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Community Development and Species-Area Relationships BIOL400 2 November 2015 Community Development: Ecological Succession Primary vs. Secondary Succession Primary succession occurs on bare soil Ex: Mt. Saint Helens Secondary succession follows disturbance in areas having some vegetation already Ex: Old-field succession Figs. 18.10—18.12 pp. 366-67 Clements’ View of Succession Community-asSuperorganism Concept • aka Relay Floristics Model Seres (= successional stages) analogous to developmental stages of an organism Monoclimax: single, predictable climax community (the community that remains stable, ending succession) Fig. 18.5 p. 358 Egler's Initial Floristic Composition Model Much more individualistic Contingency: what plant species happen to colonize first will greatly influence both the sequence and the climax Inhibition of earlier stages by later stages Connell and Slatyer's Facilitation, Inhibition, and Tolerance Model drives Clements’ relay floristics model Facilitation • A modifies environment favorably for B, etc. drives Egler’s initial floristic composition model Inhibition • B suppresses growth of A, etc. Tolerance refers to the ability of a species at any one stage to tolerate… • …resource levels (important early in sequence) • …level of competition for soil nutrients and light (important later in sequence) Succession and Plant Life Histories Whether a species predominates in early vs. late succession tends to correlate with features of its ecology and life history Ability to colonize Ability to tolerate competition with other species Table 18.1 p. 359 Fig. 18.7 p. 364 The Climax Community Climax vegetation is stable and self-perpetuating Clements—monoclimax Tansley—advocated polyclimax idea Historical contingency determines climax composition Whittaker—developed idea of pattern climax— a continuum of climax-condition possibilities based on entire suite of physical factors Other Concepts of Ecological Succession Communities Ex: that do not climax Prairies, which remain stuck in early sere because of frequency of disturbance by fire Transient succession—no climax possible due to gradually diminishing resources Exs: Dung piles Carcasses Fig. 18.14 p. 368 Fig. 18.15 p. 369 HANDOUT—Lundsten et al. 2010 Species-Area Relationships: Island Biogeography Arrhenius equation (1921) S = cAz • • • • S = species richness A = island area z = slope c = constant Linear form: logS = logc + zlogA Explaining the Arrhenius Species-Area Relationship Increased area = increased habitat heterogeneity (more niches to fill) Increased area = increased population sizes of resident species Less chance of their stochastic local extinction Increased area = larger “target” for dispersing would-be colonizing species Figs. 21.19 & 21.20 p. 442 c depends on units of area that are used, but z does not Typically, z ranges between 0.20 and 0.35 z = 0.32 z = 0.30 The Theory of Island Biogeography 1967 book by Robert A. MacArthur (Princeton) and Edward O. Wilson (Harvard) Colonization curve: declines with rising S, as fewer species are available to colonize Local extinction curve: rises with rising S, as interspecific interactions intensify Fig. 21.21 p. 442 The Dynamic Equilibrium Point at which the two lines cross Balance of new colonists arriving and resident species going locally extinct Gains = Losses Fig. 21.22 p. 443 Krakatau Indonesian island wiped clean of all life by enormous volcanic eruption in 1883 1883: 0 bird spp. 1908: 13 bird spp. 1921: 2 bird spp. lost, 16 gained (27 total) 1935: 5 bird spp. lost, 5 gained (27 total) MacArthur and Wilson’s Area Effect Larger areas house larger resident populations less prone to stochastic extinction Fig. 21.22 p. 443 MacArthur and Wilson’s Distance Effect More isolated habitat islands are more difficult to reach Fig. 21.22 p. 443 Area and Distance Effects Combined Small, isolated islands: Rare colonists High local extinction rate Low Sequilibrium Large, nearby islands: Frequent colonists Low local extinction rate High Sequilibrium Fig. 21.22 p. 443 HANDOUT Simberloff and Wilson 1969 Nonequilibrium Island Biogeography Dynamic equilibria are possible only when colonization can occur “Stranded” populations may show ecosystem decay, with an area effect on S but no distance effect on S Islands slowly “bleed” species richness More rapid loss for smaller islands Occurs naturally, but esp. important in anthropogenically-isolated habitats HANDOUT—Brown 1971 HANDOUT—Diamond 1984 HANDOUT—Newmark 1987