Scenarios of changes in biodiversity for the year 2100 can now be developed based on scenarios of changes in atmospheric carbon dioxide, climate, vegetation, and land use and the known sensitivity of biodiversity to these changes. This study identified a ranking of the importance of drivers of change, a ranking of the biomes with respect to expected changes, and the major sources of uncertainties. For terrestrial ecosystems, land-use change probably will have the largest effect, followed by climate change, nitrogen deposition, biotic exchange, and elevated carbon dioxide concentration. For freshwater ecosystems, biotic exchange is much more important. Mediterranean climate and grassland ecosystems likely will experience the greatest proportional change in biodiversity because of the substantial influence of all drivers of biodiversity change. Northern temperate ecosystems are estimated to experience the least biodiversity change because major land-use change has already occurred. Plausible changes in biodiversity in other biomes depend on interactions among the causes of biodiversity change. These interactions represent one of the largest uncertainties in projections of future biodiversity change.

Localized ecological systems are known to shift abruptly and irreversibly from one state to another when they are forced across critical thresholds. Here we review evidence that the global ecosystem as a whole can react in the same way and is approaching a planetary-scale critical transition as a result of human influence. The plausibility of a planetary-scale 'tipping point' highlights the need to improve biological forecasting by detecting early warning signs of critical transitions on global as well as local scales, and by detecting feedbacks that promote such transitions. It is also necessary to address root causes of how humans are forcing biological changes.

Traditional approaches to the study of food webs emphasize the transfer of local primary productivity in the form of living plant organic matter across trophic levels. However, dead organic matter, or detritus, a common feature of most ecosystems plays a frequently overlooked role as a dynamic heterogeneous resource and habitat for many species. We develop an integrative framework for understanding the impact of detritus that emphasizes the ontogeny and heterogeneity of detritus and the various ways that explicit inclusion of detrital dynamics alters generalizations about the structure and functioning of food webs. Through its influences on food web composition and dynamics, detritus often increases system stability and persistence, having substantial effects on trophic structure and biodiversity. Inclusion of detrital heterogeneity in models of food web dynamics is an essential new direction for ecological research.

It has been suggested that differences in body size between consumer and resource species may have important implications for interaction strengths, population dynamics, and eventually food web structure, function, and evolution. Still, the general distribution of consumer-resource body-size ratios in real ecosystems, and whether they vary systematically among habitats or broad taxonomic groups, is poorly understood. Using a unique global database on consumer and resource body sizes, we show that the mean body-size ratios of aquatic herbivorous and detritivorous consumers are several orders of magnitude larger than those of carnivorous predators. Carnivorous predator-prey body-size ratios vary across different habitats and predator and prey types (invertebrates, ectotherm, and endotherm vertebrates). Predator-prey body-size ratios are on average significantly higher (1) in freshwater habitats than in marine or terrestrial habitats, (2) for vertebrate than for invertebrate predators, and (3) for invertebrate than for ectotherm vertebrate prey. If recent studies that relate body-size ratios to interaction strengths are general, our results suggest that mean consumer-resource interaction strengths may vary systematically across different habitat categories and consumer types.

