ECOLOGICAL SUCCESSION CONCEPTS

Dictionary definition of succession: Sequence; a following of one thing after another in time

Ecological Succession:
The gradual and orderly process of ecosystem change over time brought about by changes in biotic and abiotic components What changes as a result of ecological succession?

Primary Succession Occurs on primary sites: bare areas devoid of biotic communities Examples: bare sand dunes lava flows ash deposits mud flows glacial till mine spoils

Secondary Succession Occurs on sites where the existing biological community has been disturbed Examples: logged areas grazed areas burned areas windstorm

Historical development of the concept of succession: Henry Cowles (1869-1939); Univ. Chicago; 1899: Ecological relations of the vegetation on the sand dunes of Lake Michigan Frederick E. Clements (1874-1945); Univ. Nebraska; 1916: Plant succession: an analysis of the development of vegetation. Carnegie Inst. Wash. Pub. No. 242
What Henry Cowles did:

What Frederick Clements did: defined succession as a sequence of plant communities marked by the change from lower to higher life forms 5 Key points: 1. Replacement of species by other species over time

2. Early species change the environment, making it more suitable for species that follow

3. Only one uni-directional succession trajectory

4. Succession is deterministic (predictable)

5. Succession leads to a single terminal steady-state called climax

Example characteristics of early-seral environments:

Low soil moisture Low soil organic matter Poor soil structure Low vegetation cover Temperature extremes Therefore unstable and unpredictable environment: weakly functioning or inoperative primary processes (water cycle; nutrient cycle) succession driven mostly by abiotic factors

Characteristics of early-seral plants: Tolerant of unstable/unpredictable environments Fast growth Short-lived Many seeds Widely-dispersed seeds Resources put into reproduction Small size

Example characteristics of late-seral environments:

Well-developed soil Maximum soil organic matter Soil water available Increased vegetation cover Modulated temperatures Therefore stable and predictable environment: functioning primary processes driven by biotic factors

Characteristics of late-seral plants: Intolerant of environmental disturbance Slow growth Long-lived Large size Infrequent reproduction Resources put into permanent tissues

Mechanisms of Succession
6 stages of succession, according to Clements (Facilitation model):
1. Nudation: creation of a bare substrate

2. Migration:

arrival of propagules

3. Ecsesis:

germination, establishment and reproduction

4. Competition:

leads to species replacement

5. Reaction:

vegetation modifies the environment making it suitable for new colonists

6. Stabilization:

attainment of climax

3 alternative mechanisms producing the sequence of species change during plant succession:
Connell, J.H. and R.O. Slatyer. 1977. Mechanisms of succession in natural communities and their role in community stability and organization. American Naturalist (111):1119-1144. 1. Facilitation 2. Tolerance 3. Inhibition See Figure 4.1 of text (Whisenant, Repairing Damaged Wildland), p. 105

1. Facilitation Model:
Disturbed area exploited by pioneer species that modify the site making it suitable for other species. Once established, the later-successional species eliminate the pioneers through competition.

2. Tolerance Model:
Any species able to survive on the disturbed site will establish (early or late seral). These species may modify the site, but such modification does not affect recruitment of latesuccessional species. Then, species compete for resources and competition-intolerant species are eliminated.

3. Inhibition Model:
Like the tolerance model, both early- and late-seral species can establish after disturbance. But, some species make the site less suitable for other species. As long as inhibitory species persist they will suppress recruitment of other species.

Secondary Succession occurs on areas where an existing biotic community has been altered or destroyed by disturbance.

Models of secondary succession: 1. Single equilibrium endpoint 2. Multiple equilibrium states 3. Persistent non-equilibrium

1. Single Equilibrium Endpoint Based on F.E. Clements Application to Range Management: Range Succession Model
Sampson, Arthur W. 1919. Plant succession in relation to range management. USDA Bull.791. 76 p.

Key Assumptions of the Single Equilibrium Endpoint Model: 1. 2. Only one stable state (climax) Succession is linear and two-directional moving either toward climax (progression) or away from climax (retrogression) Once disturbance agent is removed, succession trajectory will follow same pathway as during primary succession Highly predictable and deterministic

3.

4.

