.11.1 Recommended reading 1176.11.2 Questions for the future 1177 Patterns in species richness 1217.1 The big question 1217.2 Area 1217.3 Local and regional species richness 1247.4 Local patterns in species richness 1317.5 Congruence 1377.6 Assembling a model 1397.7 Conclusions 1397.7.1 Recommended reading 1407.7.2 Questions for the future 1418 Drivers of diversity 1478.1 The big question 1478.2 Coexistence or co-occurrence? 1488.3 Energy and resources 1488.4 Diversity begets diversity 1548.4.1 Heterogeneity in space 1548.4.2 Heterogeneity in time 1588.5 Disturbance 1608.6 Top-down control 1628.7 Expanding our model 1688.8 Conclusions 1698.8.1 Recommended reading 1698.8.2 Questions for the future 1699 Does diversity matter? 1759.1 The big question 1759.2 Ecosystems 1769.3 What shape is the relationship? 1789.4 Field experiments 1819.5 A problem with peas 1859.6 Other measures of diversity 1869.7 Multifunctionality 1889.8 The real world 1899.9 The link between species richness and productivity 1949.10 Conclusions 1949.10.1 Recommended reading 1959.10.2 Questions for the future 196PART III COMMUNITIES10 Organisation at the community scale 20310.1 The big question 20310.2 Definitions 20410.3 Communities in the field 20510.4 Quantitative approaches 20610.5 Community structure 21010.6 Food chains 21310.7 Food webs 21610.8 Complexity and stability 22110.9 Trophic cascades 22210.10 SAD again 22510.11 Complex systems 22810.12 Unified Neutral Theory 23110.13 Metabolic Theory of Ecology 23410.14 Conclusions 23610.14.1 Recommended reading 23710.14.2 Questions for the future 23711 Stability 24511.1 The big question 24511.2 Stable states 24511.3 Changing environments 24911.4 Hysteresis 25311.5 Predicting changes 25611.6 Coral reefs 25711.7 Shifting baselines 25911.8 Conclusions 26311.8.1 Recommended reading 26411.8.2 Questions for the future 26511.9 Coda: the seduction of Gaia 26512 Changes through time 27312.1 The big question 27312.2 Succession 27412.3 Succession and niche theory 27512.4 Examples of succession 27912.5 Disturbance 28112.6 Modelling succession 28312.7 Regeneration 28612.8 Plants and animals 28712.9 Case study: Mpala, Kenya 28812.10 Conclusions 29012.10.1 Recommended reading 29112.10.2 Questions for the future 29113 Changes through space 29513.1 The big question 29513.2 Community assembly 29613.2.1 Competitive exclusion 29713.2.2 Historical processes 30013.2.3 Habitat checkerboards 30213.2.4 Chance & contingency 30213.3 Metacommunities 30413.4 Dispersal limitation 31313.5 Combining environment and dispersal 31813.6 Conclusions 32213.6.1 Recommended reading 32213.6.2 Questions for the future 323PART IV BIOGEOGRAPHY14 Global patterns of life 33114.1 The big question 33114.2 Biogeography 33114.3 Phytogeography 33614.4 Ecoregions 34014.5 Empirical approaches 34114.6 The oceans 34514.7 Fresh water 34914.8 Conclusions 34914.8.1 Recommended reading 35014.8.2 Questions for the future 35015 Regional species richness 35515.1 The big question 35515.2 Climate and productivity 35715.3 Other processes 36015.4 Scale and productivity 36215.5 Latitudinal gradients 36715.6 Centres of origin 36915.7 Regional Species-Area Relationships 37015.8 Confounding effects 37115.9 Conclusions 37315.9.1 Recommended reading 37315.9.2 Questions for the future 37416 Latitudinal gradients 38116.1 The big question 38116.2 Hypotheses 38216.3 Geographic Area 38216.4 Climatic stability 38516.5 Productivity 38616.6 Niche size 38716.7 Evolutionary speed 39016.8 Out of the tropics 39316.9 Conclusions 39816.9.1 Recommended reading 39916.9.2 Questions for the future 39917 Earth history 40717.1 The big question 40717.2 Geological history 40817.3 Continental drift 40917.4 Echoes of Pangæa 41217.5 Climatic effects 41617.6 Ice Ages 41917.7 Sea level 42417.8 Extinctions 42517.9 Conclusions 42917.9.1 Recommended reading 43117.9.2 Questions for the future 43118 Dispersal 43718.1 The big question 43718.2 Range expansion 43818.3 Mechanisms of dispersal 44018.4 Barriers 44218.5 Case studies 44518.5.1 New Zealand 44518.5.2 Madagascar 44818.6 Conclusions 45418.6.1 Recommended reading 45518.6.2 Questions for the future 45519 Life on islands 46119.1 The big question 46119.2 Types of island 46219.3 Island biotas 46419.4 Evolution of endemics 46519.5 Size changes 46719.6 Reproduction and dispersal 47019.7 Super-generalists 47419.8 Endemic communities 47519.9 Disharmony 47519.10 Assembly rules 47719.11 Island species richness 47819.12 The Equilibrim Model of Island Biogeography 48119.13 Testing the theory 48519.14 Conclusions 48619.14.1 Recommended reading 48719.14.2 Questions for the future 48720 Reinventing islands 49320.1 The big question 49320.2 A critique of EMIB 49420.3 Rival hypotheses 49720.4 Disturbance 49820.5 Relaxation 50220.6 Extinctions 50420.7 Invasions 50520.8 A new theory? 50620.9 Evolution 50820.10 Conclusions 51520.10.1 Recommended reading 51520.10.2 Questions for the future 51621 What is a natural system? 52121.1 The big question 52121.2 Lessons learnt 52321.2.1 Ecological processes are scale-dependent 52321.2.2 All interactions are nested 52321.2.3 There is no such thing as the balance of nature 52421.2.4 Everything is contingent 52421.3 Processes not systems 525A Diversity analysis case study: Butterfly conservation in the Rocky Mountains 527A.1 Software resources 528A.2 Calculations 529A.3 Synthesis 536A.4 Conclusions 537Glossary 541Index 547

"Eichhorn�s book is a bold step...and shows the way ecology should be taught to students at all levels. At the same time, it presents an extensive overview of almost all important questions regarding biodiversity and the organization of living organisms at different levels that are currently under discourse...a work that is at the same time comprehensive, informative, educational, stimulating and pleasant to read. The standards this book has set will not be easily met by future authors." (Frontiers of Biogeography, 9(1), 27/03/2017)