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Evolutionary Ecology of Weeds

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Evolutionary Ecology of Weeds Evolutionary Ecology of Weeds Jack Dekker Weed Biology Laboratory Agronomy Department Iowa State University Ames, Iowa 50010 USA EMail: [email protected] WWW URL: www.public.iastate.edu/~jdekker 2011 ClassUse Version Copyright 1.1.11 Jack Dekker 1 Evolutionary Ecology of Weeds 2011 SHORT TABLE OF CONTENTS UNIT 1: THE NATURE OF WEEDS 13 1 The nature of weeds 14 1.1 What is a weed? 1.2 The definition of a weed 1.3 Weeds and human nature 1.4 Weedy traits 1.5 The origins of weeds 1.6 World origins and centers of agriculture, crop domestication and cultivation 1.7 World crop-weed species groups UNIT 2: THE EVOLUTION OF WEED POPULATIONS 45 2 Evolution, natural selection and weedy adaptation 47 2.1 Introduction 2.2 Evolution 2.3 Natural selection and elimination 2.4 The process of natural selection 2.5 Adaptation 3 Formation of the local weed population (deme): Precondition to natural selection 55 3.1 Introduction: opportunity and the formation of the local deme 3.2 The structure of local weedy opportunity 3.3 Habitat heterogeneity and dynamics 3.4 Limiting resources and pervasive conditions in local opportunity 3.5 The nature of plant invasions of local opportunity 4 Generation of genotypic and phenotypic variation: First process of natural selection 79 4.1 Genotypes and phenotypes 4.2 Generate genetic variation 4.3 Generate phenotypic variation 5 Survival, reproduction and inheritance: Second process of natural selection 97 5.1 Survive, Avoid Mortality 5.2 Reproduce the fittest, eliminate the others 5.3 Inheritance: transmit parental traits to offspring UNIT 3: ADAPTATION IN WEED LIFE HISTORY 114 6 Weed life history 115 6.1 Introduction to life history 6.2 Plant life history classification systems 6.3 Representing weed life history 6.4 The ecological demography of plant population life history dynamics 6.5 Evolutionary, trait-based, weed life history population dynamics 7 Reproductive adaptation 137 7.1 Introduction 7.2 Flowering, anthesis, fertilization and birth 7.3 Embryo adaptation: embryogenesis and dormancy induction 7.4 Propagule adaptation: post-abscission fecundity 8 Propagule dispersal in space and time 157 8.1 Introduction 8.2 Dispersal in space 8.3 Dispersal in time: formation of seed pools in the soil 8.4 Propagule germination and recruitment 2 Evolutionary Ecology of Weeds UNIT 4: ADAPTATION IN LOCAL PLANT COMMUNITIES 182 Weed-crop communities as complex adaptive systems 9 Neighbor interactions in local plant communities 186 9.1 Adaptation to neighbors in the community 9.2 The nature of neighbor interactions in the community 9.3 Strategic roles and traits of interference and facilitation with neighbors 9.4 Effects of neighborhood interactions on plant density, growth and form 10 Weed community structure, dynamics and biodiversity 209 10.1 Weed communities 10.2 Weed community structure 10.3 Exploiting opportunity: weed community dynamics 10.4 Weed community biodiversity 3 Evolutionary Ecology of Weeds 2011 TABLE OF CONTENTS & SYLLABUS Forward 9 Introduction 11 UNIT 1: THE NATURE OF WEEDS 13 1 The nature of weeds 14 Summary 1.1 What is a weed? 1.2 The definition of a weed 1.3 Weeds and human nature 1.4 Weedy traits 1.5 The origins of weeds 1.5.1 Weeds, planting and crop domestication 1.5.2 Biogeographic prehistory of agriculture 1.5.3 Pre-agricultural preadapted wild colonizing species 1.5.4 Wild-crop-weed complexes 1.6 World origins and centers of agriculture, crop domestication and cultivation 1.7 World crop-weed species groups UNIT 2: THE EVOLUTION OF WEED POPULATIONS 45 Introduction 2 Evolution, natural selection and weedy adaptation 47 Summary 2.1 Introduction 2.2 Evolution 2.2.1 Micro- and macroevolution 2.2.2 Units of evolution and natural selection 2.3 Natural selection and elimination 2.4 The process of natural selection 2.4.1 Population formation: Precondition to natural selection 2.4.2 Generate variation: Process of natural selection, step 1 2.4.3 Survival and reproduction: Process of natural selection, step 2 2.5 Adaptation 3 Formation of the local weed population (deme): Precondition to natural selection 55 Summary 3.1 Introduction: opportunity and the formation of the local deme 3.2 The structure of local weedy opportunity 3.2.1 Weedy habitats 3.2.2 