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John H. Lawton; Moshe Shachak Ecology, Vol Positive and Negative Effects of Organisms as Physical Ecosystem Engineers Clive G. Jones; John H. Lawton; Moshe Shachak Ecology, Vol. 78, No. 7. (Oct., 1997), pp. 1946-1957. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28199710%2978%3A7%3C1946%3APANEOO%3E2.0.CO%3B2-Q Ecology is currently published by Ecological Society of America. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/esa.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Tue Oct 30 13:55:18 2007 SPECIAL FEATURE Ecology Vol. 78, No. 7 Ecology, 78(7), 1997, pp. 1946-1957 0 1997 by the Ecological Society of America POSITIVE AND NEGATIVE EFFECTS OF ORGANISMS AS PHYSICAL ECOSYSTEM ENGINEERS CLIVEG. JONES,'JOHN H. LAW TON,',^ AND MOSHESHACHAK'~~ 'Institute of Ecosystem Studies, Box AB, Millbrook, New York I2545 USA 2Centrefor Population Biology, Imperial College, Silwood Park, SL5 7PY, UK 3Mitrani Center for Desert Ecology, Blaustein Institute for Desert Research, Ben-Gurion University of the Negev, Sede Boqer 84900, Israel Abstract. Physical ecosystem engineers are organisms that directly or indirectly control the availability of resources to other organisms by causing physical state changes in biotic or abiotic materials. Physical ecosystem engineering by organisms is the physical modi- fication, maintenance, or creation of habitats. Ecological effects of engineers on many other species occur in virtually all ecosystems because the physical state changes directly create nonfood resources such as living space, directly control abiotic resources, and indirectly modulate abiotic forces that, in turn, affect resource use by other organisms. Trophic in- teractions and resource competition do not constitute engineering. Engineering can have significant or trivial effects on other species, may involve the physical structure of an organism (like a tree) or structures made by an organism (like a beaver dam), and can, but does not invariably, have feedback effects on the engineer. We argue that engineering has both negative and positive effects on species richness and abundances at small scales, but the net effects are probably positive at larger scales encompassing engineered and nonen- gineered environments in ecological and evolutionary space and time. Models of the pop- ulation dynamics of engineers suggest that the engineerlhabitat equilibrium is often, but not always, locally stable and may show long-term cycles, with potential ramifications for community and ecosystem stability. As yet, data adequate to parameterize such a model do not exist for any engineer species. Because engineers control flows of energy and materials but do not have to participate in these flows, energy, mass, and stoichiometry do not appear to be useful in predicting which engineers will have big effects. Empirical observations suggest some potential generalizations about which species will be important engineers in which ecosystems. We point out some of the obvious, and not so obvious, ways in which engineering and trophic relations interact, and we call for greater research on physical ecosystem engineers, their impacts, and their interface with trophic relations. Key words: cascades, coupled trophic and engineering; community stability; ecosystem engineers; ecosystem function: feedbacks; habitat formation and destruction; population dynamics; positive and negative effects; species diversi ty. branches cast shade, reduce the impact of rain and wind, moderate temperature extremes, and increase hu- What does a tree do in a forest? Of course the living midity for organisms in the understory and the soil and dead tissues are eaten by many animals and mi- (Holling 1992, Callaway and Walker 1997). Root croorganisms, and the tree competes with other plants growth aerates the soil, alters its texture, and affects for light, water, and nutrients. But a tree does much the infiltration rate of water (Bouma and Anderson more than provide food and directly compete for re- 1973, Tisdall and Oades 1982, Smiles 1988, Juma sources. The branch, bark, root, and living and dead 1993). Dead leaves fall to the forest floor altering rain- drop impact, drainage, and heat and gas exchange in leaf surfaces make shelter, resting locations, and living the soil habitat, and make barriers or protection for space. Small ponds full of organisms form where seeds, seedlings, animals, and microbes (Facelli and throughfall gets channelled into crotches (Kitching Pickett 1991, Callaway and Walker 1997). The trunk, 1971, 1983), and the soil cavities that form as roots branches, and leaves can fall into forest streams cre- grow provide animals with places to live and cache ating debris dams and ponds for species to live in (Lik- food (Foster 1988, Vander Wall 1990). The leaves and ens and Bilby 1982, Hedin et al. 1988). The roots can Manuscript received 1 December 1995; accepted 3 January bind around rocks, stabilizing the substrate and ame- 1997. For reprints of this Special Feature, see footnote 1, liorating hurricane impacts on other species (Basnet et page 1945. al. 1992). If the tree falls, the downed trunk, branches, October 1997 POSITIVE INTERACTIONS IN COMMUNITIES 1947 and resulting tip-up soil pit and mound create habitats and Meadows 1991, Wilson and Agnew 1992, Jones et for numerous organisms (Collins and Pickett 1982, Pe- al. 1994, Butler 1995, Jones and Lawton 1995, Flecker terson and Pickett 1990, Peterson et al. 1990). 1996), including many effects caused by Homo sapiens, a physical ecosystem engineer par excellence (see Jones et al. 1994). We have argued that physical eco- It is probable that more species are affected by these system engineering by organisms plays a major role in things that a tree does than directly use the tree for determining the structure and functioning of most eco- food or compete with it for light, water, and nutrient systems, and we have yet to find an ecosystem in which resources. And yet, these diverse ecological effects are physical ecosystem engineering by organisms does not not trophic. Nor is the tree in the forest unique. Most play some role, even in such hostile environments as plants have similar effects, and many animals and mi- Arctic ice (Buynitskiy 1968, Arrigo et al. 1991). Here, croorganisms cause ecologically significant physical we refine the concept of engineering, distinguishing it changes in their environments (Jones et al. 1994). A from trophic relations and competitive interactions. We woodpecker or rot fungus may make holes in the tree explore the probable net effects of physical ecosystem that are then used by other species (Kitching 1971, engineers on species diversity and abundances, and Bradshaw and Holzapfel 1985, 1992, Daily et al. 1993), upon population, community, and ecosystem stability. and a beaver may come along and cut down the tree, We also ask whether there are ways to predict which making a dam and pond in which hundreds of species species will be important physical engineers, and which live (Pollock et al. 1995). ecosystems will be the most affected by them. Lastly, If these effects are not trophic, what are they? De- we explore ways in which physical engineering and spite their diversity, all of them involve changes from trophic interactions can be integrated. one physical state or condition to another in the tree (e.g., a tree without crotch ponds to a tree with them) or its local environment (e.g., a stream without a debris We first need to formally define what we mean by dam to a dam and pond). These physical state changes engineering (the definition that follows is modified are caused by the tree itself or by an organism, like slightly from Jones et al. [1994], further clarifying the the woodpecker, fungus, or beaver, that changes the concept and should be read in consultation with Fig. physical state of the tree (e.g., a tree without holes to 1). "Physical ecosystem engineers are organisms that a tree with holes) or the local environment (e.g., a directly or indirectly control the availability of re- stream without a beaver dam to a tree dam and a pond). sources to other organisms by causing physical state In all of these cases the physical state changes can have changes in biotic or abiotic materials. Physical eco- both positive and negative ecological consequences for system engineering by organisms is the physical mod- other species that live in the old or the new environment ification, maintenance, or creation of habitats.
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