Ecology: From Individuals to Ecosystems

Description

A definitive guide to the depth and breadth of the ecological sciences, revised and updated The revised and updated fifth edition of Ecology: From Individuals to Ecosystems – now in full colour – offers students and practitioners a review of the ecological sciences.  The previous editions of this book earned the authors the prestigious ‘Exceptional Life-time Achievement Award’ of the British Ecological Society – the aim for the fifth edition is not only to maintain standards but indeed to enhance its coverage of Ecology. In the first edition, 34 years ago, it seemed acceptable for ecologists to hold a comfortable, objective, not to say aloof position, from which the ecological communities around us were simply material for which we sought a scientific understanding. Now, we must accept the immediacy of the many environmental problems that threaten us and the responsibility of ecologists to play their full part in addressing these problems. This fifth edition addresses this challenge, with several chapters devoted entirely to applied topics, and examples of how ecological principles have been applied to problems facing us highlighted throughout the remaining nineteen chapters.  Nonetheless, the authors remain wedded to the belief that environmental action can only ever be as sound as the ecological principles on which it is based. Hence, while trying harder than ever to help improve preparedness for addressing the environmental problems of the years ahead, the book remains, in its essence, an exposition of the science of ecology. This new edition incorporates the results from more than a thousand recent studies into a fully up-to-date text.  Written for students of ecology, researchers and practitioners, the fifth edition of Ecology: From Individuals to Ecosystems is anessential reference to all aspects of ecology and addresses environmental problems of the future.  Cover Title Page Copyright Page Contents Preface Introduction: Ecology and its Domain Chapter 1 Organisms in their Environments: the Evolutionary Backdrop 1.1 Introduction: natural selection and adaptation 1.2 Specialisation within species 1.2.1 Geographic variation within species: ecotypes 1.2.2 Genetic polymorphism 1.3 Speciation 1.3.1 What do we mean by a `species´? 1.3.2 Allopatric speciation 1.3.3 Sympatric speciation 1.4 The role of historical factors in the determination of species distributions 1.4.1 Movements of landmasses 1.4.2 Island history 1.4.3 Climatic history 1.5 The match between communities and their environments 1.5.1 Terrestrial biomes of the earth 1.5.2 The `life form spectra´ of communities 1.6 The diversity of matches within communities Chapter 2 Conditions 2.1 Introduction 2.2 Ecological niches 2.3 Responses of individuals to temperature 2.3.1 What do we mean by `extreme´? 2.3.2 Metabolism, growth, development and size 2.3.3 Ectotherms and endotherms 2.3.4 Life at low temperatures 2.3.5 The genetics of cold tolerance 2.3.6 Life at high temperatures 2.3.7 Temperature as a stimulus 2.4 Correlations between temperature and the distribution of plants and animals 2.4.1 Spatial and temporal variations in temperature 2.4.2 Typical temperatures and distributions 2.4.3 Distributions and extreme conditions 2.4.4 Distributions and the interaction of temperature with other factors 2.5 pH of soil and water 2.6 Salinity 2.6.1 Conditions at the boundary between the sea and land 2.7 Hazards, disasters and catastrophes: the ecology of extreme events 2.8 Environmental pollution 2.9 Global change 2.9.1 Industrial gases and the greenhouse effect 2.9.2 Global warming Chapter 3 Resources 3.1 Introduction 3.2 Radiation 3.2.1 Variations in the intensity and quality of radiation 3.2.2 Net photosynthesis 3.2.3 Sun and shade plants of an evergreen shrub 3.3 Water 3.3.1 Photosynthesis or water conservation? Strategic and