Plant Ecology

Michael J. Crawley, FRS, is with the Department of Biological Sciences, Imperial College of Science, Technology and Medicine. He is the author of three bestselling Wiley statistics titles and five life science books.

"The second edition Plant Ecology is outstandingly good. If you enjoyed the first edition, read the second. It is new. It can be recommended to teachers, research workers and able students. Complex theory is explained clearly; current issues are brought to the fore. Above all, it is inspiring. As Crawley says of plant taxonomy, 'It's fun'." Journal of Ecology "The first edition of the book was one of the most successful publishing events in the field of plant ecology in the 1980s. As the new edition seems to be improved in several directions it has a good chance to become a benchmark text in plant ecology even at the beginning of the new millennium. I can recommend it to anybody who wants to refresh his/her knowledge on the particular topic of plant ecology and as a supplementary text to students of a general ecology course." Leos Klimes, Folia Geobotanica, 2002, (37)

• List of Contributors xiPreface to the Second Edition xiiiPreface to the First Edition xv1 Photosynthesis 1Harold A. Mooney And James R. Ehleringer1.1 Introduction 1.2 Background: 1.2.1 Photochemical reactions; 1.2.2 Biochemical reactions. 1.3 Environmental influences on photosynthetic capacity: 1.3.1 Light; 1.3.2 Carbon dioxide; 1.3.3 Temperature; 1.3.4 Photosynthesis with respect to water use; 1.3.5 Energy balance considerations; 1.3.6 Nutrients; 1.3.7 Atmospheric pollutant. 1.4 Seasonality of photosynthesis: 1.4.1 Individual leaves; 1.4.2 Whole plants. 1.5 Photosynthetic capacity and defence against herbivores. 1.6 Variations on the basic photosynthetic pathway. 1.7 Ecological consequences of different photosynthetic pathways: 1.7.1 Water-use efficiency; 1.7.2 Significance of temperature. 1.8 Climate change and photosynthesis: 1.8.1 Photosynthesis in the recent past and near-future CO2 environments; 1.8.2 Climate change and the evolution of photosynthetic pathways. 1.9 Conclusions.2 Plant Water Relations 28John Grace2.1 Introduction: water and life: 2.1.1 Water as a physical and chemical medium; 2.1.2 State of water in the plant; 2.1.3 Acquiring and conserving water on land; 2.1.4 Water as a limiting resource. 2.2 Transpiration rate: 2.2.1 Energetics; 2.2.2 Stomatal conductance. 2.3 Soil–plant–atmosphere continuum: 2.3.1 Pathway; 2.3.2 Pipe model of hydraulic architecture; 2.3.3 How vulnerable is the pipeline? 2.4 Water relations and plant distribution patterns. 2.5 Water, carbon and nutrient relations. 2.6 Concluding remarks.3 Nutrient Acquisition 51Alastair Fitter3.1 Availability of nutrients. 3.2 Nutrient uptake by root system: 3.2.1 Transport through the soil; 3.2.2 Transport across the root. 3.3 Responses to nutrient deficiency: 3.3.1 Modifying the rhizosphere; 3.3.2 Resource allocation; 3.3.3 Symbioses. 3.4 Heterogeneity: 3.4.1 Patchiness; 3.4.2 Responses to patches; 3.4.3 Turnover. 3.5 Summary.4 Life History and Environment 73Michael J. Crawley4.1 Introduction_4.2 Neighbourhoods. 4.3 Lite history: 4.3.1 The growth forms of plants; 4.3.2 Annual plants; 4.3.3 Monocarpic perennials; 4.3.4 Herbaccous perennial plants; 4.3.5 Trees and shrubs. 4.4 Trade-offs: 4.4.1 Colonization competitive ability; 4.4.2 Root growth shoot growth; 4.4.3 Palatability competitive ability; 4.4.4 Seed size seed number; 4.4.5 Seed size seedling performance; 4.4.6 Seed size dormancy; 4.4.7 Dormancy dispersal; 4.4.8 Longevity growth rate; 4.4.9 .Longevity reproductive output; 4.4.10 Resource extraction growth rate; 4.4.11 Defence growth rate; 4.4.12 Growth reproduction; 4.4.13 Male female reproductive function; 4.4.14 Shade growth rate shade death rate; 4.4.15 Gap forest regeneration niche; 4.4.16 Sun leaves shade leaves and water light; 4.4.17 Growth rate nutrient retention; 4.4.18 Fruit weight seed weight; 4.4.19 Pollen quantity pollen quality; 4.4.20 Flammability competitive ability. 