We dedicate this paper to the memory of John P. Sutherland, a great scientist, a great human being.Abstract. The usefulness and generality of the keystone species concept has recently been questioned. We investigated variation in interaction strength between the original keystone predator, the seastar Pisaster ochraceus, and its primary prey, mussels (Mytilus ca/ifornianus and M. trossulus). The study was prompted by differences in community structure at two low zone sites along the central Oregon coast, Boiler Bay (BB) and Strawberry Hill (SH). Predators, especially seastars, were larger and more abundant at SH than at BB. Further, sessile animals were more abundant and macrophytes were less abundant at SH. Predators were more abundant at wave-exposed sites at both sites, and at SH, sessile invertebrates were more abundant at the wave-exposed location and sand cover was high at the wave-protected location. To test the hypothesis that variation in predation strength explained some of these differences, we examined the seastar-mussel interaction at locations with high and low wave exposure at both sites. Predation intensity was quantified by determining the survival of mussels in clumps (50 mussels per clump, shell length 4-7 em) transplanted to large plots (18-163 m 2 ) with or without seastars in the low intertidal zone. Predation effects were quantified by determining prey recolonization rates in marked quadrats in the same large plots. Spatial variation in interaction strength was quantified by examining predation at scales of metres (among transplants within plots), lO's of metres (between replicate plots within each exposure at each site), 1 OO's of metres (between wave exposures within locations), and 10 OOO's of metres (between sites). Temporal variation was evaluated by performing the experiments in 1990 and 1991. The relation between prey (mussel) recruitment and growth to differences in community structure was evaluated by quantifying recruitment density in plastic mesh balls (collectors) and growth of individually marked transplanted mussels, respectively, at each site x exposure x tide level combination each month for 4 yr.Predation intensity varied greatly at all spatial scales. At the two largest spatial scales ( 1 O's of kilometres, 1 OO's of metres), differences in both survival of transplanted mussels and prey recolonization depended on variation in seastar abundance with site, wave exposure, prey recruitment and growth, and at SH protected, the extent of sand burial. Variation at the two smallest scales (metres, 10's of metres) was high when seastars were scarce and low when seastars were abundant. Transplanted mussels suffered 100% mortality in 2 wk at wave-exposed SH, but took >52 wk at wave-protected BB. Seastar effects on prey recolonization were detected only at the SH wave-exposed site. Here, where prey recruitment and growth were unusually high, the mussel M. trossulus invaded and dominated space within 9 mo. After 14 mo, whelks, which increased in both size and abundance in the absence o...

Summary1. Recent efforts to understand how the patterning of interaction strength affects both structure and dynamics in food webs have highlighted several obstacles to productive synthesis. Issues arise with respect to goals and driving questions, methods and approaches, and placing results in the context of broader ecological theory. 2. Much confusion stems from lack of clarity about whether the questions posed relate to community-level patterns or to species dynamics, and to what authors actually mean by the term 'interaction strength'. Here, we describe the various ways in which this term has been applied and discuss the implications of loose terminology and definition for the development of this field. 3. Of particular concern is the clear gap between theoretical and empirical investigations of interaction strengths and food web dynamics. The ecological community urgently needs to explore new ways to estimate biologically reasonable model coefficients from empirical data, such as foraging rates, body size, metabolic rate, biomass distribution and other species traits. 4. Combining numerical and analytical modelling approaches should allow exploration of the conditions under which different interaction strengths metrics are interchangeable with regard to relative magnitude, system responses, and species identity. 5. Finally, the prime focus on predator-prey links in much of the research to date on interaction strengths in food webs has meant that the potential significance of nontrophic interactions, such as competition, facilitation and biotic disturbance, has been largely ignored by the food web community. Such interactions may be important dynamically and should be routinely included in future food web research programmes.

Darwin's classic image of an ''entangled bank'' of interdependencies among species has long suggested that it is difficult to predict how the loss of one species affects the abundance of others. We show that for dynamical models of realistically structured ecological networks in which pair-wise consumer-resource interactions allometrically scale to the 3 ⁄4 power-as suggested by metabolic theory-the effect of losing one species on another can be predicted well by simple functions of variables easily observed in nature. By systematically removing individual species from 600 networks ranging from 10 -30 species, we analyzed how the strength of 254,032 possible pair-wise species interactions depended on 90 stochastically varied species, link, and network attributes. We found that the interaction strength between a pair of species is predicted well by simple functions of the two species' biomasses and the body mass of the species removed. On average, prediction accuracy increases with network size, suggesting that greater web complexity simplifies predicting interaction strengths. Applied to field data, our model successfully predicts interactions dominated by trophic effects and illuminates the sign and magnitude of important nontrophic interactions.

Organisms eating each other are only one of many types of well documented and important interactions among species. Other such types include habitat modification, predator interference and facilitation. However, ecological network research has been typically limited to either pure food webs or to networks of only a few (<3) interaction types. The great diversity of non-trophic interactions observed in nature has been poorly addressed by ecologists and largely excluded from network theory. Herein, we propose a conceptual framework that organises this diversity into three main functional classes defined by how they modify specific parameters in a dynamic food web model. This approach provides a path forward for incorporating non-trophic interactions in traditional food web models and offers a new perspective on tackling ecological complexity that should stimulate both theoretical and empirical approaches to understanding the patterns and dynamics of diverse species interactions in nature.