BIOS 6150: Ecology Dr. Stephen Malcolm, Department of Biological Sciences • Week 13: Community Structure. • Lecture summary: • What is a community: • Chapters 17 & 24 in Begon, et al. (1996) 3rd ed. • Chapters 16 & 21 in 4th ed. Pine forest, Sweden (Begon et al. 1996) • Species diversity. • Succession. • Patterns of community structure. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Beech forest, Scotland (Begon et al.1996) Slide - 1 2. What is a community?: • The community is an assemblage of species populations that occur together in space and time. • Ecological interest is in: • How assemblages of species are distributed. • Ways in which interactions among species influence group assembly. • Community-level properties of assemblages. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 2 3. Community properties: • Collective properties: • Include species diversity, community biomass & productivity. • Emergent properties: • Product of population-level processes such as predation, herbivory, parasitism and mutualism. • Not evident from simply summing the interactions: • Like limits to similarity of competing species or food web stability when disturbed. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 3 4. Nested hierarchy of habitats: • Recognition of community patterns has dominated community ecology at different levels of scale (Fig. 16.2). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 4 5. Species diversity: • Counting numbers of species: • Ideally should be exhaustive as in Fig. 16.3 but is comparable only if based on similar sampling effort (time or area). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 5 6. Diversity indices: • Simply counting species ignores rarity and abundance of individuals within species. • Simpson's diversity index includes: • Richness: • Number of species • Equitability: • Proportion of individuals or biomass that each species contributes to total in the sample. • D = 1/ΣPi2, (for i = 1 to S) • where Pi is proportion of numbers or biomass contribution by ith species to the total number of species (S) in the community sample (richness). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 6 7. Change in Shannon index with time: • Fig. 16.4: Species diversity (H) and equitability (J) of control and fertilized plots in Rothamsted ‘parkgrass’ experiment. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 7 8. Rank abundance plots: • Relationship between equitability and richness (Fig. 16.5) with full array of ranked Pi values. • (a, b) rank-abundance patterns of different models; (c) change in Rothamsted grassland with continuous fertilizer application. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 8 9. Community patterns in space: • 1. Gradient analysis: • Distribution of species (Fig. 16.6) according to abiotic conditions (overlap in abundances over gradients, not sharp boundaries). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 9 10. Distribution gradients: • Fig. 17.6 (3rd ed.): (a) grasses along pH gradient, (b) macrofauna in oyster bed. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 10 11. Community patterns in space • 2. Ordination: • Organization of communities by both species composition & relative abundance (Fig. 17.7, 3rd ed.): • Examine patterns and correlations to develop hypotheses and controlled experiments to determine what processes most contribute to these patterns. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 11 12. Community patterns in space • 3. Classification: • Organization by similarity (Fig. 17.8) • see Fig 16.7 in 4th ed. • Both ordination and classification are objective and help to understand community associations and perturbations. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 12 13. Community patterns in time Succession: • Begon et al., (p 479) define succession as: • “the nonseasonal, directional and continuous pattern of colonization and extinction on a site by species populations.” • Succession describes the process of community change from: • Colonization of a new habitat (e.g. sandbar, volcanic island, etc.) by the “pioneer community”, to, • Steady state called a “climax community”, in which changes occur continually but they tend to perpetuate the same community structure rather than alter it. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 13 14. Classification of successional sequences within communities: • Degradative Succession: • Occurs over short time scales via heterotrophic organisms exploiting units of dead organic matter (feces, carrion, detritus etc.) until the resource is completely utilized: • Fig. 17.9 (3rd ed.). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 14 15. Classification of successional sequences within communities • Allogenic Succession: • Result of changing, external geophysicochemical forces which alter conditions (abiotic) such as silt deposition in estuaries and marshes (Fig. 17.10, 3rd ed.) BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 15 16. Classification of successional sequences within communities • Autogenic Succession: • Result of internal biological processes which can modify both conditions and resources (biotic). • Primary (new habitat, e.g. after glacial retreat - Fig. 17.11, 3rd), or, • Secondary (e.g. old field after agricultural abandonment). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 16 17. Secondary, autogenic succession: • Fig. 17.12 (3rd ed.): Marine algal colonization of concrete blocks BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 17 18. Secondary, autogenic succession: • Fig. 17.13 (3rd ed.): (a) Effect of removal of Ulva on Gigartina recruitment; (b) Algal survivorship curves. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 18 19. Old field secondary, autogenic succession Annual weeds Herbaceous perennials Shrubs • Fugitives. • Germinate under high light intensity. • Shade intolerant. • r-selected. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Early success -ional trees Late success -ional trees • Germinate under low light intensity. • Shade tolerant. • K-selected. Slide - 19 20. Life history characteristics of early and late successional plants (Table 17.4, 3rd ed.): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 20 21. Community changes with time: • Fig. 17.15 (3rd ed.): • Changes with time since abandonment of old fields. (a) bare soil, (b) annuals, (c) herbaceous perennials, (d) woody plants, (e) change in soil nitrogen. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 21 22. Tilman’s resource-ratio hypothesis of succession (Fig. 16.14): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 22 23. Three successional mechanisms: • Facilitation: • Early species make habitat more suitable for new species. • Tolerance: • Different species exploit resources in different ways. • Inhibition: • Invasions resisted by habitat modification: • e.g. allelopathy. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 23 24. Connell & Slatyer’s overview of successional mechanisms (Fig. 17.16, 3rd ed.): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 24 25. Simple patterns in community structure: • Why do some communities contain more species than others? • Are there patterns or gradients of species richness? • If so, what are the reasons for these patterns? • “Geographical”, “Climatic”, “Independent” and “Biological” reasons. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 25 26. Species richness & resource availability: • Richness related to resource availability axis (R) as in Fig. 21.1 that is partitioned (saturated) according to niche breadths (n) that overlap by varying amounts (o) as a measure of specialization. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 26 27. Dr Seuss, nitches and nutches: Geisel, T.S. (“Dr. Seuss”) 1955. On beyond Zebra. Random House Publishing, New York. http://shade.grove.iup.edu/~rgendron/Seuss.htmlx BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 27 28. Richness relationships: • Resource productivity more productive environments have more species (Fig. 24.2, 3rd ed.) with narrower niches but higher densities. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 28 29. North American tree species richness (Fig. 24.3) (see Fig 21.3 in 4th ed): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 29 30. North American animal richness relationships with tree species richness (a-d) and potential evapotranspiration (e-h) (Fig. 24.4, see Fig. 21.4 in 4th ed.): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 30 31. North American richness relationships (Fig. 24.5, 3rd ed.): Ants & rodents BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Lizards Chydorid cladocera Slide - 31 32. Highest richness at intermediate productivity in some communities (Fig. 24.6, 3rd ed.): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 32 33. Effect of spatial heterogeneity: • Increased spatial heterogeneity also generates more species diversity (Fig. 24.8, 3rd ed.). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 33 34. Species diversity and habitat structural diversity for (a) fish in Wisconsin lakes and (b) birds (Fig. 24.9, 3rd ed.): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 34 35. Lizard richness against desert plant measures (Fig. 24.10, 3rd ed.): BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 35 36. Effect of climatic variation: • Unpredictable climatic variation is a form of disturbance and species richness may be highest at intermediate levels. • But some work shows that species richness increases as climatic variation decreases (Fig. 21.9). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 36 37. Gradients of richness: • Latitude: • Increase in species diversity with a decrease in latitude towards the tropics (Fig. 24.13, 3rd ed.). • Perhaps because of more intense predation (topdown) and also reduced competition, as well as increased productivity (bottom-up) but with most productivity locked up in biomass releasable by high decomposition rates after death. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 37 38. Gradients of richness: • Altitude: • Decrease in species richness with increased altitude (Fig. 24.17, 3rd ed.). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 38 39. Gradients of richness: • Depth: • Generally species richness in lakes decreases with water depth. • But for benthic invertebrates richness is highest on the continental shelf at about 2000m (Fig. 24.19, 3rd ed.). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 39 40. Gradients of richness: • Succession: • Species richness increases with time through successional series because of a shift in dominance (Fig. 24.20, 3rd ed.) from smaller numbers of dominant species to more species of equivalent dominance. BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 40 41. Gradients of richness: • Succession: • Decline in richness at successional maturity (Fig. 24.21, 3rd ed.). BIOS 6150: Ecology - Dr. S. Malcolm. Week 13: Community structure Slide - 41