tactical solutions 3.3.2 Roots as water foragers 3.4 Carbon dioxide 3.4.1 C3, C4 and CAM 3.4.2 The response of plants to changing atmospheric concentrations of CO2 3.5 Mineral nutrients 3.6 Oxygen – and its alternatives 3.7 Organisms as food resources 3.7.1 The nutritional contents of plants and animals and their extraction 3.8 A classification of resources, and the ecological niche 3.8.1 Categories of resources 3.8.2 Resource dimensions of the ecological niche 3.9 A metabolic theory of ecology Chapter 4 Matters of Life and Death 4.1 An ecological fact of life 4.2 Individuals 4.2.1 Unitary and modular organisms 4.2.2 Growth forms of modular organisms 4.2.3 Senescence – or the lack of it – in modular organisms 4.2.4 Integration 4.3 Counting individuals 4.4 Life cycles 4.5 Dormancy 4.5.1 Dormancy in animals: diapause 4.5.2 Dormancy in plants 4.6 Monitoring birth and death: life tables, survivorships curves and fecundity schedules 4.6.1 Cohort life tables 4.6.2 Survivorship curves 4.6.3 Static life tables 4.6.4 The importance of modularity 4.7 Reproductive rates, generation lengths and rates of increase 4.7.1 Relationships between the variables 4.7.2 Estimating the variables from life tables and fecundity schedules 4.8 Population projection models 4.8.1 Population projection matrices 4.8.2 Life table response experiments 4.8.3 Sensitivity and elasticity analysis Chapter 5 Intraspecific Competition 5.1 Introduction 5.1.1 Exploitation and interference 5.2 Intraspecific competition, and density-dependent mortality, fecundity and growth 5.2.1 Density-dependent mortality and fecundity 5.2.2 Intraspecific competition and density-dependent growth 5.2.3 Density or crowding? 5.3 Quantifying intraspecific competition 5.4 Intraspecific competition and the regulation of population size 5.4.1 Carrying capacities 5.4.2 Net recruitment curves 5.4.3 Sigmoidal growth curves 5.5 Mathematical models: introduction 5.6 A model with discrete breeding seasons 5.6.1 Basic equations 5.6.2 What type of competition? 5.6.3 Time lags 5.6.4 Incorporating a range of competition 5.6.5 Chaos 5.6.6 Stochastic models 5.7 Continuous breeding: the logistic equation 5.8 Individual differences: asymmetric competition 5.8.1 Size inequalities 5.8.2 The generation and dilution of size inequalities 5.8.3 Asymmetry enhances regulation 5.8.4 Territoriality 5.9 Self-thinning 5.9.1 Dynamic thinning lines 5.9.2 Species and population boundary lines 5.9.3 A single boundary line for all species? 5.9.4 An areal basis for self-thinning 5.9.5 A resource-allocation basis for thinning boundaries Chapter 6 Movement and Metapopulations 6.1 Introduction 6.2 Patterns of migration 6.3 Modes of dispersal 6.3.1 Passive dispersal 6.3.2 An active–passive continuum 6.3.3 Clonal dispersal 6.4 Patterns of dispersion 6.4.1 Patchiness 6.4.2 Forces favouring aggregation 6.4.3 Forces diluting aggregations: density-dependent dispersal 6.5 Variation in dispersal within populations 6.5.1 Dispersal polymorphism 6.5.2 Sex- and age-related differences 6.6 The demographic significance of dispersal 6.6.1 Dispersal and the demography of single populations 6.6.2 Invasion dynamics 6.6.3 Modelling dispersal: the distribution of patches 6.7 The dynamics of metapopulations 6.7.1 Uninhabited habitable patches 6.7.2 The development of metapopulation theory: islands and metapopulations 6.7.3 When is a population a metapopulation? 