High frequency of functional extinctions in ecological networks.

Intensified exploitation of natural populations and habitats has led to increased mortality rates and decreased abundances of many species. There is a growing concern that this might cause critical abundance thresholds of species to be crossed, with extinction cascades and state shifts in ecosystems as a consequence. When increased mortality rate and decreased abundance of a given species lead to extinction of other species, this species can be characterized as functionally extinct even though it still exists. Although such functional extinctions have been observed in some ecosystems, their frequency is largely unknown. Here we use a new modelling approach to explore the frequency and pattern of functional extinctions in ecological networks. Specifically, we analytically derive critical abundance thresholds of species by increasing their mortality rates until an extinction occurs in the network. Applying this approach on natural and theoretical food webs, we show that the species most likely to go extinct first is not the one whose mortality rate is increased but instead another species. Indeed, up to 80% of all first extinctions are of another species, suggesting that a species' ecological functionality is often lost before its own existence is threatened. Furthermore, we find that large-bodied species at the top of the food chains can only be exposed to small increases in mortality rate and small decreases in abundance before going functionally extinct compared to small-bodied species lower in the food chains. These results illustrate the potential importance of functional extinctions in ecological networks and lend strong support to arguments advocating a more community-oriented approach in conservation biology, with target levels for populations based on ecological functionality rather than on mere persistence.

Stepping in Elton's footprints: a general scaling model for body masses and trophic levels across ecosystems.

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.

Response of ecosystems to realistic extinction sequences.

Recent research suggests that effects of species loss on the structure and functioning of ecosystems will critically depend on the order with which species go extinct. However, there are few studies of the response of natural ecosystems to realistic extinction sequences. Using an extinction scenario based on the International Union for Conservation of Nature (IUCN) Red List, de Visser et al. sequentially deleted species from a topological model of the Serengeti food web. Under this scenario, large-bodied species like top predators and mega-herbivores go extinct first. The resulting changes in the trophic structure of the food web might affect the robustness of the ecosystem to future disturbances.

The importance of species interactions in eco-evolutionary community dynamics under climate change.

Eco-evolutionary dynamics are essential in shaping the biological response of communities to ongoing climate change. Here we develop a spatially explicit eco-evolutionary framework which features more detailed species interactions, integrating evolution and dispersal. We include species interactions within and between trophic levels, and additionally, we incorporate the feature that species' interspecific competition might change due to increasing temperatures and affect the impact of climate change on ecological communities. Our modeling framework captures previously reported ecological responses to climate change, and also reveals two key results. First, interactions between trophic levels as well as temperature-dependent competition within a trophic level mitigate the negative impact of climate change on biodiversity, emphasizing the importance of understanding biotic interactions in shaping climate change impact. Second, our trait-based perspective reveals a strong positive relationship between the within-community variation in preferred temperatures and the capacity to respond to climate change. Temperature-dependent competition consistently results both in higher trait variation and more responsive communities to altered climatic conditions. Our study demonstrates the importance of species interactions in an eco-evolutionary setting, further expanding our knowledge of the interplay between ecological and evolutionary processes.

A potential role for rare species in ecosystem dynamics.

The ecological importance of common species for many ecosystem processes and functions is unquestionably due to their high abundance. Yet, the importance of rare species is much less understood. Here we take a theoretical approach, exposing dynamical models of ecological networks to small perturbations, to explore the dynamical importance of rare and common species. We find that both species types contribute to the recovery of communities following generic perturbations (i.e. perturbations affecting all species). Yet, when perturbations are selective (i.e. affects only one species), perturbations to rare species have the most pronounced effect on community stability. We show that this is due to the strong indirect effects induced by perturbations to rare species. Because indirect effects typically set in at longer timescales, our results indicate that the importance of rare species may be easily overlooked and thus underrated. Hence, our study provides a potential ecological motive for the management and protection of rare species.

Trophically unique species are vulnerable to cascading extinction.