Niches in the local community 3.2.3 The niche hypervolume 3.3 Habitat heterogeneity and dynamics 3.3.1 Spatial heterogeneity and patchiness 3.3.2 Temporal division of the environment 3.3.3 Disturbance 3.4 Limiting resources and pervasive conditions in local opportunity 3.4.1 Limiting resources 3.4.1.1 Light 3.4.1.2 Water 3.4.1.3 Mineral nutrients 3.4.1.4 Gases 3.4.2 Pervasive conditions in the environment 3.4.2.1 Heat 3.4.2.2 Terroir 3.4.3 Environmental signal spacetime 4 Evolutionary Ecology of Weeds 3.5 The nature of plant invasions of local opportunity 3.5.1 The plant invasion process: seizing, exploiting and occupying opportunity 3.5.2 Dispersal 3.5.3 Colonization 3.5.4 Enduring occupation of a locality 3.5.5 Extinction 3.5.6 The perception of plant invasion 4 Generation of genotypic and phenotypic variation: First process of natural selection 79 Summary 4.1 Genotypes and phenotypes 4.2 Generate genetic variation 4.2.1 Sources of genetic diversity 4.2.1.1 Forces increasing population variability 4.2.1.2 Forces decreasing population variability 4.2.2 Speciation 4.2.2.1 Process of speciation 4.2.2.2 Reproductive isolating mechanisms 4.2.2.3 Modes of speciation 4.3 Generate phenotypic variation 4.3.1 Phenotypic plasticity 4.3.2 Somatic polymorphism 5 Survival, reproduction and inheritance: Second process of natural selection 97 Summary 5.1 Survive, Avoid Mortality 5.2 Reproduce the fittest, eliminate the others 5.2.1 Timing of reproduction 5.2.2 Plant age and stage structure 5.2.3 Reproductive value 5.2.4 Risk of death determines life history 5.2.5 Influences of plant density on mortality 5.2.6 Modes of selection and population diversity 5.3 Inheritance: transmit parental traits to offspring 5.4 Mating system and inheritance 5.4.1 Mating system and opportunity 5.4.2 Evolution of mating systems 5.4.3 Sex classification systems 5.4.4 Types of mating systems 5.4.4.1 Self-pollenating species 5.4.4.2 Out-crossing species 5.4.4.3 Apomictic species 5.4.4.4 Vegetative clone reproducing species UNIT 3: ADAPTATION IN WEED LIFE HISTORY 114 Introduction 6 Weed life history 115 Summary 6.1 Introduction to life history 6.1.1 Phenotypic life history traits 6.1.2 Processes of life history adaptation 6.2 Plant life history classification systems 6.2.1 Life span 6.2.2 Growth and life form 6.2.3 Life history strategies 6.3 Representing weed life history 6.4 The ecological demography of plant population life history dynamics 6.4.1 Weed life history models 6.4.2 Demographic weed life history population dynamics models 6.4.3 Representation and information, inference and prediction 5 Evolutionary Ecology of Weeds 6.4.3.1 Representation and information 6.4.3.2 Inference 6.4.3.2.1 The deme 6.4.3.2.2 Life history development and behavior 6.4.3.2.3 Model formalization and measurement metrics 6.4.3.3 Predicting weed population dynamics 6.5 Evolutionary, trait-based, weed life history population dynamics 7 Reproductive adaptation 137 Summary 7.1 Introduction 7.2 Flowering, anthesis, fertilization and birth 7.2.1 Parental plant architecture 7.2.2 Mating systems 7.3 Embryo adaptation: embryogenesis and dormancy induction 7.3.1 Induction of seed dormancy 7.3.2 The evolutionary ecology of seed dormancy 7.3.3 Weed seed dormancy variability and somatic polymorphism 7.3.4 Evolutionary ecology of seed heteroblasty 7.3.5 Weed species seed heteroblasty examples 7.3.6 Observable seed dormancy-germinability regulation life forms 7.3.6.1 Non-dormant 7.3.6.2 Vegetative, perinating buds 7.3.6.3 Environmental seed germination control mechanisms 7.3.6.4 Hard, gas- and water-impermeable, seed envelope germination inhibition 7.3.6.5 Light-phytochrome and nitrate stimulated germination 7.3.6.6 Species with multiple interacting germination control mechanisms 7.3.6.7 Other seed germination control mechanisms 7.3.7 Experimental weed seed science 7.4 Propagule adaptation: post-abscission fecundity 150 7.4.1 Five roles of seeds 7.4.2 Principle of strategic allocation 7.4.3 Trade-offs among seed roles 7.4.4 Seed size trade-offs 7.4.4.1 Seed size plasticity and stability 7.4.4.2 Relationship of seed size to habitat 8 Propagule dispersal in space and time 157 Summary 8.1 Introduction 8.1.1 The evolutionary ecology of dispersal structures 8.1.2 Seed dispersal trade-offs 8.1.3 Cost of dispersal 8.1.4 Space-time dimensions of dispersal 8.2 Dispersal in space 8.2.1 Dispersal and post-dispersal