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Mechanistic models for macroecolgy: moving beyond correlation. Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405. ?? What causes geographic variation in species richness ??. Understanding species richness patterns. Data sources
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Mechanistic models for macroecolgy: moving beyond correlation Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405
Understanding species richness patterns • Data sources • A critique of current methods • Range cohesion and the mid-domain effect • Mechanistic models for species richness • Model selection • Summary
Gary EntsmingerAcquired Intelligence Rob ColwellUniversity of Connecticut Nicholas Gotelli, University of Vermont Thiago Rangel Federal University of Goiás Carsten RahbekUniversity of Copenhagen Gary GravesSmithsonian
Understanding species richness patterns • Data sources • A critique of current methods • Range cohesion and the mid-domain effect • Mechanistic models for species richness • Model selection • Summary
Data sources • Gridded map of domain
Avifauna of South America “There can be no question, I think, that South America is the most peculiar of all the primary regions of the globe as to its ornithology.” P.L. Sclater (1858)
South American Avifauna • 2891 breeding species • 2248 species endemic to South America and associated land-bridge islands
Minimum: 18 species
Maximum: 846 species Minimum: 18 species
Data sources • Gridded map of domain • Species occurrence records within grid cells
Anas puna Geographic Ranges For Individual Species Phalacrocorax brasilianus Myiodoorus cardonai
Geographic Ranges Species Richness
Geographic Ranges Species Richness
Data sources • Gridded map of domain • Species occurrence records within grid cells • Quantitative measures of potential predictor variables within grid cells (NPP, temperature, habitat diversity)
Understanding species richness patterns • Data sources • A critique of current methods • Range cohesion and the mid-domain effect • Mechanistic models for species richness • Model selection • Summary
How are these macroecological data typically analyzed?Curve-fitting!
Criticisms of Curve-Fitting • “Correlation does not equal causation”
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology!
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology! • Non-linearity & non-normal, spatially correlated errors
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology! • Non-linearity & non-normal, spatially correlated errorsLOESS, Poisson, Spatial Regression (SAM)
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology! • Non-linearity & non-normal, spatially correlated errorsLOESS, Poisson, Spatial Regression (SAM) • Choosing among correlated predictor variables
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology! • Non-linearity & non-normal, spatially correlated errorsLOESS, Poisson, Spatial Regression (SAM) • Choosing among correlated predictor variablesModel selection strategies, stepwise regression, AIC
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology! • Non-linearity & non-normal, spatially correlated errorsLOESS, Poisson, Spatial Regression (SAM) • Choosing among correlated predictor variablesModel selection strategies, stepwise regression, AIC • Sensitivity to spatial scale, taxonomic resolution, geographic range size
Criticisms of Curve-Fitting • “Correlation does not equal causation”Common to all of macroecology! • Non-linearity & non-normal, spatially correlated errorsLOESS, Poisson, Spatial Regression (SAM) • Choosing among correlated predictor variablesModel selection strategies, stepwise regression, AIC • Sensitivity to spatial scale, taxonomic resolution, geographic range sizeStratify analysis
Predicted Species Richness (S / grid cell) Observed Species Richness (S / grid cell) Conceptual Weakness of Curve-Fitting Paradigm Potential Predictor Variables (tonnes/ha, C°)
Predicted Species Richness (S / grid cell) Observed Species Richness (S / grid cell) Conceptual Weakness of Curve-Fitting Paradigm Potential Predictor Variables (tonnes/ha, C°) minimizeresiduals
Predicted Species Richness (S / grid cell) Observed Species Richness (S / grid cell) Conceptual Weakness of Curve-Fitting Paradigm Potential Predictor Variables (tonnes/ha, C°) minimizeresiduals ??MECHANISM??
Predicted Species Richness (S / grid cell) Observed Species Richness (S / grid cell) Alternative Strategy:Mechanistic Simulation Models Potential Predictor Variables (tonnes/ha, C°) ExplicitSimulationModel
Predicted Species Richness (S / grid cell) Observed Species Richness (S / grid cell) Alternative Strategy:Mechanistic Simulation Models Potential Predictor Variables (tonnes/ha, C°) mechanism ExplicitSimulationModel
How can we build explicit simulation models for macroecology?
Understanding species richness patterns • Data sources • A critique of current methods • Range cohesion and the mid-domain effect • Mechanistic models for species richness • Model selection • Summary
One-dimensional geographic domain Species geographic ranges randomly placed line segments within domain
One-dimensional geographic domain Species geographic ranges randomly placed line segments within domain Peak of species richness in geographic center of domain
Species Number One-dimensional geographic domain Species geographic ranges randomly placed line segments within domain Peak of species richness in geographic center of domain
geographic range domain
Pancakus spp. der Pfankuchen Guild
Mid-domain peak of species richness in the center of the domain
2-dimensional MDE Model • Random point of originationwithin continent (speciation) • Random spread of geographicrange into contiguousunoccupied cells • Spreading dye model (Jetz & Rahbek 2001) predicts peak richness incenter of continent (r2 = 0.17)
Assumptions of MDE models • Placement of ranges within domain is random with respect to environmental gradients • Controversial, but logical for a null model for climatic effects
Assumptions of MDE models • Placement of ranges within domain is random with respect to environmental gradients • Controversial, but logical for a null model for climatic effects • Geographic ranges are cohesive within the domain • Rarely discussed, but important as the basis for a mechanistic model of species richness