4.5 Canopy architecture: 4.5.1 Modular growth; 4.5.2 Integration of plant growth; 4.5.3 Allometry; 4.5.4 Plant height; 4.5.5 Leaf arrangement; 4.5.6 Phyllotaxis; 4.5.7 Switch from growth to reproduction; 4.5.8 Ageing and senescence. 4.6 Environmental factors affecting plant performance: 4.6.1 Fire; 4.6.2 Drought; 4.6.3 Waterlogging; 4.6.4 Shade; 4.6.5 Disturbance; 4.6.6 Low nutrient availability; 4.6.7 Soil acidity; 4.6.8 Heavy metals in soil; 4.6.9 Salinity; 4.6.10 Atmospheric pollutants; 4.6.11 Exposure; 4.6.12 Trampling; 4.6.13 Extremes of heat; 4.6.14 Mutualists; 4.6.15 Enemies; 4.6.16 Nurse plant. 4.7 Conclusions.5 Plant Secondary Metabolism 132Jeffrey R. Harborne.5.1 Introduction. 5.2 Secondary metabolites. 5.3 Terpenoid metabolites: 5.3.1 Mononterpenoids; 5.3.2 Sesquiterpenoids; 5.3.3 Triterpenoids. 5.4 Nitrogen-containing metabolites. 5.5 Phenolic metabolites. 5.6 Conclusions.6 Sex 156Michael J. Crawley6.1 Introduction. 6.2 Sex: why bother?: 6.2.1 Costs of sex; 6.2.2 Benefits of sex; 6.2.3 Variable progeny and individual fitness 6.3 Mating systems. 6.4 Inbreeding and outbreeding: 6.4.1 Population genetics of inbreeding; 6.4.2 Inbreeding depression; 6.4.3 Heterosis (hybrid vigour); 6.4.4 Outbreeding depression, 6.4.5 Kinds of self-pollination. 6.5 Sex types. 6.6 Incompatibility systems. 6.7 Prevention of self-pollination: 6.7.l Evolution of self-pollination from a cross-pollinating ancestor. 6.8 Limits to reproductive output: 6_8 1 Resource-limited fecundity; 6.8.2 Pollen-limited fecundity; 6.8.3 Population regulation. 6.9 Monocarpy and polycarpy. 6.10 Pollination by wind. 6.11 Pollination by animals: 6.11.1 Flowering phenology; 6.11.2 Nectar reward; 6.11.3 Pollen reward; 6.11.4 Plant spatial pattern. 6.12 Sexual investment by hermaphrodites: 6.12.1 Measuring the costs of male and female function; 6.12.2 Theory of male and female investment. 6.13 Agamospermy: seeds without sex. 6.14 Sex ratios and variable sex expression: 6.14.1 Sex determination in plants; 6.14.2Labile sex expression and environmental conditions; 6.14.3 Monoecy; 6.14.4 Dioecy. 6.15 Population genetics and genetic neighbourhoods; 6.15.l Minimum viable population (MVP); 6.l5.2 Genetic drift; 6.15.3 Effective population size; 6.15.4 Muration; 6.15.5 Selection; 6.15.6 Components of variance. 6.16 Gene flow through migration: 6.16.1 Gene flow through pollen; 6.16.2 Assortative and disassortative mating; 6.16.3 Venereal diseases of plants; 6.16.4 Gene flow through seed dispersal. 6.17 Sex on islands. 6.18 Local mate competition. 6.19 Mate choice in plants. 6.20 Conflicts of interest. 6.21 Case studies: 6.21.1 Paternity analysis; 6.21.2 Male fitness and pollen flow; 6.21.3 Selfing and inbreeding depression. 6.22 Conclusions.7 Seed Dormancy 214Mark. Rees7.1 Introduction: 7.1.1 Types of seeds; 7.1.2 Definitions of dormancy. 7.2 Seeds and the environment: 7.2.l Effects of light; 7.2.2 Effects of the chemical environment; 7.2.3 Effects of temperature; 7.2.4 Other germination cues. 7.3 Seed banks: 7.3.1 Temporal dynamics; 7.3.2 Physical structure. 7.4 Population persistence. 7.5 Population dynamics and coexistence. 7.6 Evolution of dormancy: 7.6.l Relationships between regenerative and established plant traits. 7.7 Conclusions.8 Mechanisms of Plant Competition 239David Tilman8.1Introduction. 8.2 Competition in natural plant communities: 8.2.1 Competition in a grassland field; 8.2.2 Limiting resources; 8.2.3 Competition for nitrogen and light. 8.3 A single limiting resource: 8.3.l The R* concept (‘R star’); 8.3.2 Resource dynamics; 8.3.3 Competition for a limiting resource; 8.3.4 Tests of the R* hypothesis. 8.4 Competition for two resources: 8.4.1 Resource isoclines; 8.4.2 Resource consumption vectors; 8.4.3 Resource supply vector; 8.4.4 Coexistence and displacement; 8.4.5 Experimental tests. 