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Readings

Readings. Chapter 11 textbook Connell, J. H., and R. O. Slatyer. 1977. Mechanisms of succession in natural communities and their role in community stability and organization. The American Naturalist 111 :1119-1144. Outline. Intermediate disturbance hypothesis Dynamic equilibrium hypothesis

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Readings

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  1. Readings • Chapter 11 textbook • Connell, J. H., and R. O. Slatyer. 1977. Mechanisms of succession in natural communities and their role in community stability and organization. The American Naturalist 111:1119-1144.

  2. Outline • Intermediate disturbance hypothesis • Dynamic equilibrium hypothesis • Basic characteristics of succession • Mechanisms of succession • Disturbance and succession in range and forest ecosystems: scale • Large scale disturbance (fire, logging, landslides etc) • Gap dynamics and replacement models • Herbivory and holistic grazing management

  3. Intermediate disturbance hypothesis • Recall that: • Disturbance is creation of space • Plant species classified as r- or K selected • Intermediate disturbance hypothesis: at very high and very low rates of disturbance, the pool of species adapted to conditions is small. At intermediate levels, frequencies, or intensities, both r- and K species could survive, hence diversity highest.

  4. Intermediate disturbance hypothesis Species diversity Low disturbance High disturbance

  5. Dynamic equilibrium hypothesis • An extention of intermediate disturbance –Huston 1979, Am. Nat 113:81-101 • Species coexistence (therefore diversity) determined by RATE at which competitive equilibrium reached. • Therefore, disturbance increases richness in high productivity environments and reduces it in low productivity environments.

  6. Succession • Directional plant change after disturbance (2o) or on new substrate (1o) • May be AUTOGENIC (driven by plants) or ALLOGENIC (driven by environment)

  7. Characteristics of successional communities • Some common characteristics of communities as succession proceeds (table 11.3 text) • Often increase in diversity and biomass (though only to a point – recall I.D.H.) • Increase in structural complexity • Increase in resistance, decrease in resilience • Switch from r to K species • Decrease in productivity • Decrease in light available • More mesic environment (less stress) • Nutrients tied up in biomass; tight, slow cycling

  8. Mechanisms of succession • Relay Floristics (Clements): succession requires arrival at and site preparation by previous species. 6 steps: nudation, migration, ecesis, competition, reaction, and stabilization. • Initial Floristic Composition (Egler): plants involved in succession are already at site; differences in rate of establishment result in ‘seres’.

  9. Other explanations • Connell and Slatyer 1977: • Facilitation (RF; site preparation) • Tolerance (IFC; plants don’t interact much) • Inhibition (succession slowed by current vegetation) • Grime 1977: C-S-R life history • Shift in composition from R to C to S through time • Higher productivity, larger change in composition • Tilman 1985: Resource ratio hypothesis • Shift in biomass and resource availability allows change in competitive interactions throughout succession (different species “win” based on ability to deplete resources)

  10. Forest Succession • Three mechanisms: • Catastrophic disturbance (fire, landslide, clearcut, insect outbreak) • Gap dynamics (small-scale succession in treefall gaps) • Continuous recruitment without disturbance What sorts of species or species interactions might drive each of these mechanisms?

  11. Successional pathways may differ based on landscape properties, environment, etc (see handout) • Several ways of looking at secondary succession in forest stand • Stand development (initiation, stem exclusion, understory reinitiation, old growth) • Population model (establishment, thinning, transition, steady-state)

  12. Catastrophic disturbance • “Traditional” secondary succession • Previously discussed mechanisms of succession apply, including facilitation • Secondary succession models of stands based on this disturbance mechanism • Understory intolerant species dominate early • Transition to more tolerant species • Handout – fire in PIPO forest in Montana.

  13. Gap dynamics • “Regeneration process whereby small-scale, localized disturbance gaps form in the canopy, as one or a few trees die, and release resources of light, nutrients, water, etc. for colonization of the gap by seedlings or saplings of the same or replacement species”. • Smaller scale, less change in environmental conditions

  14. Gap dynamics • Can affect structure and composition of forest • Important in mesic forests with low rates of disturbance and long canopy turnover times (e.g. Appalachian mts – 0.9% disturbance and 125-year canopy replacement time) • Tend to find understory tolerant species • Seedling banks may come into play • Similar to tolerance or IFC models of succession • See handouts

  15. Gap dynamics • Note: can model canopy composition/structure dynamics using transition matrices. • Different forest species have different probabilities of colonizing treefall gaps of different sizes. • Given a starting composition, rate of gap formation for each species, and probability of replacement for each species, can model canopy composition.

  16. Disturbance in rangeland • Rangeland disturbances? • Role of disturbance? • Plant responses to disturbance?

  17. Disturbance in rangeland • Rangeland disturbances: • Fire • Soil disturbance (e.g. prairie dogs, hoofprints) • herbivory • Role of disturbance: • Reduce competition • Create regeneration microsites (Grubb 1979) • Release resources • Plant responses to disturbance: • Resprouting, seedling germination in microsites, increase or decrease.

  18. Plant functional groups • Functional group: usually defined based on • Role of species in ecosystem (eg N-fixer) • Physiognomy or growth form (e.g. bunchgrass, rhizomatous grass) • Response to disturbance (e.g. increaser, decreaser) • Characteristics of increasers and decreasers?

  19. Disturbance in rangeland • “Chronic disturbance” in grazed rangeland: dynamic equilibrium or non-equilibrium model. • Intermediate disturbance hypothesis: intermediate grazing pressure – increased productivity and diversity • Dynamic equilibrium hypothesis: more productive sites can handle higher grazing pressure

  20. Parallels with forest succession? • Gap dynamics at small scale? (e.g. level of a bite or hoofprint) • “catastrophic” disturbance – heavy grazing, fire, flood. Differences between responses and successional mechanisms under chronic grazing and intense rotational grazing (“holistic management” style)?

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