6.7.4 Metapopulation dynamics Chapter 7 Life History Ecology and Evolution 7.1 Introduction 7.2 The components of life histories 7.2.1 Reproductive value 7.3 Trade-offs 7.3.1 Observing trade-offs 7.3.2 The cost of reproduction 7.3.3 The number and fitness of offspring 7.4 Life histories and habitats 7.4.1 Options sets and fitness contours 7.4.2 High and low CR habitats: a comparative classification 7.4.3 Reproductive investment and its timing 7.5 The size and number of offspring 7.5.1 The number of offspring: clutch size 7.6 Classifying life history strategies 7.6.1 r- and K-selection 7.6.2 A fast–slow continuum 7.6.3 Grime´s CSR triangle 7.7 Phylogenetic and allometric constraints 7.7.1 Effects of size and allometry 7.7.2 Effects of phylogeny Chapter 8 Interspecific Competition 8.1 Introduction 8.2 Some examples of interspecific competition 8.2.1 Competition among phytoplankton species for phosphorus 8.2.2 Competition among plant species for nitrogen 8.2.3 Coexistence and exclusion of competing salmonid fishes 8.2.4 Some general observations 8.2.5 Coexistence of competing diatoms 8.2.6 Coexistence of competing birds 8.2.7 Competition between unrelated species 8.3 Some general features of interspecific competition – and some warnings 8.3.1 Unravelling ecological and evolutionary aspects of competition 8.3.2 A further warning: coexistence without niche differentiation? 8.3.3 Exploitation and interference competition and allelopathy 8.4 The Lotka–Volterra model of interspecific competition 8.4.1 The Lotka–Volterra model 8.4.2 Lessons from the Lotka–Volterra model 8.5 Consumer-resource models of competition 8.5.1 A model for a single resource 8.5.2 A model for two resources 8.5.3 Models with complex dynamics 8.5.4 Consumer–resource competition in practice 8.5.5 Spatial and temporal separation of niches 8.6 Models of niche overlap 8.6.1 Combining niche overlap and competitive similarity – a route to `neutral´ coexistence 8.6.2 A model of limiting similarity 8.7 Heterogeneity, colonisation and pre-emptive competition 8.7.1 Unpredictable gaps: the poorer competitor is a better coloniser 8.7.2 Unpredictable gaps: the pre-emption of space 8.7.3 Fluctuating environments 8.7.4 Aggregated distributions 8.8 Apparent competition: enemy-free space 8.9 Ecological effects of interspecific competition: experimental approaches 8.10 Evolutionary effects of interspecific competition 8.10.1 Natural experiments 8.10.2 Experimenting with natural experiments 8.10.3 Selection experiments Chapter 9 The Nature of Predation 9.1 Introduction 9.1.1 The types of predators 9.1.2 Patterns of abundance and the need for their explanation 9.2 Foraging: widths and compositions of diets 9.2.1 Food preferences 9.2.2 Switching 9.2.3 The optimal foraging approach to diet width 9.2.4 Foraging in the presence of predators 9.3 Plants´ defensive responses to herbivory 9.3.1 Plant defences 9.3.2 Apparency theory 9.3.3 The timing of defence: induced chemicals 9.3.4 Defending what´s most valuable 9.3.5 Defence when times are hard 9.4 Effects of herbivory and plants´ tolerance of those effects 9.4.1 Herbivory, defoliation and plant growth 9.4.2 Herbivory and plant survival 9.4.3 Herbivory and plant fecundity 9.4.4 Meta-analyses of herbivory 9.5 Animal defences 9.6 The effect of predation on prey populations 9.6.1 Intimidation: the non-consumptive effects of risk Chapter 10 The Population Dynamics of Predation 10.1 The underlying dynamics of consumer-resource systems: a tendency towards cycles 10.1.1 The Lotka–Volterra model 10.1.2 Delayed density dependence 10.1.3 The Nicholson–Bailey model 10.1.4 Predator–prey cycles in nature: or are they? 10.2 Patterns of consumption: functional responses and interference 10.2.1 The type 1 functional response 10.2.2 The type 2 functional response 10.2.3 The type 3 functional response 10.2.4 Individual and population-level satiation 10.2.5 Food quality 10.2.6 The effects of conspecifics – interference and ratio-dependent predation 10.3 The population dynamics of interference, functional responses and intimidation: equations and isoclines 10.3.1 The population dynamics of interference 10.3.2 The population dynamics of functional responses 10.3.3 The population dynamics of intimidation 10.4 Foraging in a patchy environment 10.4.1 Behaviour that leads to aggregated distributions 10.4.2 The optimal foraging approach to patch use 10.4.3 Ideal