8.5 Multispecies communities: 8.5.1 Spatially discrete individuals; 8.5.2 Spatial heterogeneity; 8.5.3 Resource fluctuations and non-equilibrium conditions; 8.5.4 Multiple trophic levels. 8.6 Conclusion.9 Ecology of Pollination and Seed Dispersal 262Itenry F Howe And Lynn C. Westley9.1 Introduction. 9.2 Challenges of a sedentary existence. 9.3 Adaptive trends: 9.3.1 Flowers and pollinators; 9.3.2 Fruits and frugivores; 9.3.3 Cocvolution or co-occurrence? 9.4 Reproductive imperatives of success and failure: 9.4.1 Pollen success and failure; 9.4.2 Fertilized, unfertilized and aborted ovules; 9.4.3 Dispersed and undispersed seeds. 9.5 Adjusting tophysical and biological reality; 9.5.1 Physical environment; 9.5.2 Adjusting to neighbours. 9.6 Conclusions.10 Plant Chemistry and Herbivorv, or Why the World is Green 284Susan E. Hartley And Clive G. Jones10.1 Why is the world green? 10.2 Plants are poor food: they have ‘cruddy’ ingredient: 10.2.1 Nitrogen limitation of herbivores; 10.2.2 Secondary metabolites and herbivores; 10.2.3 Last thoughts on secondary metabolism and how green the world is. 10.3 Plants are poor food: they are unpredictable: 10.3.1 Intrinsic heterogencity; 10.3.2 Extrinsic heterogeneity; 10.3.3 Last thoughts on unpredictability and how green the world is. 10.4 Herbivores are between the devil and the deep blu1: sea. 10.5 Conclusions.11 The Structure of Plant Populations 325Michael J. Hutchings11.1 Introduction. 11.2 Performance structure in plant populations: 11.2.1 Plant weights; 11.2.2 Other aspects of performance. 11.3 Spatial structure of plant populations: 11.3.l Spatial structure of seed and seedling populations; 11.3.2 Spatial structure of population of established plants. 11.4 Age structure in plant populations: 11.4.1 The seed bank: dispersal in time; 11.4.2 Age structure or the growing plants in populations; 11.4.3 Age structure of populations of modules. 11.5 Generic structure of plant populations. 11.6 Abiotic influences on population structure.12 Plant Population Dynamics 359Andrew R. Watkinson12.l Introduction. 12.2 Population flux. 12.3 Population regulation. 12.4 The individual and the population. 12.5 The fates of individuals: 12.5.1 Fates of seeds; 12.5.2 Fates of individuals classified according to age and stage 12.6 Population models: 12.6.1 Matrix models; 12.6.2 Difference equations. 12.7 Density-dependence. 12.8 Population dynamics: 12.8.1 Annual plants; 12.8.2 Perennial plants. 12.9 Interactions in mixtures of species: 12.9.1 Interspecific competition; 12.9.2 Mutualism. 12.10 Concluding remarks.13 Plant–Herbivore Dynamics 401Michael J. Crawley13.1 Introduction. 13.2 Herbivores and plant performance: 13.2.1 Seedling growth and survival; 13.2.2 Shoot growth; 13.2.3 Root growth; 13.2.4 Plant shape; 13.2.5 Flowering; 13.2.6 Fruiting and fruit dispersal; 13.2.7 Seed production; 13.2.8 Seed predation; 13.2.9 Mast fruiting and predator satiation; 13.2.10 Mature plant death rate. 13.3 Herbivores and plant vigour: 13.3.1 Herbivory and plant productivity; 13.3.2 Plant stress hypothesis; 13.3.3 Plant vigour hypothesis; 13.3.4 Herbivore: plant–herbivore interactions. 13.4 Plant compensation: 13.4.1 Reduced rate, of fruit and seed abortion; 13.4.2 Grasses; 13.4.3 Trees; 13.4.4 Shrubs; 13.4.5 Herbs. 13.5 Herbivores and plant fitness. 13.6 Overgrazing. 13.7 Herbivores and plant genetics. 13.8 Herbivores and atmospheric CO2. 13.9 Herbivores and plant population dynamics: 13.9.1 Herbivory and plant competition; 13.9.2 Herbivores and plant demography; 13.9.3 Generalists and specialists; 13.9.4 Plant growth; 13.9.5 Herbivore functional responses; 13.9.6 Herbivore numerical responses; 13.9. 7 Herbivore density dependence; 13.9.8 Granivory: the dynamics of seed predation. 13.10 Case studies: 13.10.1 Keystonc herbivores: the kangaroo rats of southern Arizona; 13.10.2 Exclusion experiments using fences against large vertebrate herbivores; 13.10.3 Cyclic herbivore populations; 13.l0.4 Weed biocontrol; 13.10.5 Exclusion experiments involving insect herbivores and chemical pesticides. 