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The structure of community ecology Mark Vellend Departments of Botany & Zoology Biodiversity Research Centre University of British Columbia Is community ecology a mess? 1 …with a key difference in how process is treated Patterns we want to understand Population Genetics Community Ecology Diversity and composition of alleles/genotypes Diversity and composition of species Processes to Mutation explain Drift patterns Migration Selection Competition Predation Niches Neutrality Island biogeography Resource-use trade-offs Colonization-competition trade-offs Disturbance Productivity Evolutionary history Etc. But is the difference real or necessary? Patterns we want to understand Population Genetics Community Ecology Diversity and composition of alleles/genotypes Diversity and composition of species Processes to Mutation explain Drift patterns Migration Selection Speciation Drift Dispersal Selection 2 Question on Ecology 101 final exam: What processes influence species composition and diversity? Answer: Question on Evolution 101 final exam: What processes influence genetic variation? Answer: Mutation Drift Migration Selection 3 Processes that influence community composition and diversity Speciation: The creation of new species. Drift: Random changes in the relative frequencies of species. Dispersal: Movement of organisms/propagules among communities. Selection: Deterministic differential survival or reproduction of species. THE LAST 40 YEARS OF COMMUNITY ECOLOGY The MacArthur school (Marlboro Circle) Simple, general, deterministic models Compare equilibrium to data – draw inferences Island Biogeography That’s fraught with pattern-process problems Most of the world is not at equilibrium most of the time (Strong, Simberloff criticize “assembly rules”) SELECTION Lawton’s “traditional” Manipulative field experiments community ecology Patch dynamics and disequilibrium Neutral theory: the ultimate null model? DRIFT Better “null” models for community assembly studies Spatial ecology DISPERSAL Niche-neutral reconciliation Metacommunities Experiments often give good answers to trivial questions, are system specific and local Back to history and geography Macroecology SPECIATION (Ricklefs’ regional effects) Better inferential tools Metabolic theory (community ecology?) (ML, Bayes) 4 Conceptual umbrella Key contribution Competition theory Experimental ecology SELECTION Regional & historical effects SPECIATION (+ DISPERSAL) Neutral theory DRIFT Metacommunities DISPERSAL The Structure of Community Ecology Morin (1999) Community Ecology Putman (1993) Community Ecology I: Communities, Patterns and Processes: Competition I, Competition II Predation I, Predation II Food Webs Mutualisms Indirect Effects II: Factors Influencing Interactions: Temporal Patterns Habitat Selection Spatial Dynamics III: Large Scale Phenomena: Causes and Consequences of Diversity Succession Applied Community Ecology Ecological communities - definitions and a search for pattern. Population interactions Food webs and connectance. Compartments in food webs. Food web topology. Niche theory Guilds and guild structure. Spp composition, community assembly A question of equilibrium. Stability. 5 The Structure of Community Ecology Morin (1999) Community Ecology Putman (1993) Community Ecology Commonalities Ecological communities - definitions and a search for pattern. CommunityPopulation patterns interactions Food webs and connectance. Space and time Compartments in food webs. Competition Food web topology. Niche theory Predation Guilds and guild structure. Spp composition, community assembly Food webs A question of equilibrium. Stability. Niches I: Communities, Patterns and Processes: Competition I, Competition II Predation I, Predation II Food Webs Mutualisms Indirect Effects II: Factors Influencing Interactions: Temporal Patterns Habitat Selection Spatial Dynamics III: Large Scale Phenomena: Causes and Consequences of Diversity Succession Applied Community Ecology Logical difficulties with this organization Community patterns Space and time Competition Predation Food webs Niches Pattern Consideration Process Process Concept/thing Concept 6 The Structure of Population Genetics Hedrick (2000) Genetics of Populations Hartl & Clark (2007) Principles of Pop. Genetics General Background The Diversity of Genetic Variation Measures of Genetic Variation Selection: An Introduction Selection: Advanced Topics Inbreeding Genetic Drift Mutation Neutral Theory, Coalescence Gene Flow, Population Structure Linkage Disequilibrium, Recombination Molecular Genetics and Evolution Genetic & Phenotypic Variation Organization of Genetic Variation Random Genetic Drift Mutation & the Neutral Theory Darwinian Selection Inbreeding, Pop. Subdivision, Migration Molecular Population Genetics Evolutionary Quantitative Genetics Population Genomics Human Population Genetics The Structure of Population Genetics Hedrick (2000) Genetics of Populations Hartl & Clark (2007) Principles of Pop. Genetics Gene Flow, Population Structure Linkage Disequilibrium, Recombination Molecular Genetics and Evolution Human Population Genetics Commonalities General Background & Phenotypic Variation MeasurementGenetic of diversity The Diversity of Genetic Variation Organization of Genetic Variation Measures of Genetic Variation Mutation Random Genetic Drift Selection: An Introduction Mutation & the Neutral Theory Selection: Advanced Topics Drift Darwinian Selection Inbreeding Inbreeding, Pop. Subdivision, Migration Genetic Drift Migration Molecular Population Genetics Mutation Evolutionary Quantitative Genetics Neutral Theory, Coalescence Selection Population Genomics 7 Logical consistency of this organization Diversity measures Mutation Drift Migration Selection Pattern Process Process Process Process Why the difference? The architects of the modern synthesis were not distracted by data! (Vieulle and Slatkin 2002 ) 8 Ecology is a discipline “whose material study is part of everyday encounters: birds, bees, trees, and rivers”. “It is, however, a mistake to imagine that this familiarity makes ecology and easy pursuit…the very familiarity of ecological objects presents the difficulties”. (Allen & Hoekstra 1992, Toward a Unified Ecology) A Different Structure for Community Ecology Primary patterns Processes • Drift • Speciation • Dispersal • Selection (across space & time) • Species diversity • Species composition Abiotic env. Biotic interactions • Competition • Predation • Etc. (identity, traits, abundances) Emergent patterns • Productivity • Stability • Food web connectance • Whatever you can think of 9 Relating the many to the few Ecological idea/model/theory Processes Island biogeography Dispersal and drift Resource competition Selection (frequency dependent) Metacommunity theory Dispersal, selection, and drift Neutral theory Speciation and drift Assembly rules Dispersal and selection Spatial/temporal heterogeneity Selection (variable in space/time) Food webs Selection (frequency dependent) Species pool theory Speciation Productivity/stress/disturbance Selection (and speciation) The Evolutionary Modern Synthesis (Stebbins 1971, modified by Kutschera & Niklas 1994) Dispersal A (not so) Modern Synthesis for Community Ecology? Speciation Drift Dispersal (Selection) Selection & Drift Selection Selection (Ricklefs & Schluter 1993) 10 Global community Speciation Drift Selection Regional community Dispersal Dispersal Speciation Drift Selection Dispersal Dispersal Local Community (Speciation) Drift Selection Note: Extinction results from drift & selection Teaching Community Ecology Processes • Drift • Speciation • Dispersal • Selection Primary patterns (across space & time) • Species diversity • Species composition (identity, traits, abundances) Advantages (I hope): • Logical coherence Emergent patterns • Allow links among sub-fields to be seen more clearly • Productivity • Stability • Food web connectance • Whatever you can think of • Facilitate communication of the conceptual core of the discipline more clearly 11