Foundation Species: Consequences for the Structure and Dynamics of Forested Ecosystems Author(S): Aaron M
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Loss of Foundation Species: Consequences for the Structure and Dynamics of Forested Ecosystems Author(s): Aaron M. Ellison, Michael S. Bank, Barton D. Clinton, Elizabeth A. Colburn, Katherine Elliott, Chelcy R. Ford, David R. Foster, Brian D. Kloeppel, Jennifer D. Knoepp, Gary M. Lovett, Jacqueline Mohan, David A. Orwig, Nicholas L. Rodenhouse, William V. Sobczak, Kristina A. Stinson, Jeffrey K. Stone, Christopher M. Swan, Jill Thompson, Betsy Von Holle and Jackson R. Webster Source: Frontiers in Ecology and the Environment, Vol. 3, No. 9 (Nov., 2005), pp. 479-486 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/3868635 . Accessed: 19/09/2014 10:31 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Frontiers in Ecology and the Environment. http://www.jstor.org This content downloaded from 158.135.136.72 on Fri, 19 Sep 2014 10:31:20 AM All use subject to JSTOR Terms and Conditions REVIEWS REVIEWS REVIEWS Loss of foundation species: consequences I for the structure and dynamics of forested ecosystems Aaron M Ellisonl*, Michael S Bank1, Barton D Clinton2, Elizabeth A Colburnm,Katherine Elliott2, Chelcy R Ford2, David R Foster1, Brian D Kloeppel3, Jennifer D Knoepp2, Gary M Lovett4, Jacqueline Mohan1, David A Orwig1, Nicholas L Rodenhouse5, William V Sobczak6, Kristina A Stinson', Jeffrey K Stone7, Christopher M Swan8, Jill Thompson9, Betsy Von Holle1, and Jackson R Webster10 In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species-specific traits control ecosystem dynamics. Such foundation tree species are declining throughoutthe world due to introductionsand outbreaksof pests and pathogens,selective removal of individual taxa, and over-harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environmenton which a varietyof other speciesdepend; how this disruptsfun- damental ecosystem processes,including rates of decomposition, nutrient fluxes, carbon sequestration,and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynam- ics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small pertur- bations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, antic- ipate the cascade of immediate, short- and long-term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management. I Front Ecol Environ 2005; 3(9): 479-486 WV e are living in an era of unprecedented and rapid life on Earth (Eldridge 1998). The loss of a common or ecological change (Reid et al. 2005). Through abundant foundation species (sensu Dayton 1972; see habitat conversion, over-consumption of resources, and Panel 1), which by virtue of its structural or functional worldwide introductions of pests and pathogens, humans attributes creates and defines an entire ecological com- are causing species extinctions at a record rate: the sixth munity or ecosystem, can have dramatic effects on our extinction crisis in the billion-year history of eukaryotic perception of the landscape and broad consequences for associated biota, ecosystem function, and stability. In a nutshell: Foundation species differ from keystone predators (Paine * In manyecosystems, a singlefoundation species controls pop- 1966) in that the former usually occupy low trophic levels ulationand communitydynamics and modulatesecosystem whereas the latter are usually top predators.They are also processes distinct from core (Hanski 1982) in that foundation * species The lossof foundationspecies acutely and chronically impacts are not abundant and com- fluxesof andnutrients, foodwebs, and bio- species only locally regionally energy hydrology, mon but also create stable conditions diversity locally required by * Humanactivities, including logging and the introductionof many other species. They also serve to stabilizefundamental exoticpests and pathogens, often functionally remove founda- ecosystemprocesses such as productivityand water balance. tion treespecies from forests Trees are most to be foundation in * likely species Foundationspecies that are currentlybeing lost fromNorth forested as their architecture and functional Americanforests include eastern hemlock, Port-Orford cedar, ecosystems, andoaks and physiological characteristics define forest