Ecology

Ecology
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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.

Оглавление

Michael Begon. Ecology

Table of Contents

List of Tables

List of Illustrations

Guide

Pages

ECOLOGY. From Individuals to Ecosystems

Preface. A science for everybody – but not an easy science

Thirty‐four years on: the urgent problems facing us

About this fifth edition

Technical and pedagogical features

Acknowledgments

Introduction: Ecology and its Domain. Definition and scope of ecology

Explanation, description, prediction and control

Pure and applied ecology

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

APPLICATION 1.1 Selection of ecotypes for conservation

1.2.2 Genetic polymorphism

APPLICATION 1.2 Variation within a species with man‐made selection pressures

1.3 Speciation

1.3.1 What do we mean by a ‘species’?

1.3.2 Allopatric speciation

1.3.3 Sympatric speciation

APPLICATION 1.3 Conservation significance of hot spots of endemism

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

APPLICATION 1.4 Global warming and species distributions and extinctions

APPLICATION 1.5 Human history and species invasions

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

APPLICATION 1.6 Stream invertebrate species traits and agricultural pollution

1.6 The diversity of matches within communities

Chapter 2 Conditions. 2.1 Introduction

2.2 Ecological niches

APPLICATION 2.1 Ecological niche modelling and ordination as management tools

APPLICATION 2.2 Judging the fundamental niche of a species driven to extreme rarity

2.3 Responses of individuals to temperature. 2.3.1 What do we mean by ‘extreme’?

2.3.2 Metabolism, growth, development and size

APPLICATION 2.3 Getting predictions right in the face of climate change

2.3.3 Ectotherms and endotherms

2.3.4 Life at low temperatures

2.3.5 The genetics of cold tolerance

APPLICATION 2.4 Selection for cold tolerance in crops to increase their productivity and geographic range

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

APPLICATION 2.5 Tropical species at particular risk from climate change

2.4.4 Distributions and the interaction of temperature with other factors

APPLICATION 2.6 Farmers’ choice of cover crops in relation to temperature and soil water potential

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

APPLICATION 2.7 Coral reefs and mangrove forests may ameliorate the impact of tsunamis

2.8 Environmental pollution

APPLICATION 2.8 Bioremediation and phytomining

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

APPLICATION 3.1 Bioengineering crops for accelerated recovery from photoprotection

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

APPLICATION 3.2 Turning to CAM crops

APPLICATION 3.3 Genetic engineering of CCMs into crops

3.4.2 The response of plants to changing atmospheric concentrations of CO2

APPLICATION 3.4 Harmful effects of plants’ responses to CO2 enrichment

3.5 Mineral nutrients

3.6 Oxygen – and its alternatives

APPLICATION 3.5 Permafrost, methanogenic anaerobic respiration and global warming

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

APPLICATION 4.1 Seed banks and the restoration of forested wetlands

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

APPLICATION 4.2 The survivorship curves of captive mammals

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

APPLICATION 4.3 Customised conservation of northern wheatears

4.8.3 Sensitivity and elasticity analysis

APPLICATION 4.4 Elasticity analysis and population management

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

APPLICATION 5.1 Optimal sowing rates for conservation

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

APPLICATION 5.2 Human population growth and a global carrying capacity

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

APPLICATION 5.3 Reintroduction of territorial vultures

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

APPLICATION 5.4 Density management diagrams

Chapter 6. Movement and Metapopulations. 6.1 Introduction

6.2 Patterns of migration

APPLICATION 6.1 Tracking the tracking migrations of locusts

APPLICATION 6.2 The conservation of migratory species

6.3 Modes of dispersal. 6.3.1 Passive dispersal

6.3.2 An active–passive continuum

APPLICATION 6.3 Winds predict the arrival of midges carrying bluetongue virus

6.3.3 Clonal dispersal

APPLICATION 6.4 Invasive fragmenting aquatic weeds

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

APPLICATION 6.5 Habitat restoration for a declining squirrel population

6.6.2 Invasion dynamics

APPLICATION 6.6 Invaders of the Great Lakes

6.6.3 Modelling dispersal: the distribution of patches

APPLICATION 6.7 Reaction–diffusion modelling of shifting species’ distributions under climate change

6.7 The dynamics of metapopulations. 6.7.1 Uninhabited habitable patches

APPLICATION 6.8 Species distribution modelling for (re)introductions and invasions