free and related distributions: aggregation and interference 10.5 The population dynamics of heterogeneity, aggregation and spatial variation 10.5.1 Aggregative responses to prey density 10.5.2 Heterogeneity in predator–prey models 10.5.3 Patch and lattice models 10.5.4 Aggregation, heterogeneity and spatial variation in practice 10.6 Beyond predator–prey Chapter 11 Decomposers and Detritivores 11.1 Introduction 11.2 The organisms 11.2.1 Decomposers: bacteria, archaea and fungi 11.2.2 Detritivores and specialist microbivores 11.2.3 The relative roles of decomposers and detritivores 11.2.4 Are local communities predisposed to deal effectively with local litter? 11.2.5 Ecological stoichiometry and the chemical composition of decomposers, detritivores and their resources 11.3 Detritivore–resource interactions 11.3.1 Consumption of plant detritus 11.3.2 Feeding on invertebrate faeces 11.3.3 Feeding on vertebrate faeces 11.3.4 Consumption of carrion Chapter 12 Parasitism and Disease 12.1 Introduction: parasites, pathogens, infection and disease 12.2 The diversity of parasites 12.2.1 Microparasites 12.2.2 Macroparasites 12.3 Hosts as habitats 12.3.1 The distribution of parasites within host populations: aggregation 12.3.2 Host specificity: host ranges and zoonoses 12.3.3 Hosts as resources and reactors 12.3.4 Hosts as reactors: resistance and recovery 12.3.5 Hosts as reactors: the cost of resistance 12.3.6 Hosts as reactors: resistance, tolerance and virulence 12.3.7 Competition among parasites for host resources 12.3.8 The power of coinfection 12.4 Coevolution of parasites and their hosts 12.5 The transmission of parasites amongst hosts 12.5.1 Transmission dynamics 12.5.2 Contact rates: density- and frequency-dependent transmission 12.5.3 Host diversity and the spread of disease 12.6 The effects of parasites on the survivorship, growth and fecundity of hosts 12.7 The population dynamics of infection 12.7.1 The basic reproductive number and the transmission threshold 12.7.2 Directly transmitted microparasites: R0 and the critical population size 12.7.3 Epidemic curves 12.7.4 Dynamic patterns of different types of parasite 12.7.5 Immunisation and herd immunity 12.7.6 Crop pathogens: macroparasites viewed as microparasites 12.7.7 Parasites in metapopulations 12.8 Parasites and the population dynamics of hosts 12.8.1 Red grouse and nematodes 12.8.2 An integral role for parasites? Chapter 13 Facilitation: Mutualism and Commensalism 13.1 Introduction: facilitation, mutualists and commensals 13.2 Commensalisms 13.3 Mutualistic protectors – a behavioural association 13.3.1 Cleaners and clients 13.3.2 Ant–plant mutualisms 13.4 Farming mutualisms 13.4.1 Human agriculture 13.4.2 Farming of insects by ants 13.4.3 Farming of fungi by beetles and ants 13.5 Dispersal of seeds and pollen 13.5.1 Seed dispersal mutualisms 13.5.2 Pollination mutualisms 13.5.3 Brood site pollination: figs and yuccas 13.6 Mutualisms involving gut inhabitants 13.6.1 Vertebrate guts 13.6.2 The vertebrate gut metagenome 13.6.3 Insect guts 13.7 Mutualism within animal cells: insect bacteriocyte symbioses 13.8 Photosynthetic symbionts within aquatic invertebrates 13.9 Mutualisms involving higher plants and fungi 13.9.1 Arbuscular mycorrhizas 13.9.2 Ectomycorrhizas 13.9.3 Ericoid mycorrhizas 13.9.4 Orchid mycorrhizas 13.9.5 Mycorrhizal networks 13.10 Fungi with algae: the lichens 13.11 Fixation of atmospheric nitrogen in mutualistic plants 13.11.1 Mutualisms of rhizobia and leguminous plants 13.11.2 Nitrogen-fixing mutualisms in non-leguminous plants 13.11.3 Nitrogen-fixing plants and succession 13.12 Models of mutualisms Chapter 14 Abundance 14.1 Introduction 14.2 Fluctuation or stability? 