processes 8.2.2 Seed flux at a locality 8.2.3 Modes of seed and propagule dispersal 8.2.3.1 Gravity 8.2.3.2 Wind and air 8.2.3.3 Water 8.2.3.4 Animal, non-human 8.2.3.5 Human 8.2.3.6 Other modes of dispersal 8.3 Dispersal in time: formation of seed pools in the soil 8.3.1 Adaptative roles of soil seed pools 8.3.2 Population dynamics in the soil seed pool 8.3.2.1 Life history of a seed 6 Evolutionary Ecology of Weeds 8.3.2.2 Seed states, fates and seed state transition processes 8.3.2.3 Seed pool additions, losses and continuity 8.3.3 Structure of soil seed pools 8.3.3.1 Spatial distribution in the soil profile 8.3.3.2 Floral seed community compostion 8.3.3.3 Seed pool size 8.3.3.4 Seed longevity in the soil 8.4 Propagule germination and recruitment 8.4.1 Introduction 8.4.2 Process of recruitment 8.4.3 Germination micro-sites and safe sites 8.4.4 Patterns of seedling emergence 8.4.5 Relationship
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    ISTA List of Stabilized Plant Names th 7 Edition ISTA Nomenclature Committee Chair: Dr. M. Schori Published by All rights reserved. No part of this publication may be The Internation Seed Testing Association (ISTA) reproduced, stored in any retrieval system or transmitted Zürichstr. 50, CH-8303 Bassersdorf, Switzerland in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior ©2020 International Seed Testing Association (ISTA) permission in writing from ISTA. ISBN 978-3-906549-77-4 ISTA List of Stabilized Plant Names 1st Edition 1966 ISTA Nomenclature Committee Chair: Prof P. A. Linehan 2nd Edition 1983 ISTA Nomenclature Committee Chair: Dr. H. Pirson 3rd Edition 1988 ISTA Nomenclature Committee Chair: Dr. W. A. Brandenburg 4th Edition 2001 ISTA Nomenclature Committee Chair: Dr. J. H. Wiersema 5th Edition 2007 ISTA Nomenclature Committee Chair: Dr. J. H. Wiersema 6th Edition 2013 ISTA Nomenclature Committee Chair: Dr. J. H. Wiersema 7th Edition 2019 ISTA Nomenclature Committee Chair: Dr. M. Schori 2 7th Edition ISTA List of Stabilized Plant Names Content Preface .......................................................................................................................................................... 4 Acknowledgements ....................................................................................................................................... 6 Symbols and Abbreviations ..........................................................................................................................
  • Allium Paradoxum (M.Bieb.) G. Don (Amaryllidaceae) – a New Invasive Plant Species for the Flora of Baltic States

    Allium Paradoxum (M.Bieb.) G. Don (Amaryllidaceae) – a New Invasive Plant Species for the Flora of Baltic States

    Acta Biol. Univ. Daugavp. 20 (1) 2020 ISSN 1407 - 8953 ALLIUM PARADOXUM (M.BIEB.) G. DON (AMARYLLIDACEAE) – A NEW INVASIVE PLANT SPECIES FOR THE FLORA OF BALTIC STATES Pēteris Evarts-Bunders, Aiva Bojāre Evarts-Bunders P., Bojāre A. 2020. Allium paradoxum (M. Bieb.) G. Don – a new invasive plant species for the flora of Baltic States.Acta Biol. Univ. Daugavp., 20 (1): 55 – 60. Allium paradoxum (M. Bieb.) G. Don was recorded as a new species for the flora of Latvia and the Baltic States on the basis of plant material first collected by A. Bojāre and P. Evarts-Bunders in 2020. Relatively large population of this species was found in Rīga, in the Rumbula district (Eastern border of the city), on the slope of the river Daugava covered by natural vegetation, but near the private gardens zone and ruderal places. This species can be easily distinguished from other Allium species by one flat, 5-25 mm wide, keeled leaf per bulb, inflorescence with several bulbils and mostly with only one flower. The species is considered as invasive and spreading by means of bulbils. In Latvia the species has been identified in their typical habitat – disturbed forest and shrubland along the riverbank on damp soil. Key words: Allium paradoxum, Latvia, Baltic States, invasive plant, flora. Pēteris Evarts-Bunders. Institute of Life Sciences and Technology, Daugavpils University, Parādes str., 1A, Daugavpils, LV-5401, Latvia; [email protected] Aiva Bojāre. National Botanical garden, Dendroflora Department, Miera Str., 1, Salaspils, LV-2169, Latvia; [email protected] INTRODUCTION There are seven species of sedge occurring in the wild in Latvia – Allium angulosum Allium L.