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

APPLICATION 6.9 Metapopulation capacities for birds and the giant panda

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

APPLICATION 7.1 Grain size and number in wheat

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

APPLICATION 7.2 Kiwis and Operation Nest Egg

7.6 Classifying life history strategies

7.6.1 r‐ and K‐selection

7.6.2 A fast–slow continuum

APPLICATION 7.3 The fast–slow continuum, invasion and conservation

7.6.3 Grime’s CSR triangle

APPLICATION 7.4 CSR and dark diversity

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

APPLICATION 8.1 Encouraging biodiversity in field margins

APPLICATION 8.2 Invasion and heterogeneity; invasibility and impact

8.8 Apparent competition: enemy‐free space

APPLICATION 8.3 Apparent competition threatens an endangered plant

APPLICATION 8.4 Red and grey squirrels, and squirrelpox virus

8.9 Ecological effects of interspecific competition: experimental approaches

APPLICATION 8.5 Additive experiments help guide prairie restoration

APPLICATION 8.6 Response surface analysis of the consequences of intercropping

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

APPLICATION 9.1 Switching can have economic importance in horticulture

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

APPLICATION 9.2 Using maize ‘landraces’ to improve inducible indirect defence

9.3.4 Defending what’s most valuable

9.3.5 Defence when times are hard

APPLICATION 9.3 Reducing stress to resist pine bark beetle attack

APPLICATION 9.4 Elevated CO2 and plant defence in sugar cane

9.4 Effects of herbivory and plants’ tolerance of those effects

9.4.1 Herbivory, defoliation and plant growth

APPLICATION 9.5 Invasion of a tolerant seaweed

9.4.2 Herbivory and plant survival

APPLICATION 9.6 The effects of aphids and the viruses they carry in response to climate change

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

APPLICATION 10.1 Generalist, switching predators as effective biocontrol agents?

APPLICATION 10.2 Human exploitation and a destabilising Allee effect

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

APPLICATION 10.3 What’s required of a good biological control agent?

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

APPLICATION 11.1 The importance of earthworms

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

APPLICATION 11.2 The value of dung beetles to agriculture

11.3.4 Consumption of carrion

APPLICATION 11.3 Forensic entomology and microbiology

APPLICATION 11.4 Ecosystem services provided by vultures

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

APPLICATION 12.1 Zoonotic infections

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

APPLICATION 12.2 Indirect effects of therapeutic treatments

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

APPLICATION 12.3 Myxomatosis

12.5 The transmission of parasites amongst hosts. 12.5.1 Transmission dynamics

12.5.2 Contact rates: density‐ and frequency‐dependent transmission

APPLICATION 12.4 Superspreaders and their identification

12.5.3 Host diversity and the spread of disease

APPLICATION 12.5 Lyme disease and the dilution effect

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

APPLICATION 12.6 Epidemic forecasting for Ebola

12.7.4 Dynamic patterns of different types of parasite

12.7.5 Immunisation and herd immunity

APPLICATION 12.7 Critical vaccination coverage

12.7.6 Crop pathogens: macroparasites viewed as microparasites

APPLICATION 12.8 Other classes of parasite and their control

12.7.7 Parasites in metapopulations

APPLICATION 12.9 Phocine distemper virus in harbour seals

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

APPLICATION 13.1 Commensalism, restoration and intercropping agriculture

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

APPLICATION 13.2 A mutualistic ant‐scale insect interaction may indirectly benefit coffee plants

APPLICATION 13.3 The large blue – a butterfly in danger

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

APPLICATION 13.4 Restoration of pollination networks

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

APPLICATION 13.5 Disruption of the honey bee gut metagenome by a pesticide

13.7 Mutualism within animal cells: insect bacteriocyte symbioses

APPLICATION 13.6 Novel uses of insect microbial symbionts to advance human welfare

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

APPLICATION 13.7 A role for lichens in medicine

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

APPLICATION 16.1 A parasitic threat to Darwin’s finches

16.4 Plurality in the structuring of communities

Chapter 17. Food Webs

17.1 Food chains

17.1.1 Trophic cascades

APPLICATION 17.1 Mesopredator release

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

APPLICATION 17.2 Sea otters – keystone initiators of a trophic cascade

APPLICATION 17.3 Humans as hyper‐keystones

17.2.3 Complexity and stability in model communities

17.2.4 Relating theory to data: aggregate properties

APPLICATION 17.4 Complementarity and food security

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

APPLICATION 17.5 A permafrost tipping point in the global ecosystem?

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

APPLICATION 18.1 Size spectra as indicators of human impairment of communities

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

APPLICATION 18.2 The application of succession theory to restoration

Invoking the theory of competition–colonisation trade‐offs

Invoking successional niche theory

Invoking facilitation theory

Invoking enemy interaction theory

Managing succession to restore a cultural tradition

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

APPLICATION 18.3 Managing successional mosaics for conservation

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

APPLICATION 19.1 Setting aside protected areas to conserve biodiversity

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

APPLICATION 19.2 Resolving conflicting requirements of agriculture versus conservation

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

APPLICATION 19.3 Nature reserves as ecological ‘islands’

19.6 Gradients of species richness

19.6.1 Latitudinal gradients

19.6.2 Gradients with elevation and depth

APPLICATION 19.4 Marine protected areas

19.6.3 Gradients during community succession

19.7 Selecting areas for conservation

APPLICATION 19.5 Site selection based on complementarity and irreplaceability

19.8 Managing for multiple objectives – beyond biodiversity conservation

APPLICATION 19.6 Marine zoning plans

APPLICATION 19.7 Holistic landscape planning for Catalonia, Spain

Chapter 20 The Flux of Energy through Ecosystems. 20.1 Introduction

APPLICATION 20.1 Ecosystem services

20.1.1 The fundamentals of energy flux

20.2 Patterns in primary productivity

APPLICATION 20.2 Human appropriation of net primary production

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?

