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Fundamentals of Restoration Ecology Ecological Succession: The key To Restoration • Succession is the process of change in ecosystem structure and composition over time. • Succession drives ecosystem response to restoration efforts. • Restoration, then, is the attempt to initiate and/or direct successional processes. Linear model of Primary Succession H.C. Cowles’ successional stages at the southern end of Lake Michigan Figure from Keeton, W.T. (1980) Models of Succession • Relay floristics or “facilitation model” – Facilitative replacement of species – Sequential, directional, somewhat predictable • Initial floristics or “tolerance model” – All or most species present at early successional stage – Change in relative abundance over time • Intermediary models – Some aspects of facilitation and fluctuations in relative abundance due to competition and environmental modification as a community develops Primary Succession Sere 1 (colonization): Initially by specialist stress-tolerant species able to cope with rudimentary soils and extreme moisture/nutrient availability, e.g. mosses, lichens. Sere 2 (development): Soils improve, organic content increases, productivity increases. Environment less stressful, but still vulnerable to disturbance. Stress-tolerant species replaced by more competitive and productive sere 2 species, which tolerate some disturbance, e.g. grasses and weeds. Sere 3 (mature): soil now developed, soil conditions more stable, nutrient and water system is not stressful. Competitive species predominate(many still short life cycle), e.g. grasses, bushes and shrubs. Productive system with more complex trophic structure and cycling. Sere 4 (climax): relatively stable vegetation community (???), productivity is high and ecosystem structure complex. Often dominated by competitors, which are long-lived species (e.g. trees). Little evidence of initial conditions or stress/disturbance tolerant species (but occur locally: river banks, gaps left by tree-fall). Stable climax may not exist. Grimes (1979) life history strategies for plants • Competitive – reduce allocation towards vegetative growth and reproduction. This is not necessary in an environment where there is little interspecific competition. Instead they invest in features that ensure the endurance of mature individuals, i.e. special adaptations in growth form and structure (below ground biomass). • Stress-tolerant – maximize the capture of resources in productive but relatively undisturbed habitats. Bracken fern (Pteridium aquilinium) is a classic competitor. It has large reserves of energy stored in underground organs that can be mobilized rapidly in the growing season to produce large vegetative canopies. • Ruderal – are usually herbs having a short life-span and high seed production. They are found in highly disturbed, but potentially productive environments (e.g trampled but arable ground). Initially, competition is reduced in disturbed environments. Ruderals invest in regenerative phases (e.g. seeds, vegetative propagules or runners, protective growth forms/structures). Many such species are considered to be weeds. Rapid growth and development means that they mature and set seed quickly, which ensures population persistence through subsequent disturbances. Important for the selection of plants in restoration efforts Secondary Succession Figure adapted from Franklin and Spies (1991). Succession difficult to predict • Multiple determinants of early succession • Multiple pathways of succession are possible Multiple Pathways of Succession: f (Timing, Type, and Intensity of Disturbances + Masting and Seed Availability) From Hemstrom and Logan (1986), in Spies (1997) Altered Successional Pathways Resulting from a Complex History of Land-use Figure from Foster (1992) Why is an understanding of natural disturbance regimes so important for restoration? • Potential for disturbances to impair restoration success • Potential for disturbances to facilitate restoration success • Desired future condition must be dynamic • Mimic the role of biological legacies in post-disturbance ecosystem recovery Natural Disturbance Regimes • • • • • • Type Intensity Frequency Spatial extent and pattern Specificity Synergisms Types of Natural Disturbances Ice Storms Insect and Pathogens Outbreaks Floods Fine-scale Windthrow Large-scale Windthrow: Hurricanes, Tornadoes, etc. Timber volume blown down by the 1938 hurricane per each township Legend Estimated volume of timber blown down Over 10,000 board feet 1,000 to 10,000 board feet Sources: Figure from Boose et al (1994); Data compiled by the Northeastern Timber Salvage Administration (1943) < 1,000 board feet Not affected or no report Hurricanes in New England 1635 (8/25) Great Colonial Hurricane* 1879 (8/18-19) Cape Cod Hurricane of '79 1893 (8/24) 1638 (8/3) 1675 (9/7) Second Great Colonial Hurricane 1893 (8/29) passed well-inland 1683 (8/23) Hurricane and Flood of 1683 1896 (10/12-13) offshore hurricane 1713 (8/30) 1916 (7/21) excessive rain+all 1727 (9/27) 1924 (8/26) Off-shore Hurricane of '24 1743 (11/2) Ben Franklin's Eclipse Hurricane 1933 (9/17-18) 13.27 inches rain at Provincetown 