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Diversity and Stability By Ariana Strandburg-Peshkin QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Does Increased Diversity Lead to Increased Stability? What is stability? - MacArthur • Abundances of species staying relatively constant – Perturb one species --> Others relatively unaffected • The amount of choice which the energy has in following paths up through the food web Complexity & Stability - MacArthur It’s Intuitive! QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. VS. Implications of MacArthur’s Stability Definition • Stability can be increased by… – Increasing the number of links – Increasing the number of species • Complexity --> Stability is implied in the definition • Evolution optimizes for maximum efficiency subject to a certain necessary stability QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Ecological Implications, if this is true… • Small number of species --> hard to fulfill stability requirement --> must diversify diet • Large number of species --> stability requirement easily fulfilled --> can specialize Along Comes Robert May… I know math. You’re all wrong! QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. • In mathematical models, complexity tends to lead to instability Lotka-Volterra Models - May Lotka-Volterra Models - May Mathematical Models - May • Randomly-Constructed Community Matrices – Too rich web connectance --> instability – Too large average interaction strength --> instability Tilman’s Study • Effects of Nitrogen addition to soil • 207 plots of land, 9 different nutrient treatments • Sampled biomass every year • Major droughts in 1987 and 1988 Tilman - Results • Species-rich plots more drought resistant Tilman - Results • Total biomass variability (c.v.) negatively correlated with SR (both drought and non-drought) Tilman - Results • Biomass of individual species varied more in SR plots Reconciling MacArthur and May Stop arguing! You’re both right! Quic kTime™ and a TIFF (Unc ompres sed) dec ompres sor are needed to see this pic ture. • For COMMUNITIES diversity --> stability • For SPECIES diversity --> instability • But wait… How Can This Be?? How can this be?? • Competitive Release / Compensatory Growth • Monocultures - Avg. response to perturbation = Avg. of individuals • Mixtures - Avg. response to perturbation includes compensatory growth • Mixtures do better --> Species-rich plots do better b/c higher probability of perturbation resistant plants What About Omnivores? • Omnivore - species that feeds on more than one trophic level • Classical Mathematical Results: Omnivory --> Instability • Fagan: Omnivory --> Stability Fagan - Experiment • Studied arthropod communities • Manipulated level of omnivory • Perturbed populations Fagan - Results • Omnivory decreases coefficient of variation after a perturbation • Probably due to rapid response of omnivores to changing pressures References • MacArthur, R.H. Fluctuations of animal populations and a measure of community stability. Ecology 36, 533-536 (1955) • May, R.M. Stability and Complexity in Model Ecosystems (Princeton Univ. Press, 1973). • Tilman, D. Biodiversity: population versus ecosystem stability. Ecology 77, 350-363 (1996). • Fagan, W.F. Omnivory as a stabilizing feaure of natural communities. Am. Nat. 150, 554567 (1997).