APPLICATION 20.3 How important is biodiversity loss compared with other human‐induced factors?

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

APPLICATION 21.1 Nutrient pollution of aquatic ecosystems

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

APPLICATION 21.2 Managing forests to mitigate climate warming

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

APPLICATION 21.3 Constructing wetlands to reduce nitrate runoff to coastal seas

21.3.5 Open oceans

APPLICATION 21.4 Could fertilising the ocean with iron reduce global warming?

21.4 Global biogeochemical cycles

21.4.1 Hydrological cycle

APPLICATION 21.5 Flood risk, groundwater exploitation and climate change

APPLICATION 21.6 Ecohydrological responses to predicted climate change

21.4.2 Phosphorus cycle

APPLICATION 21.7 Human activities contribute the majority of phosphorus in inland waters

21.4.3 Nitrogen cycle

APPLICATION 21.8 Humans impact on the nitrogen cycle in diverse ways

21.4.4 Sulphur cycle

APPLICATION 21.9 Sulphur and acid rain

21.4.5 Carbon cycle

APPLICATION 21.10 Climate change and ocean acidification

Chapter 22 Ecology in a Changing World. 22.1 Introduction

22.2 Climate change

22.2.1 Ecological risks

APPLICATION 22.1 Combating the increased risk of forest fires in the boreal region

APPLICATION 22.2 A zero‐deforestation policy?

22.3 Acidification

22.3.1 Interactions among drivers

APPLICATION 22.3 Assisted evolution as a management response to climate warming and ocean acidification

22.4 Land‐system change

22.4.1 Expansion of the anthromes

APPLICATION 22.4 Establishing and managing protected areas

22.4.2 Perturbation of nitrogen and phosphorus cycles

22.4.3 Downstream effects of nutrient cycle perturbations

APPLICATION 22.5 Strategies for catchment management

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

APPLICATION 22.6 International action to deal with global pollution problems

22.6 Overexploitation

APPLICATION 22.7 Overfishing – the way forward

22.7 Invasions

22.7.1 Winners and losers among invaders under climate change

APPLICATION 22.8 Invasive plants in protected areas

22.7.2 Climate change, land‐use change and invasion risk

22.8 Planetary boundaries

22.9 Finale

References

Organism Index

Subject Index

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Отрывок из книги

MICHAEL BEGON

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Plants grow by developing new shoots from the buds that lie at the apices (tips) of existing shoots and in the leaf axils. Within the buds, the meristematic cells are the most sensitive part of the whole shoot – the ‘Achilles’ heel’ of plants. Raunkiaer argued that the ways in which these buds are protected in different plants are powerful indicators of the hazards in their environments and may be used to define the different plant forms (Figure 1.24). Thus, trees expose their buds high in the air, fully exposed to the wind, cold and drought; Raunkiaer called them phanerophytes (Greek phanero, ‘visible’; phyte, ‘plant’). By contrast, many perennial herbs form cushions or tussocks in which buds are borne above ground but are protected from drought and cold in the dense mass of old leaves and shoots (chamaephytes: ‘on the ground plants’). Buds are even better protected when they are formed at or in the soil surface (hemicryptophytes: ‘half hidden plants’) or on buried dormant storage organs (bulbs, corms and rhizomes – cryptophytes: ‘hidden plants’; or geophytes: ‘earth plants’). These allow the plants to make rapid growth and to flower before they die back to a dormant state. A final major category consists of annual plants that depend wholly on dormant seeds to carry their populations through seasons of drought and cold (therophytes: ‘summer plants’). Therophytes are the plants of deserts (they make up nearly 50% of the flora of Death Valley, USA), sand dunes and repeatedly disturbed habitats. They also include the annual weeds of arable lands, gardens and urban wastelands.

But there is, of course, no vegetation that consists entirely of one growth form. All vegetation contains a mixture, a spectrum, of Raunkiaer’s life forms. The composition of the spectrum in any particular habitat is as good a shorthand description of its vegetation as ecologists have yet managed to devise. Raunkiaer compared these with a ‘global spectrum’ obtained by sampling from a compendium of all species known and described in his time (the Index Kewensis), biased by the fact that the tropics were, and still are, relatively unexplored. Thus, for example, we recognise a chaparral type of vegetation when we see it in Chile, Australia, California or Crete because the life form spectrums are similar. Their detailed taxonomies would only emphasise how different they are.

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