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The Evolutionary Ecology of Weeds and Invasive Plants Dekker: The Evolutionary Ecology of Weeds and Invasive Plants v.12.12.08 2009 Course Version ___________________________________________________________________________ The Evolutionary Ecology of Weeds and Invasive Plants Jack Dekker Weed Biology Laboratory Agronomy Department Iowa State University Ames, Iowa 50011 EMail: [email protected]; Web URL: www.public.iastate.edu/~jdekker ___________________________________________________________________________ 1 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants Copyright 2009 Jack Dekker 2 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants TABLE OF CONTENTS PAGE FORWARD 11 INTRODUCTION 13 Approach to weed biology The weeds always win The evolutionary ecology of weeds and invasive plants UNIT 1: THE NATURE OF WEEDS 17 1 The Nature of Weeds 19 1.1 Definitions of a weed 1.2 Terminology of other weedy plant types 1.3 Weeds defined by their traits UNIT 2: WEED EVOLUTION 23 2 Natural Selection and Weedy Adaptation 25 2.1 Evolution, adaptation and natural selection 2.2 Evolution 2.3 Adaptation 2.4 Natural selection 2.4.1 Process of natural selection: 2.4.1.1 Reproduction 2.4.1.2 Heredity and inheritance 2.4.1.3 Variation in fitness of organisms 2.4.1.4 Variation in individuals and traits 2.4.2 Units of selection 2.4.3 Selection and population diversity 2.5 Genotypes and phenotypes 2.6 Genetic Variation 2.6.1 Mating system variation 2.6.1.1 Apomictic species 2.6.1.2 Self-pollinating species 2.6.1.3 Out-crossing species 2.6.2 Sources of genetic diversity 2.6.2.1 Forces increasing population diversity 2.6.2.3 Forces decreasing population diversity 2.7 Phenotypic Variation 2.7.1 Phenotypic plasticity 2.7.2 Somatic polymorphism 2.8 Speciation 2.8.1 Process of speciation 2.8.2 Reproductive isolating mechanisms 2.8.3 Modes of speciation 3 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants UNIT 3: AGRICULTURAL COMMUNITY DYNAMICS 57 3 Opportunity Space-Time 59 3.1 Local opportunity: space and time 3.2 Weedy habitats 3.3 Niches in the local community 3.4 Local opportunity space-time structure 3.4.1 The niche hypervolume 3.4.2 Four dimensions of the niche hypervolume 3.4.2.1 Spatial heterogeneity and patchiness 3.4.2.1.1 Lateral heterogeneity of environments 3.4.2.1.2 Vertical heterogeneity of environments 3.4.2.1.3 Selection in a patchy environment 3.4.2.1.4 Dispersal mechanisms set the scale of a plant's spatial heterogeneity 3.4.2.2 Temporal division of the environment 3.4.2.3 Disturbance 3.4.2.3.1 Disturbance: local opportunity 3.4.2.3.2 Dimensionality of disturbance 3.4.2.3.3 Proximity of disturbance effects 3.4.2.3.4 Vulnerability to disturbance 3.4.2.3.5 Temporal patterns in disturbance 3.4.2.4 Differential use of resources and conditions 3.5 Limiting resources-conditions in local opportunity space-time 3.5.1 Limiting resources in the environment 3.5.1.1 Light as resource 3.5.1.2 Water as resource 3.5.1.3 Mineral nutrients as resources 3.5.1.4 Gases as resources 3.5.2 Pervasive conditions in the environment: heat 4 Plant Invasions 73 4.1 Seizing, exploiting and occupying opportunity space-time 4.2 The plant invasion process 4.2.1 The invasion matrix 4.2.2 Dispersal 4.2.3 Colonization 4.2.4 Enduring occupation of a locality 4.2.5 Extinction 4.4 The perception of plant invasion 4.4.1 Human values 4.4.2 Conclusions 4.4.2.1 The human role in creating opportunity space for plant invasion 4 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants 4.4.2.2 Utilizing and exploiting beneficial plant invasive species 5 Plant Community Structure 81 5.1 Introduction 5.2 Weed Biodiversity 5.2.1 Introduction 5.2.2 Biodiversity levels within a habitat 5.2.3 Stability, sustainability and biodiversity in plant communities 5.3 Weed community structure 5.3.1 Population genetic structure 5.3.1.1 Introduction 5.3.1.2 Case study: The weedy Setaria species-group 5.3.1.2.1 Genetic diversity in the weedy foxtails 5.3.1.2.2 Genetic pattern of colonization and regional differentiation 5.3.1.2.3 Genetic variation and evolutionary success of colonizers. 5.3.2 Species associations for colonization 5.3.2.1 The origin of weeds: Wild-crop-weed plant complexes 5.3.2.2 Preadaptive colonizing archetypes 5.3.2.2.1 Generalist-specialist genotypes 5.3.2.2.2 Reproductive colonizing types 5.3.2.3 Genotype structuring: colonizing species associations 5.3.2.3.1 Species-groups 5.3.2.3.2 Polyploid species clusters 5.3.2.3.3 Aggregate species 5.4 Exploiting opportunity: weed community dynamics 5.4.1 Phenotypic life history traits 5.4.1.1 Introduction 5.4.1.2 Plant morphology, development and activity during life history 5.4.1.3 Preadaptation 5.4.1.4 Trait basis of the invasion process 5.4.1.5 Life history trait lists: birth, dispersal, recruitment, vegetative growth, reproductive growth 5.4.2 Ecological roles-guilds-trades in weed-crop plant community assembly