1749 (10/19) 1936 (9/18-19) 7.79 inches rain at Provincetown 1761 (10/23-24) Winthrop's Hurricane 1938 (9/21) 1770 (10/20) 1944 (9/14-15) Great Atlantic Hurricane* Stile's Hurricane Great New England Hurricane* 1778 (8/12-13) The French Storm 1950 (9/11-12) Hurricane Dog 1788 (8/19) Western New England Hurricane 1954 (8/31) Carol* 1815 (9/23) The Great September Gale* 1954 (9/11) Edna* 1821 (9/3) Redfield's Hurricane (arrived at low tide) 1954 (9/11) Hazel 1841 (10/3) The October Gale 1955 (8/17-19) Diane -- extreme floods 1856 (8/21) Charter Oak Storm 1960 (9/12) Donna 1869 (9/8) September Gale of '69 1985 (9/27) Gloria 1991 (8/19) Bob 1878 (10/23-24) Predicted Topographic Susceptibility to Windthrow Figure from Boose et al. (1994) Fire Forest fires happens in New England too! Intensity of Natural Disturbances Low Intensity Proportion of events Low Intensity High Intensity High Intensity Proportion of events Low Intensity High Intensity Frequency • High frequency regimes typically have low average intensity • Low frequency regimes typically have high average intensity Think of purposeful disturbance as a tool for restoration High-intensity disturbance = large opening or big area + high mortality sets back succession Low-intensity disturbance = small gap or area + low mortality accelerates succession Spatial extent and specificity of disturbances influence ecosystem pattern Mosaic of Patches: Fine-Scale, Dynamic Mosaic of Patches: High Contrast, Dynamic Mosaic of Patches: High Contrast, Stable Mosaic of Patches: Coarse-scale, Dynamic Mosaic of Patches: Anthropogenic disturbances Synergism What are biological legacies? • Whole organisms • Reproductive structures: seeds, spores, stumps, rhizomes • Organically derived structures: snags, logs, SOM, soil aggregates • Patterns: organic and inorganic – Microbial distribution – Soil chemistry and inhibition – Community patterns: e.g. gaps and antigaps Implications of biological legacies for restoration Restoration at Mount St. Helens: Passive and Active Approaches Aided by Biological Legacies Biological Legacies of the 1938 Hurricane in MA Biological legacies: organic matter carryover •“Lifeboats” organisms above and below-ground Organic matter helps ameliorate postdisturbance stress: •soil moisture retention •microclimate: shade, windspeed, etc. •soil stabilization •nutrient cycling Linkages between terrestrial and aquatic ecosystems: • Bank stabilization •organic matter inputs Biological legacies can persist on a site for a long-time; ecological functions change as the structure ages (e.g. decays) How do ecosystems respond to disturbance? Ecosystem Structure and Function Time Ecosystem Structure and Function Disturbance Disturbance Time Desired Future Condition • Given that ecosystems are dynamics, what should the desired future condition be? Early Successional? Mid Successional? Late-Successional? Reference Condition • What should we use as reference condition? – Historic condition – which one? – Reference site: e.g. an extant, ecologically similar but un-degraded site. But which site? What part of it? – Predicted future condition in light of global climate change. Should we take this into consideration? • The key is to understand the range of variability. Restore to something within this range. Historical Range of Variability Proportion of Landscape in Old-growth 1 HRV 0.5 0 1400 1500 1600 1700 1800 1900 Year Figure from Aplet and Keeton (1999) Proportion of Landscape in Old-growth Scale: Small Watershed 1 Hurricane HRV 0.5 0 Scale: Drainage Basin Proportion of Landscape in Old-Growth 0 100 200 300 400 500 Years 1 Hurricanes HRV 0.5 0 0 100 200 300 400 500 Scale: Region Proportion of Landscape in Old-Growth Years 1 HRV 0.5 0 0 100 200 300 Years 400 500 Source: Aplet and Keeton (1999) Restoration of Native Vegetation: Exotic Organism Control 1. Understand biology (i.e. life history) of the exotic organism 2. Identify critical life history stage What life history traits make organisms successful invasives? 3. Determine possible control practices/techniques What intensity of treatment is acceptable? 4. Map your site compartmentalize based on exotic species occurrence, density, threat, etc. 5. Develop removal program and schedule Invasive Species Management European Buckthorn Japanese Knotweed Japanese Honeysuckle Tartatian Honeysuckle Using fire to control exotics… Kudzu Case Study • Kudzu (Pueraria lobata) is a perennial vine in the legume family • Imported from Japan in 1876 to landscape a garden at the Japanese Pavilion at the Philadelphia Centennial Exposition. • In the early 1900's, this vine was discovered to be excellent forage for cows, pigs, and goats in the South in acidic soils and during droughty seasons. It was also promoted as cover for erosion control in gullies. • The distribution of kudzu in the United States today extends from Connecticut to Missouri and Oklahoma, south to Texas and Florida. Before 1970, kudzu was planted along Missouri highways to control erosion and some farmers experimented with kudzu for livestock fodder. Kudzu infestation 1. Mechanical and hand removal 3. Prescribed burning then herbicide application 4. Native grasses planted Completed restoration