14.2.1 Determination and regulation of abundance 14.2.2 Approaches to the investigation of abundance 14.3 The demographic approach 14.3.1 Key factor analysis 14.3.2 λ-contribution analysis 14.4 The mechanistic approach 14.4.1 Experimental perturbation of populations 14.5 The time series approach 14.6 Population cycles and their analysis 14.6.1 Red grouse 14.6.2 Snowshoe hares 14.6.3 Microtine rodents: lemmings and voles 14.7 Multiple equilibria: alternative stable states Chapter 15 Pest Control, Harvesting and Conservation 15.1 Managing abundance 15.2 The management of pests 15.2.1 Economic injury levels and economic thresholds 15.2.2 Chemical pesticides and their unintended consequences 15.2.3 Evolution of resistance to pesticides 15.2.4 Biological control 15.2.5 Integrated pest management 15.3 Harvest management 15.3.1 Maximum sustainable yield 15.3.2 Harvesting strategies based on MSY 15.3.3 Economic and social factors 15.3.4 Instability of harvested populations: depensation and multiple equilibria 15.3.5 Instability of harvested populations: environmental fluctuations 15.3.6 Recognising structure in harvested populations: dynamic pool models 15.3.7 Rules of thumb for sustainable harvesting 15.3.8 Ecosystem-based fisheries management? 15.4 Conservation ecology 15.4.1 Introduction 15.4.2 Small populations 15.4.3 Causes of extinction 15.4.4 Minimum viable populations and population viability analysis 15.4.5 Conservation of metapopulations 15.4.6 Decision analysis Chapter 16 Community Modules and the Structure of Ecological Communities 16.1 Introduction 16.2 The influence of competition on community structure 16.2.1 Demonstrable competition between species 16.2.2 The structuring power of competition 16.2.3 Evidence from community patterns: niche differentiation 16.2.4 Niche differentiation – apparent or real? Null and neutral models 16.2.5 Evidence from morphological patterns – community-wide character displacement 16.2.6 Evidence from negatively associated distributions 16.2.7 Intransitive competition 16.3 The influence of predation on community structure 16.4 Plurality in the structuring of communities Chapter 17 Food Webs 17.1 Food chains 17.1.1 Trophic cascades 17.1.2 Top-down or bottom-up control of food webs? 17.1.3 Why is the world green? 17.2 Food web structure, productivity and stability 17.2.1 What do we mean by `stability´? 17.2.2 Strong interactors and keystone species 17.2.3 Complexity and stability in model communities 17.2.4 Relating theory to data: aggregate properties 17.2.5 Relating theory to data: community structure 17.2.6 Compartmentalisation 17.2.7 Organisation of trophic loops 17.2.8 Food chain length: the number of trophic levels 17.2.9 Parasites in food webs 17.3 Regime shifts Chapter 18 Patterns in Community Composition in Space and Time 18.1 Introduction 18.2 Description of community composition 18.2.1 Diversity indices 18.2.2 Rank–abundance diagrams 18.2.3 Community size spectra 18.3 Community patterns in space 18.3.1 Gradient analysis 18.3.2 The ordination of communities 18.3.3 Problems of boundaries in community ecology 18.4 Community patterns in time 18.4.1 Primary and secondary successions 18.4.2 Primary succession on volcanic lava 18.4.3 Primary succession on coastal sand dunes 18.4.4 Secondary successions in abandoned fields 18.5 The mechanisms underlying succession 18.5.1 A species replacement model of succession 18.5.2 A trade-off between competition and colonisation 18.5.3 Successional niche models 18.5.4 Facilitation 18.5.5 The role of animals 18.5.6 The role of functional traits 18.5.7 The nature of the climax 18.6 Communities in a spatiotemporal context 18.6.1 Disturbance, gaps and dispersal 18.6.2 The frequency of gap formation 18.6.3 Formation and filling of gaps 18.7 The metacommunity concept 18.7.1 The patch dynamics metacommunity model 18.7.2 The neutral metacommunity model 18.7.3 The species-sorting metacommunity model 18.7.4 The mass-effects metacommunity model 18.7.5 Patterns in abundance and diversity predicted by metacommunity models 18.7.6 The value and shortcomings