and structure 5.4.2.1 Definitions 5.4.2.2 Guild structure and community organization 5.4.2.3 Parameters of weed species ecological role and niche 5.4.2.4 Trait guild: Relative seedling/bud emergence order 5.4.2.5 Other trait guilds 5.4.3 Changes in plant community structure 5.4.3.1 Weed population shifts 5.4.3.1.1 Inter-specific weed population shifts 5 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants 5.4.3.1.2 Intra-specific weed population shifts 5.4.3.1.3 Herbicide-induced population shifts 5.4.3.2 Plant community ecological succession UNIT 4: WEED LIFE HISTORY 123 6 Weed Life History Models 125 6.1 Introduction 6.2 Limitations of demographic models 7 Reproduction, Mortality, Life History Timing and Trade-Offs 127 7.1 Introduction 7.2 Definitions 7.3 Reproduction and mortality 7.4 Timing of reproduction 7.5 Reproductive value 7.6 Risk of death 7.7 Plant age stages 7.8 Trade-offs in reproduction 7.8.1 Principle of strategic allocation 7.8.2 Life history trade-offs 7.8.3 Timing of life history trade-offs 7.8.4 Seed number 7.8.5 Seed age 7.8.5.1 Seed size plasticity and stability in a species 7.8.5.2 Variable seed size 7.8.5.3 Relationship of seed size to habitat 7.8.5.4 Small seed size 7.8.5.5 Relative size of seed produced by weed species 7.8.5.6 Seed weight variation between plant species 7.8.6 Seed size and number variation and trade-offs 7.8.7 Five roles of seed 7.8.7.1 Trade-offs among seed roles 7.8.7.2 Weed examples 8 Reproduction: Flowering, Birth, Embryogenesis and Abscission 141 8.1 Introduction 8.2 Flowering, anthesis and fertilization 8.2.1 Mating systems 8.3 Seed embryogenesis 8.3.1 Induction of dormancy in seeds during embryogenesis 8.3.1.1 Seed dormancy definitions 8.3.1.2 Life history of a seed 8.3.1.3 The evolutionary and ecological basis for seed dormancy and soil seed pools 6 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants 8.3.1.4 Variability and somatic polymorphism in weed seed dormancy 8.3.1.4.1 Definitions 8.3.1.4.2 Evolutionary ecology of seed heteroblasty 8.3.1.4.3 Examples of weed species heteroblasty 8.3.1.5 Observational categories of seed dormancy mechanisms 8.3.1.5.1 Non-dormant 8.3.1.5.2 Vegetative, perinating buds 8.3.1.5.3 Hard seed coats 8.3.1.5.4 Light and nitrate stimulated soil surface germination 8.3.1.5.5 Oxygen and water signal restriction of after-ripening and germination 8.3.1.5.6 Other seed dormancy mechanisms 8.3.1.5.7 Species with more than one dormancy type, interacting mechanisms 8.3.1.6 Beware of the seed dormancy literature 8.3.1.7 Summary of seed dormancy 9 Propagule Dispersal in Space and Time 155 9.1 Introduction 9.1.1 Definition 9.1.2 The evolutionary ecology of dispersal 9.1.3 Seed trade-offs 9.1.4 Cost of dispersal 9.1.5 Space-time dimensions of dispersal 9.2 Dispersal in space 9.2.1 Dispersal and post-dispersal processes 9.2.2 Seed flux at a locality 9.2.3 Modes of seed and propagule dispersal 9.2.3.1 Gravity 9.2.3.2 Wind and air 9.2.3.3 Water 9.2.3.4 Animal, non-human 9.2.3.5 Human 9.2.3.6 Other miscellaneous types 9.2.3.7 Anecdote on dispersal 9.3 Dispersal in time: soil seed and propagule pool formation 9.3.1 Introduction 9.3.2 Seed states and processes in the soil pool 9.3.2.1 Seed states and fates 9.3.2.2 Seed state transition processes 9.3.3 Population dynamics in the soil seed pool 9.3.3.1 Additions to the seed pool 9.3.3.2 Losses from the seed pool 9.3.3.3 Continuity in the seed pool with time 9.3.4 Structure of soil seed pools 9.3.4.1 Spatial distribution in the soil profile 7 Dekker: The Evolutionary Ecology of Weeds and Invasive Plants 9.3.4.2 Floral seed community composition 9.3.4.3 Seed pool size 9.3.4.4 Seed longevity in the soil 10 Seed Germination and Seedling Recruitment 169 10.1 Introduction 10.2 Process of recruitment 10.3 Favorable germination micro-sites 10.4 Patterns of seedling emergence 10.5 The relationship between seed dormancy heteroblasty and seedling recruitment timing 11 Community Interactions: Vegetative growth and development 173 11.1 Introduction 11.2 Spatial and temporal foraging 11.3 Plant density, plant form and community diversity 11.4 Influences of plant density and growth on yield 11.4.1 Density-yield response 11.4.2 Plant to plant variation 11.5 Influences of plant density on mortality 11.6 Influence of plant density on form and reproduction 11.6.1 Plant form and diversity of community 11.6.2 Phenotypic plasticity and somatic polymorphism 11.7 Forces of selection acting on plant community dynamics and competition 11.7.1 Biological categories of selection 11.7.1.1 ‘r’ and ‘K’ selection 11.7.1.2 Selection for ecological combining ability 11.7.1.3 Selection by activity of predators and pathogens 11.7.1.4 The evolutionary consequences of disturbances 11.7.1.5 Selection in a patchy environment 11.7.2 Maximizing fitness in a spatially variable environment 11.7.3 Biodiversity, complexity and community stability [vs.
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