LECTURE 21 CH 18 + 19: COMMUNITY STRUCTURE + DEVELOPMENT MAJOR CONCEPTS 1. Ecologists hold diverse concepts of communities. Some have a holistic view; others an individualistic view of the nature of the community. 2. Gradient analysis usually shows support for the individualistic view; interactions support holistic view. 3. Communities change gradually along a continuum unless an ecotone sharply divides communities. 4. Communities may have tight connection among species – or redundant species with less strong connection. 5. Feeding relationships organize communities in food webs that can be quantified with multiple measures. 6. The activities of a keystone species may control the structure of communities. 7. Debate continues about whether greater food web complexity leads to stability. 8. Abundance within trophic levels is influenced from above by predation (top-down control) and from below by production (bottom-up control). 9. Communities can switch between alternative stable states. 10. Communities vary in whether or not they are in equilibrium. Disturbance keeps a community from reaching equilibrium 11. Qualities of disturbances vary greatly among communities. 12. Primary succession is more slow than secondary succession because of the lack of soil and seeds and harsh abiotic conditions. 13. Life history differences influence rank order of species entry during succession. 14. Various mechanisms govern succession: facilitation, inhibition, tolerance Define community: Assemblage of interacting species (populations) occurring in same area Often named after conspicuous members Questions: Spatially-defined communities Ecotone: sharp boundary between communities; caused by abrupt changes in physical feature (e.g. soil type) 18.4, 18.5 Growth on poor soils may be do tolerance or inability to compete on better soils 18.6 What is the ‘nature’ of the community? Is it a natural unit of organization? pg. 369-370 Pro: Holistic view 18.3a Closed with discrete boundaries If true, then can focus on classifying communities Con: Individualistic view 18.3b, 18.7 Haphazard assemblages of species Open with no natural boundaries; each species has unique range If true, then focus on community dynamics and functional organizatio Continuum concept: species replace one another continuously along physical gradients 18.7 Gradient analysis: most studies support continuum, hence open communities 18.9 (Interactions support holistic nature of communities) Questions: Functionally-defined in terms of interactions How strong are connections among species? Rivet model Tight linkage among species Obligate associations or exclusions Conservation implications: requires community dynamics approach Expect removal of one species to have cascading effects Redundancy model Loose linkage among species Most species have little impact on other species Multiple species can play same ecological role If remove one species, another takes over Feeding relationships organize communities in food webs 18.11 Effect of species richness on food web structure What changes with increased food web complexity? 18.12 Number of trophic levels Number of guilds per trophic level Number of interactions/species (doesn’t change) Connectance = # interactions/total possible = # interactions/[S(S-1)] Linkage density = # interactions/ # species Variety of food web types Connectedness, energy flow, function webs 18.14 Effect of food web structure on species diversity What is the role of keystone species in communities? 18.13 Non-redundant species that maintains community stability and diversity System-specific: Can be plant, herbivore, or predator What controls abundance within each trophic level? 18.19 Top-down control (by predators) Bottom-up control (by nutrients, plant productivity, plant defense) Trophic cascade: ripple effects from top level through multiple trophic levels 18.23, 24, 25 Indirect effects on lower levels; abundance alternates at each trophic level Does greater food web complexity contribute to community stability? 18.15 Constancy (resist change); Pros: Alternative resources; less dependent on fluctuations in any one resource Energy can take many routes; disrupt one pathway shunts more energy to another Cons: More links creates widespread, destabilizing time lags in pop. processes Resilience (return to prior state after disturbance) 18.17 Does species diversity help communities to bounce back from disturbance? Communities can switch between alternative stable states. 18.18 Summary Chap 18: 1-5; 7-12 CHAPTER 19: ECOLOGICAL SUCCESSION AND COMMUNITY DEVELOPMENT Are communities stable (in equilibrium) or transient (in non-equilibrium)? Historically: emphasis on equilibrium view Current: raises importance of disturbance and non-equilibrium view Not ‘either-or’; rather under what circumstances does one or other prevail? Importance for conservation: easier to conserve a predictable equilibrium community? Disturbance keeps communities in non-equilibrium state. Differ in size, frequency, intensity, level of environmental heterogeneity, and remnant species and seed sources. Succession Directional replacement of species after disturbance Sere – all stages of successional change 19.2 Succession follows as colonists alter environmental conditions Theoretically ends in climax community that is self-replacing pg. 398-9; 19,13 Primary Succession pg. 392-393 On new bare substrate; no soil 19.4, 19.5 No seed bank; seeds dispersed from outside 19.11 Extreme abiotic conditions Very slow Secondary Succession 19.3 After disturbance to existing vegetation Soil remains; seed bank may remain Life history differences influence species rank order of entry during succession 19.10 r-selected pioneer species in early succession Table 19.2 K-selected species in late succession 19.12 Mechanisms governing succession Facilitation – (drives ‘relay floristics’); common in primary succession 19.9 Early species modify environment in ways that later-stage species can establish Inhibition – can lead to ‘arrested succession’ ‘Jump-starting’ succession in restoration ecology Tolerance – (explains ‘initial floristics’); passive Animal communities follow stages of plant succession Summary: 1-8