2. Multiple Equilibrium States Also called: State and Transition Model Multiple Stable States Model References applicable to rangelands:
Briske, D.D., S.D. Fuhlendorf and F.E. Smeins. 2003. Vegetation dynamics on rangelands: A critique of the current paradigms. J. Applied Ecology 40:601-614. Friedel, M.H. 1991. Range condition assessment and the concept of thresholds A viewpoint. J. Range Manage. 44:422-426. Stringham, T.K., W.C. Krueger and P.L. Shaver. 2003. State and transition modeling: An ecological process approach. J. Range Manage. 56:106-113. Westoby, M., B. Walker, and I. Noymeir. 1989. Range management on the basis of a model which does not seek to establish equilibrium. J. Arid Environments 17:235-239.

Definitions: state = condition in which the structure and function of an ecosystem remains relatively stable. transition = event(s) which alters the structure and function of an ecosystem

Key Assumptions of the Multiple Equilibrium States Model: 1. 2. 3. Multiple stable states Succession may be non-linear If a disturbance is removed or changed the succession trajectory may not return to the same pre-disturbance state Not always predictable

4.

So, what determines the succession trajectory? 1. 2. 3. 4. 5. kind, intensity, frequency of disturbance what propagules are available what plants establish how plants compete for basic resources how plants alter the site to favor some species

3. Persistent Non-Equilibrium
1. Succession trajectories never reach a stable state;

2. Succession trajectories are unpredictable and nonlinear

3. Succession trajectories are driven by external stochastic (random, unpredictable) events and ecological legacies of the disturbed site

Head Notes on Ecological Succession: Be able to discuss the following:

1. People involved with the concept of ecological succession as applied to rangelands; what was the background and contribution of each person? Henry Cowles (1869-1939); Univ. Chicago. In 1899 published Ecological relations of the vegetation on the sand dunes of Lake Michigan, Botanical Gazette 27:95-117. Frederick E. Clements (1874-1945); Univ. Nebraska. In 1916 published Plant succession: an analysis of the development of vegetation, Carnegie Inst. Publ. 242. 512 p. Sampson, A.W. 1919. Plant succession in relation to range management. USDA Bull. 791. 76 p. 2. What is a model and why is the concept important? 3. What are 2 models that guide range management? 4. What is ecological succession? What things change as a result of succession? 5. What is primary succession? Define these terms: sere; seral stage; climax 6. What are the characteristics of early seral environments? 7. What are the characteristics of plants inhabiting early seral environments? 8. What are the characteristics of late seral environments? 9. What are the characteristics of plants inhabiting late seral environments? 10. describe 3 mechanisms of succession as defined by Connell & Slatyer. 11. What is secondary succession? 12. Describe the single equilibrium endpoint model of secondary succession 13. Describe the multiple equilibrium state model of secondary succession Terms/concepts to know: Succession: ecosystem change over time brought about changes in biotic and abiotic components. Sere: the whole collection of ecological communities, including their abiotic characteristics, along a successional trajectory from a bare site to climax. Seral stage: a specific ecological community within a sere that is recognizable by its own characteristic combination of plants

Climax: the idea of a terminal, steady-state endpoint of succession in which the established plants and animals are not replaced by other species. Progression: succession trajectory moving toward climax and becoming more complex in terms of species composition and ecological processes. Retrogression: movement of ecological communities away from complex ecological processes toward less stable and less complex communities. Deterministic: predictable, orderly Stochastic: random, unpredictable, chance Succession trajectory: the whole process of succession from the initial state to the final state; sere. 3 mechanisms of succession: facilitation, tolerance, inhibition OTHER REFERENCES:

Briske, D.D., S.D. Fuhlendorf and F.E. Smeins. 2003. Vegetation dynamics on rangelands: A critique of the current paradigms. J. Applied Ecology 40:601-614. Friedel, M.H. 1991. Range condition assessment and the concept of thresholds A viewpoint. J. Range Manage. 44:422-426. Stringham, T.K., W.C. Krueger and P.L. Shaver. 2003. State and transition modeling: An ecological process approach. J. Range Manage. 56:106-113. Westoby, M., B. Walker, and I. Noymeir. 1989. Range management on the basis of a model which does not seek to establish equilibrium. J. Arid Environments 17:235-239.