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What Do Human Economies, Large Islands and Forest Fragments Reveal About the Factors Limiting Ecosystem Evolution?

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What Do Human Economies, Large Islands and Forest Fragments Reveal About the Factors Limiting Ecosystem Evolution? doi:10.1111/j. 1420-9101,2008.01624.x REVIEW What do human economies, large islands and forest fragments reveal about the factors limiting ecosystem evolution? E. G. LEIGH, JR,* G. J. VERMEIJj & M. WIKELSKIJ * Smithsonian Tropical Research Institute, Balboa, Panama ''(Department of Geology, University of California at Davis, Davis, CA, USA %Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA Keywords: Abstract diversity; What factors limit ecosystem evolution? Like human economies, ecosystems fragmentation; are arenas where agents compete for locally limiting resources. Like econo- intensity of competition; mies, but unlike genes, ecosystems are not units of selection. In both intensity of herbivory; economies and ecosystems, productivity, diversity of occupations or species interdependence; and intensity of competition presuppose interdependence among many islands; different agents. In both, competitive dominants need abundant, varied productivity. resources, and many agents' products or services, to support the activity and responsiveness needed to maintain dominance. Comparing different-sized land masses suggests that productivity is lower on islands whose area is too small to maintain some of the interdependences that maintain diversity, productivity and competitiveness in mainland ecosystems. Islands lacking the rare, metabolically active dominants that make competition so intense in mainland ecosystems are more easily invaded by introduced exotics. Studies of islets in reservoirs identify mechanisms generating these phenomena. These phenomena suggest how continued fragmentation will affect future 'natural' ecosystems. Introduction and diversity of mainland ecosystems. Darwin (1859) had already predicted that the intensity of competition What conditions constrain ecosystem evolution? We was lower on islands, making them susceptible to begin our answer by comparing natural ecosystems with invasion by mainland exotics, because island species are human economies. Both evolve, in the sense of descent tested by fewer encounters with other species of com- with cultural or genetic modification of their members, petitors and predators. We also show that, absent human whether endogenous or driven by ideas or immigrants introductions, islands, especially smaller ones, lack rare, from outside. Unlike genes, ecosystems or economies metabolically active dominant predators like those rarely die or replicate: they are not units of selection. ensuring the rapid resource turnover that enhances Both are shaped by competition among members for productivity and intensity of competition in mainland locally limiting resources needed to survive and grow or ecosystems. Study of forest fragments isolated by reser- reproduce. For both economies and ecosystems, we show voirs confirms these conclusions and identifies some of how interdependence underpins productivity, diversity their underlying causes. As human activity is fragment- of species or occupations and intensity of competition. ing natural ecosystems at an ever-increasing rate (Lau- Next, we show that island ecosystems are less productive rance & Bierregaard, 1997), we will use our conclusions as well as less diverse than their mainland counterparts to predict how fragmentation will shape the 'natural' because the small area of an island offers less opportunity ecosystems of the future. for speciation in situ, specialization, and the relationships of interdependence that make possible the productivity Competition, economies and ecosystems Correspondence: Egbert Giles Leigh, Jr, Smithsonian Tropical Research In human economies, individuals and families compete Institute, MRC 0580-02, Unit 9100, Box 0948, DPO AA 34002-9998, USA. for the means to live. They often form coalitions such as Tel.: 507 212 8940; lax: 507 212 8937; e-mail: [email protected] tribes, businesses, city-states or republics, whose JOURNAL COMPILATION 2008 EUROPEAN SOCIETY FOR EVOLUTIONARY BIOLOGY NO CLAIM TO ORIGINAL US GOVERNMENT WORKS E. G. LEIGH ETAL. members cooperate to compete more effectively with invention, improvement and greatly reduced cost of others. Competitive success can be enhanced by finding various modes of long-distance communication amplified new ways to make a living, specializing to exploit a these developments. Larger economies support more narrower range of resources more effectively, specializing occupations (Bonner, 2004) and greater specialization of to particular techniques for procuring resources, or to occupations (Smith, 1776). They provide more opportu- particular locales for finding certain resources, or finding nity