structure and alter microclimates, while their biomass and chemi- 'HarvardUniversity, Harvard Forest, 324 NorthMain Street, Petersham, MA 01366 *([email protected]);2USDA Forest Service SouthernResearch Station, Coweeta Hydrologic Lab, Otto, NC, 28763; 3Universityof Georgia,Institute of Ecology,Athens, GA 30602; 4Instituteof EcosystemStudies, Millbrook, NY 12545;5Wellesley College, Department of BiologicalSciences, Wellesley, MA 02481;6College of theHoly Cross, Department of Biology,Worcester, MA 01610;70regon State University, Department of Botanyand Plant Pathology, Corvallis, OR 97331;8University of Maryland,Baltimore County, Dept of Geography& EnvironmentalSystems, Baltimore, MD 21250;9University of PuertoRico, Institute for Tropical Ecosystem Studies, San Juan, PR 00931; 'VirginiaPolytechnic Institute and State University, Department of Biology,Blacksburg, VA 24061 www.frontiersinecology.org © The Ecological Society of America www.frontiersinecology.org This content downloaded from 158.135.136.72 on Fri, 19 Sep 2014 10:31:20 AM All use subject to JSTOR Terms and Conditions I Loss of foundation species AM Ellison et al. Figure 1. (a) An old-growtheastern hemlock (Tsuga canadensis) stand, (b and inset) a stand decliningfollowing 10 years of infestationby the introducedhemlock woolly adelgid (Adelges tsugae), and (c) denseregeneration of blackbirch (Betula lenta) saplingson a siteformerly dominated by easternhemlock in southernConnecticut. Nearly all hemlocktrees in this 150-hectareforest in southernConnecticut were killed in themid-1 990s by hemlockwoolly adelgid. cal makeup contribute substantially to ecosystem canadensis)resulting from an introduced insect and pre- processes. Foundation tree species are declining through- emptive salvage logging; the local extirpation of white- out the world due to a number of factors, including intro- bark pine (Pinus albicaulis)caused by interactions among ductions and outbreaks of nonindigenous pests and a nonnative pathogen, a native insect, and human alter- pathogens, irruptions of native pests, over-harvesting and ation of fire regimes; and the functional removal of high-intensity forestry,and deliberate removal of individ- American chestnut (Castanea dentata) by an introduced ual species from forests. We use three examples from pathogen. Our examples focus on trees in systems we North America to illustrate consequences for both terres- know best, but they are broadly representative of a wide trial and aquatic habitats of the loss of foundation tree range of foundation species and illustrative of their role in species: the ongoing decline of eastern hemlock (Tsuga forests throughout the world (Panel 2). Panel 1. The many definitions of foundation species * The rise and fall of eastern hemlock Followingnomenclatural priority, we adopt Dayton's(1972) ter- minology and general definitionof a foundationspecies: a single Majestic hemlock groves (Figure 1) evoke reverence, speciesthat definesmuch of the structureof a communityby creating affection, and poetry (Frost 1923). Eastern hemlock stableconditions other and and sta- locally for species, by modulating (Tsuga canadensis),one of the most long-lived, shade-tol- bilizingfundamental ecosystem processes. erant trees in North America, dominates about 1 x 106ha Subsequentauthors, working in differenthabitats and appar- to southern ently unaware of historical antecedents, have suggested terms of forest from the southern Appalachians with some or all of the attributesof foundationspecies, including: Canada and west to the central Lake states (McWilliams Core species (Hanski 1982) are locallyabundant and region- and Schmidt 2000). In the north, hemlock typically ally common;associated satellite species are sparse and rare. occurs in stands with species-poor understo- An associated model core-satellite nearly pure metapopulation (the ries. In the south, hemlock in mixed stands in nar- hypothesis) explains relationships between a species' local grows abundanceand its regionaldistribution. row riparian strips and moist coves, often with dense Dominant species (Grime 1984) competitively exclude understories of rhododendron (Rhododendronmaximum). subordinate species by garneringa disproportionate share of In hemlock-dominated stands, the combination of deep