13.11 Herbivores and plant diversity: 13.11.1 Selective herbivory and the identity of the dominant plant species; 13.11.2 Selective herbivory and plant species richness. 13.12 Herbivores and plant succession: 13.12.1 Primary succession; 13.12.2 Secondary succession. 13.13 Summary14 The Structure of Plant Communities 475Michael J. Crawley14.1 Introduction. 14.2 Definition of plant community: 14.2.1 Clements’ view of community structure; 14.2.2 Gleason’, view of community structure; 14.2.3 The modern synthesis. 14.3 The niche concept. 14.4 Species richness: 14.4.1 Spatial heterogeneity; 14.4.2 Temporal variation; 14.4.3 Competitive ability dispersal trade-off; 14.4.4 Niche separation and resource partitioning; 14.4.5 Herbivory and the palatability competitive ability trade-off; 14.4.6 Disturbance; 14.4.7 Refuges; 14.4.8 Alpha, beta and gamma diversity; 14.4.9 Species richness in the: Park Grass Experiment: a case study. 14.5 Evenness and relative abundance: 14.5.1 Species–area effects; 14.5.2 Biogcography; 14.5.3 Species abundance distributions. 14.6 Physical structure of plant communities: 14.6.1Life-forms in plant communities; 14.6.2 Vertical structure: of plant communities; 14.6.3 Spatial structure of plant communities; 14.6.4 Allelopathy and spatial patterns; 14.6.5 Quantitative methods for describing spatial patterns; 14.6.6 Spatial patterns and quadrat size; 14.6.7 Spatial patterns reflecting temporal changes. 14.7 Succession: 14.7.1 Interglacial cycles; 14.7.2 Primary succession; 14.7.3 Secondary succession. 14.8 Models or spatial dynamics: 14.8.1 Metapopulation models; 14.8.2 Patch models; 14.8.3 Reaction diffusion models; 14.8.4 Cellular automata; l4.8.5 Coupled map lattice. 14.9 Conclusions.15 Dynamics of Plant Communities 532Stephen W. Pacala15.1 Introduction. 15.2 Simple models of ideas: 15.2.l Competition colonization trade-off; 15.2 2 Resource partitioning; 15.2.3 Temporal partitioning: the storage effect; 15.2.4 Janzen–Connell hypothesis. 15.3 Empirical tests. 15.4 Models of natural systems. 15.5 Spatial segregation hypothesis. 15.6Empirical evidence for the spatial segregation hypothesis. 15.7 Conclusions.16 Plants in Trophic Webs 556James P. Grover And Robert D. Holt17 Plants and Pollution 568Mike Ashmore17.1 Introduction. 17.2 Effects on individual plants. 17.3 Effects on species interactions. 17.4 Evolutionary responses. 17.5 Community-level effects. 17.6 Concluding remarks.18 Climate Change and Vegetation 582J. Philip Grime18.1 Introduction. 18.2 Importance of land use. 18.3 Current predictions: 18.3.1World vegetation patterns; 18.3.2 Regional vegetation patterns. 18.4 Current research: 18.4.1 A research protocol; 18.4.2 Screening of plant attributes; 18.4.3 Formal searches for plant functional types; 18.4.4 Monitoring of vegetation responses to climate; 18.4.5 Manipulative experiments 18.5 Conclusions.19 Biodiversity 595Michael. J. Crawley19.1 Introduction. 19.2 The number of plant species. 19.3 Origins of plant biodiversity. 19.4 Postglacial changes in plant biodiversity. 19.5 Current geographical distribution of biodiversity: 19.5.1 Biodiversity hot-spots; 19.5.2 Cape floral kingdom of South Africa; 19.5.3 Island floras. 19.6 Variation in plant biodiversity within the British Isles. 19.7 Threats to biodiversity: 19.7.1 Species loss in Britain; 19.7.2 Species loss in tropical environments; 19.7.3 Urbanization; 19.7.4 Enforcement of conservation legislation. 19.8 Alien plants: 19.8.1 Notions of invasive and non-invasive: species; 19.8.2 Problem plants; 19.8.3 What are the problem plants?; 19.8.4 Problem plants in other countries; 19.8.5 Overview of problem plants. 19.9 Plant conservation: 19.9.1Parks and nature reserves; 19.9.2 Habitat restoration; 19.9.3 Botanic gardens; 19.9.4 Gene banks. 19.10 Food plant conservation. 19.11 Economics, of plant conservation. 19.12 Conclusions.References 633Index 701Colour plates fall between pp. 366 and 367