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Towards a Food-web Perspective on Biodiversity and Ecosystem Functioning

One of the most common questions asked by researchers across a variety of scientific disciplines is ‘How does the number of nodes connected together into a network influence the efficiency and reliability of that network?’. While social scientists and epidemiologists might think of ‘nodes’ and ‘connections’ as people interacting within a social network, computer scientists, neurologists, and civil engineers would instead think of servers connected together in a world-wide web, synapses connecting neurons in the brain, or hubs connecting to other hubs in a transportation network or telecommunications grid (Albert and Barabasi 2002, Newman 2003). Regardless of the particular study system, all of these individuals ask similar questions about how the number of nodes and connections among nodes influence the efficiency and reliability by which information, disease, energy, or matter is transmitted throughout that network. Within the field of ecology, one of the oldest and most fundamental questions asked by researchers is ‘How does the number of species interacting within a food web influence the efficiency and reliability by which energy and matter are transmitted through that web?’. Research on this topic can be broadly divided into two foci. Historically, much attention in ecology has focused on identifying those taxa that are the most influential nodes in a food web. For many years, it has been thought that some subset of species might represent ‘hubs’ of interactions and/or exhibit such strong interactions that they exert a disproportionate influence over food web dynamics. This idea has fueled much debate over the prevalence of omnivory in food webs (Polis and Strong 1996, Thompson et al. 2007, Yodzis 1984) and whether the increased number of feeding links that result from omnivory increases or decreases the stability of energy flow through a food web (McCann et al. 1998, MacArthur 1955). Identifying species that represent influential nodes has also been one of the primary goals in the search for ‘ecosystem engineers’ (Jones et al. 1994), ‘keystone species’ (Paine 1966, Power et al. 1996) or other types of ‘strong interactors’ (Wootton and Emmerson 2005) that might have cascading effects on the diversity and biomass of species at a variety of different trophic levels (Paine 1966, Carpenter et al. 1987, Elser et al. 1988). In the 1990s, ecologists began to pursue a slightly different perspective on food webs. This perspective focused not on the cascading impacts of individual species, but rather on how the number of species that comprise any single trophic level might control fluxes of energy and matter. Research in this area was generally referred to as Biodiversity effects on Ecosystem Functioning (BEF for short), and was often justified on grounds that (1) loss of biological diversity ranks among the most pronounced changes to the global environment (Sala et al. 2000, Pimm et al. 1995), and (2) reductions in diversity, and corresponding changes in species composition, may alter fluxes of energy and matter that underlie important services that ecosystems provide to