Understanding which species might become extinct and the consequences of such loss is critical. One consequence is a cascade of further, secondary extinctions. While a significant amount is known about the types of communities and species that suffer secondary extinctions, little is known about the consequences of secondary extinctions for biodiversity. Here we examine the effect of these secondary extinctions on trophic diversity, the range of trophic roles played by the species in a community. Our analyses of natural and model food webs show that secondary extinctions cause loss of trophic diversity greater than that expected from chance, a result that is robust to variation in food web structure, distribution of interactions strengths, functional response, and adaptive foraging. Greater than expected loss of trophic diversity occurs because more trophically unique species are more vulnerable to secondary extinction. This is not a straightforward consequence of these species having few links with others but is a complex function of how direct and indirect interactions affect species persistence. A positive correlation between a species' extinction probability and the importance of its loss defines high-risk species and should make their conservation a priority.

Early onset of secondary extinctions in ecological communities following the loss of top predators.

The large vulnerability of top predators to human-induced disturbances on ecosystems is a matter of growing concern. Because top predators often exert strong influence on their prey populations their extinction can have far-reaching consequences for the structure and functioning of ecosystems. It has, for example, been observed that the local loss of a predator can trigger a cascade of secondary extinctions. However, the time lags involved in such secondary extinctions remain unexplored. Here we show that the loss of a top predator leads to a significantly earlier onset of secondary extinctions in model communities than does the loss of a species from other trophic levels. Moreover, in most cases time to secondary extinction increases with increasing species richness. If local secondary extinctions occur early they are less likely to be balanced by immigration of species from local communities nearby. The implications of these results for community persistence and conservation priorities are discussed.

Adaptive rewiring aggravates the effects of species loss in ecosystems.

Loss of one species in an ecosystem can trigger extinctions of other dependent species. For instance, specialist predators will go extinct following the loss of their only prey unless they can change their diet. It has therefore been suggested that an ability of consumers to rewire to novel prey should mitigate the consequences of species loss by reducing the risk of cascading extinction. Using a new modelling approach on natural and computer-generated food webs we find that, on the contrary, rewiring often aggravates the effects of species loss. This is because rewiring can lead to overexploitation of resources, which eventually causes extinction cascades. Such a scenario is particularly likely if prey species cannot escape predation when rare and if predators are efficient in exploiting novel prey. Indeed, rewiring is a two-edged sword; it might be advantageous for individual predators in the short term, yet harmful for long-term system persistence.

Confronting the paradox of enrichment to the metacommunity perspective.

Resource enrichment can potentially destabilize predator-prey dynamics. This phenomenon historically referred as the "paradox of enrichment" has mostly been explored in spatially homogenous environments. However, many predator-prey communities exchange organisms within spatially heterogeneous networks called metacommunities. This heterogeneity can result from uneven distribution of resources among communities and thus can lead to the spreading of local enrichment within metacommunities. Here, we adapted the original Rosenzweig-MacArthur predator-prey model, built to study the paradox of enrichment, to investigate the effect of regional enrichment and of its spatial distribution on predator-prey dynamics in metacommunities. We found that the potential for destabilization was depending on the connectivity among communities and the spatial distribution of enrichment. In one hand, we found that at low dispersal regional enrichment led to the destabilization of predator-prey dynamics. This destabilizing effect was more pronounced when the enrichment was uneven among communities. In the other hand, we found that high dispersal could stabilize the predator-prey dynamics when the enrichment was spatially heterogeneous. Our results illustrate that the destabilizing effect of enrichment can be dampened when the spatial scale of resource enrichment is lower than that of organismss movements (heterogeneous enrichment). From a conservation perspective, our results illustrate that spatial heterogeneity could decrease the regional extinction risk of species involved in specialized trophic interactions. From the perspective of biological control, our results show that the heterogeneous distribution of pest resource could favor or dampen outbreaks of pests and of their natural enemies, depending on the spatial scale of heterogeneity.

Species-rich ecosystems are vulnerable to cascading extinctions in an increasingly variable world.

