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Patterns of Succession Succession is: • Non-seasonal, directional change in plant species living in a particular place through time. • Involves: – Colonization – Establishment – Ultimate local extinction • Initiated by a disturbance that opens up space and resources Types of Succession • Primary: Occurs when new bare rock is generated by geologic activity (no organic material) – Mt. St. Helens, Krakatoa, glacial retreat • Secondary: vegetation invades areas that have been previously occupied, but has been removed due to disturbance (fire, agriculture, etc) Primary Succession after glacial retreat Example of primary succession: lichensmoss small forbsshrubstrees QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Lichens and moss growing on rock (primary succession) Secondary Succession QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Secondary Succession in YNP after 1988 fires Classic Study: Billings (1938) • Studied succession from old field to oak forest (150 years) – – – – – Annual plants = pioneer species Biennial plants, grasses Perennial plants, shrubs After about 15 years: softwood trees After about 50 years hardwood trees, primarily oak “Old-field Succession” QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Successional Progression • Annuals: Pioneer Species – – – – Tolerate low nutrient soils, need high light Good dispersers Grow rapidly, short life span (1 year) Produce many offspring • Biennial plants, grasses – Share attributes with pioneer species, but longer lived, grow more slowly • Perennials and shrubs – Slower growing, longer lived – Can reproduce asexually as well as sexually – Better competitors • Softwood trees (Pines) – Slow growing – Eventually produce canopy – Shade out light-loving shrubs and perennials in understory – Also shades out new baby pines • Hardwood trees (Oaks and hickory) – Longer lived – Better competitors – Eventually dominated forest as softwood trees died. Oak forest in this case = climax community Climax Communities • Final successional stage: self-replacing, persistant over very long periods of time • Idea of climax community controversial these days because of recognition of role of disturbance – Shifting mosaic steady state model: • Majority of patches in habitat in some stage of recovering from disturbance • Landscape is in a steady-state because roughly constant portions of landscape are in each stage of succession • Thus there is a dynamic equilibrium across the landscape Disturbance Specialists: Often Weeds QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Mechanisms of Succession Connell and Slayter, 1977 1. Facilitation - Changes in abiotic conditions are caused by the plants currently occupying an area - These changes favor new invaders over the current occupants (current occupants pave the way for future occupants) Examples: Lichens make soil, nitrogenfixing plants enrich soil, nurse plants support others 2. Tolerance • Plants in different stages of succession don’t have much impact on each other • Position in the scheme of succession depends on their different life histories • Example: Old field Succession 3. Inhibition • Members of one stage of succession resist invasion of later stages • Succession proceeds when one stage dies Example: Allelopathy Succession and Chemical Cycling • Biomass, production, diversity and chemical cycling change during succession • Biomass and diversity peak in midsuccession, increasing at first to a maximum, then declining and varying over time. K-selection and r-selection • Pioneer species generally have particular set of characteristics - all together these are called r-selected traits • Climax species generally have different set - represent a different life history strategy Succession and Plant Diversity Succession and Diversity Just focusing on woody plants: Corresponding changes in animal species Succession in the Intertidal Diversity through Succession in Intertidal Succession in Streams Points to Consider: • Change through time is a natural characteristic of most ecosystems - when thinking about conservation need to incorporate understanding of natural disturbance regimes • Mosaic of Patches in different stages of succession maximizes habitat heterogeneity and thus diversity Ecological Restoration • Attempt to return impacted land to natural state • What exactly should be returned? • Increasingly are trying to replace the original disturbance regime so that succession can occur naturally – Need to restore disturbance and succession processes both temporally and spatially – What area of land is big enough to encompass these processes? Stream Restoration • What ecosystem functions/services need to be restored? – – – – – – Habitat Conduit Barrier Filter Source Sink • Processes need to be restored, not just individual components – Hydrologic functions: • How much flow, how deep, timing – Geomorphological functions • Stream profile, sedimentation patterns, erosion – Physical and Chemical Properties of Water • Toxins? • Nutrients • PH – Biological Processes (eg large woody debris, habitat structure, heterogeneity in stream) – Role of Disturbance/Dynamic Equilibrium