for the origin of competitive dominants, enhance the better ways to manage resources. Competition is most intensity of competition, and provide more opportunities likely to enhance an economy's range of resource use, for different specialists to pool complementary abilities to and its effectiveness in using them, if improvements compete better with third parties (Smith, 1776). These differentially benefit their inventors, and if market- phenomena jointly increased the gross product, diversity distorting monopolies do not restrict innovation (Smith, of occupations and levels of interdependence in the 1776; Netting, 1993). Thus competition favours innova- world economy during historic time (Smith, 1776; Leigh tion, specialization and diversification of occupations. For & Vermeij, 2002; Vermeij, 2004). example, the invention and progressive improvement of Although these principles are most obvious and best agriculture enabled higher production by fewer people of known for human economies, they apply equally to more easily stored food, freeing people for other occu- ecosystems comprised of organisms of other species. pations. Effective agriculture made possible greater divi- Other organisms also need resources to survive and sion of labour - a greater diversity of more specialized reproduce: they must also find ways to 'make a living.' As occupations, with greater efficiency in each specialty - in human economies, competition among individuals, thereby increasing interdependence among individuals coalitions of cooperating individuals, and species, for with different specialties (Smith, 1776). The usefulness or resources needed to survive and reproduce has driven an attractiveness of products of other specialists or different increase of productivity, diversity and intensity of com- lands made it possible to live by trade, an occupation petition in the world's ecosystems over evolutionary time based on interdependence. (Vermeij, 1987, 2004). In natural ecosystems, competi- Larger economies offer greater scope for interdepen- tion quickly breaks down most monopolies that 'distort dence and innovation, and more opportunity for the the market' by defending poorly used resources: poorly evolution of competitively dominant individuals, families used resources soon find better users. or businesses. Stratified societies where cities with Species diversify because organisms face trade-offs in monumental buildings depended on extensive surround- the ability to exploit different resources or habitats, ing agriculture first developed on continents: the north where adaptations enabling better use of one diminish China plain, the Indus valley, the 'Fertile Crescent' the capacity to use another. Speciation normally begins stretching from Mesopotamia to Palestine, Egypt, with genetic divergence of different parts of a population Mexico and Peru. Occupations connected with trade in response to a trade-off (Fisher, 1930: 125-129). favoured, and were favoured by, innovations enhancing Experiments and careful monitoring show that, where mobility, and multiplying the power of human work, there are trade-offs between the capacity to use different such as domesticating carriage animals and developing resources, 'divergence of character' (Darwin, 1859: 111), water and road transport. The invention of machines for now called character displacement (Brown & Wilson, manufacture and transport gave rise to vehicles such as 1956), reduces competition for shared resources, steamships, trains, trucks and airplanes. Inventing promoting coexistence among the species involved machines to produce these objects, and manufacturing (Schluter, 1994, 2005; Grant & Grant, 2007). As in them in the quantities needed, however, required the human economies, where the specialization involved in assembly of resources from a great many people and the division of labour allows great increases in efficiency coordination of work by specialists of many different (Smith, 1776), the specialization resulting from adaptive types. These achievements are far easier in large econ- divergence leads to more thorough exploitation of omies, with the resources to support competitively resources. Moreover, enhanced sensory capacities permit dominant business firms that respond to more different more refined choice of mates, allowing more opportunity stimuli and depend on a wider variety of resources and for divergent sexual selection, which promotes speciation markets. In turn, manufacture and transport favoured and diversification (Fisher, 1930; Losos, 1994: 486; further specialization and diversity of occupations and Jiggins etal, 2001; Grant & Grant, 2007; Puebla et al, faster turnover of a greater variety of resources. Manu- 2007). Diversity therefore increases over evolutionary facture and transport also increased productivity by time. providing a greater variety of useful or desirable prod- Innovation
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