of metacommunity models Chapter 19 Patterns in Biodiversity and their Conservation 19.1 Introduction 19.1.1 Estimating richness: rarefaction and extrapolation 19.2 A simple model of species richness 19.3 Spatially varying factors that influence species richness 19.3.1 Productivity and resource richness 19.3.2 Energy 19.3.3 Spatial heterogeneity 19.3.4 Environmental harshness 19.4 Temporally varying factors that influence species richness 19.4.1 Climatic variation 19.4.2 Environmental age: evolutionary time 19.5 Habitat area and remoteness: island biogeography 19.5.1 MacArthur and Wilson´s `equilibrium´ theory 19.5.2 Habitat diversity alone – or a separate effect of area? 19.5.3 Remoteness 19.5.4 Which species? Turnover 19.5.5 Which species? Disharmony 19.5.6 Which species? Evolution 19.6 Gradients of species richness 19.6.1 Latitudinal gradients 19.6.2 Gradients with elevation and depth 19.6.3 Gradients during community succession 19.7 Selecting areas for conservation 19.8 Managing for multiple objectives – beyond biodiversity conservation Chapter 20 The Flux of Energy through Ecosystems 20.1 Introduction 20.1.1 The fundamentals of energy flux 20.2 Patterns in primary productivity 20.2.1 Latitudinal trends in productivity 20.2.2 Temporal trends in primary productivity 20.2.3 Autochthonous and allochthonous production 20.2.4 Variations in the relationship of productivity to biomass 20.3 Factors limiting primary productivity in terrestrial communities 20.3.1 Inefficient use of solar energy 20.3.2 Water and temperature as critical factors 20.3.3 Drainage and soil texture can modify water availability and thus productivity 20.3.4 Length of the growing season 20.3.5 Productivity may be low because mineral resources are deficient 20.3.6 Do community composition and species richness affect ecosystem productivity? 20.4 Factors limiting primary productivity in aquatic communities 20.4.1 Limitation by light and nutrients in streams 20.4.2 Lakes and estuaries: the importance of nutrients and of autochthonous production 20.4.3 Nutrients and the importance of upwelling in oceans 20.4.4 Productivity varies with depth in aquatic communities 20.5 The fate of energy in ecosystems 20.5.1 Patterns among trophic levels 20.5.2 Possible pathways of energy flow through a food web 20.5.3 The importance of transfer efficiencies in determining energy pathways 20.5.4 Energy flow: spatial and temporal variation Chapter 21 The Flux of Matter through Ecosystems 21.1 Introduction 21.1.1 Relationships between energy flux and nutrient cycling 21.1.2 Biogeochemistry and biogeochemical cycles 21.1.3 Nutrient budgets 21.2 Nutrient budgets in terrestrial communities 21.2.1 Inputs to terrestrial communities 21.2.2 Outputs from terrestrial communities 21.2.3 Carbon inputs and outputs may vary with forest age 21.2.4 Importance of nutrient cycling in relation to inputs and outputs 21.3 Nutrient budgets in aquatic communities 21.3.1 Streams 21.3.2 Lakes 21.3.3 Estuaries 21.3.4 Continental shelf regions of the oceans 21.3.5 Open oceans 21.4 Global biogeochemical cycles 21.4.1 Hydrological cycle 21.4.2 Phosphorus cycle 21.4.3 Nitrogen cycle 21.4.4 Sulphur cycle 21.4.5 Carbon cycle Chapter 22 Ecology in a Changing World 22.1 Introduction 22.2 Climate change 22.2.1 Ecological risks 22.3 Acidification 22.3.1 Interactions among drivers 22.4 Land-system change 22.4.1 Expansion of the anthromes 22.4.2 Perturbation of nitrogen and phosphorus cycles 22.4.3 Downstream effects of nutrient cycle perturbations 22.5 Pollution 22.5.1 Chlorofluorocarbons, ozone depletion and UVB radiation 22.5.2 Mercury and persistent organic pollutants 22.5.3 Plastic waste 22.6 Overexploitation 22.7 Invasions 22.7.1 Winners and losers among invaders under climate change 22.7.2 Climate change, land-use change and invasion risk 22.8 Planetary boundaries 22.9 Finale References Organism Index Subject Index EULA