Ecology
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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
WILEY END USER LICENSE AGREEMENT
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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