Global warming leads to increased intensity and frequency of weather extremes. Such increased environmental variability might in turn result in increased variation in the demographic rates of interacting species with potentially important consequences for the dynamics of food webs. Using a theoretical approach, we here explore the response of food webs to a highly variable environment. We investigate how species richness and correlation in the responses of species to environmental fluctuations affect the risk of extinction cascades. We find that the risk of extinction cascades increases with increasing species richness, especially when correlation among species is low. Initial extinctions of primary producer species unleash bottom-up extinction cascades, especially in webs with specialist consumers. In this sense, species-rich ecosystems are less robust to increasing levels of environmental variability than species-poor ones. Our study thus suggests that highly species-rich ecosystems such as coral reefs and tropical rainforests might be particularly vulnerable to increased climate variability.

Species loss and secondary extinctions in simple and complex model communities.

1. The loss of a species from an ecological community can trigger a cascade of secondary extinctions. Here we investigate how the complexity (connectance) of model communities affects their response to species loss. Using dynamic analysis based on a global criterion of persistence (permanence) and topological analysis we investigate the extent of secondary extinctions following the loss of different kinds of species. 2. We show that complex communities are, on average, more resistant to species loss than simple communities: the number of secondary extinctions decreases with increasing connectance. However, complex communities are more vulnerable to loss of top predators than simple communities. 3. The loss of highly connected species (species with many links to other species) and species at low trophic levels triggers, on average, the largest number of secondary extinctions. The effect of the connectivity of a species is strongest in webs with low connectance. 4. Most secondary extinctions are due to direct bottom-up effects: consumers go extinct when their resources are lost. Secondary extinctions due to trophic cascades and disruption of predator-mediated coexistence also occur. Secondary extinctions due to disruption of predator-mediated coexistence are more common in complex communities than in simple communities, while bottom-up and top-down extinction cascades are more common in simple communities. 5. Topological analysis of the response of communities to species loss always predicts a lower number of secondary extinctions than dynamic analysis, especially in food webs with high connectance.

Using community viability analysis to identify fragile systems and keystone species.

Owing to interdependences among species in ecological communities, the loss of one species can trigger a cascade of secondary extinctions with potentially dramatic effects on the functioning and stability of the community. It is, therefore, important to assess the risk and likely extent of secondary extinctions. Community viability analysis is a new technique that can be used to accomplish this goal. The analysis can also be used to identify fragile community structures and keystone species and, hence, to provide guidelines for conservation priorities. Here, we describe the principles underlying community viability analysis and review its contributions to our understanding of the response of ecological communities to species loss.

Keystone species and vulnerable species in ecological communities: strong or weak interactors?

The loss of a species from an ecological community can trigger a cascade of secondary extinctions. The probability of secondary extinction to take place and the number of secondary extinctions are likely to depend on the characteristics of the species that is lost--the strength of its interactions with other species--as well as on the distribution of interaction strengths in the whole community. Analysing the effects of species loss in model communities we found that removal of the following species categories triggered, on average, the largest number of secondary extinctions: (a) rare species interacting strongly with many consumers, (b) abundant basal species interacting weakly with their consumers and (c) abundant intermediate species interacting strongly with many resources. We also found that the keystone status of a species with given characteristics was context dependent, that is, dependent on the structure of the community where it was embedded. Species vulnerable to secondary extinctions were mainly species interacting weakly with their resources and species interacting strongly with their consumers.

Body size in ecological networks.

Body size determines a host of species traits that can affect the structure and dynamics of food webs, and other ecological networks, across multiple scales of organization. Measuring body size provides a relatively simple means of encapsulating and condensing a large amount of the biological information embedded within an ecological network. Recently, important advances have been made by incorporating body size into theoretical models that explore food web stability, the patterning of energy fluxes, and responses to perturbations. Because metabolic constraints underpin body-size scaling relationships, metabolic theory offers a potentially useful new framework within which to develop novel models to describe the structure and functioning of ecological networks and to assess the probable consequences of biodiversity change.