Weak interactions, omnivory and emergent food-web properties

Empirical studies have shown that, in real ecosystems, species-interaction strengths are generally skewed in their distribution towards weak interactions. Some theoretical work also suggests that weak interactions, especially in omnivorous links, are important for the local stability of a community at equilibrium. However, the majority of theoretical studies use uniform distributions of interaction strengths to generate artificial communities for study. We investigate the effects of the underlying interaction-strength distribution upon the return time, permanence and feasibility of simple Lotka-Volterra equilibrium communities. We show that a skew towards weak interactions promotes local and global stability only when omnivory is present. It is found that skewed interaction strengths are an emergent property of stable omnivorous communities, and that this skew towards weak interactions creates a dynamic constraint maintaining omnivory. Omnivory is more likely to occur when omnivorous interactions are skewed towards weak interactions. However, a skew towards weak interactions increases the return time to equilibrium, delays the recovery of ecosystems and hence decreases the stability of a community. When no skew is imposed, the set of stable omnivorous communities shows an emergent distribution of skewed interaction strengths. Our results apply to both local and global concepts of stability and are robust to the definition of a feasible community. These results are discussed in the light of empirical data and other theoretical studies, in conjunction with their broader implications for community assembly.     

Extinction cascades and the distribution of species interactions

The distribution of interaction strengths among community members has important consequences for assembly processes and community responses to perturbations. Species deletion from communities can trigger cascading extinction events, with strong evidence from empirical and theoretical work. I examined model competitive communities, sequentially assembled using species drawn from a global pool with interaction strengths described by different distribution shapes (uniform or beta), with the same mean and variance. As community size increased, it became harder to assemble communities drawn from a uniform distribution compared to a beta distribution. The distribution of interaction values in the assembled communities differed from the shape of the initial distribution. The distribution shape and the relative abundance of the deleted species also had strong impacts on the probability of extinction cascades following primary species removal. Extinction cascades occurred in communities with a higher mean and variance of interaction strengths before the primary extinction. Those species lost had negative equilibrium densities and tended to be the least abundant, when assessed following the reorganisation that occurred after the primary and subsequent extinctions. Knowledge of the shape of the distribution of interaction strengths from real communities will allow us to make better predictions about which species are most at risk in extinction cascades under natural circumstances.     

Stability of a diamond-shaped module with multiple interaction types

Indirect interactions among species emerge from the complexity of ecological networks and can strongly affect the response of communities to disturbances. To determine these indirect interactions and understand better community dynamics, ecologists focused on the interactions within small sets of species or modules. Thanks to their analytical tractability, modules bring insights on the mechanisms occurring in complex interaction networks. So far, most studies have considered modules with a single type of interaction although numerous species are involved in mutualistic and antagonistic interactions simultaneously. In this study, we analyse the dynamics of a diamond-shaped module with multiple interaction types: two resource species sharing a mutualist and a consumer. We describe the different types of indirect interaction occurring between the resource species and the conditions for a stable coexistence of all species. We show that the nature of indirect interactions between resource species (i.e. apparent facilitation, competition or antagonism), as well as stable coexistence, depend on the species generalism and asymmetry of interactions, or in other words, on the distribution of interaction strengths among species. We further unveil that a balance between mutualistic and antagonistic interactions at the level of resource species favours stable coexistence, and that species are more likely to coexist stably if there is apparent facilitation between the two resource species rather than apparent competition. Our results echo existing knowledge on the trophic diamond-shaped module, and confirm that our understanding of communities combining different interaction types can gain from module analyses.     

Disentangling the effect of hybrid interactions and of the constant effort hypothesis on ecological community stability

In the last years, a remarkable theoretical effort has been made in order to understand the relation between stability and complexity in ecological communities. Yet, what maintains species diversity in real ecological communities is still an open question. The non‐random structures of ecological interaction networks have been recognized as one key ingredient impacting the maximum number of coexisting species within the ecological community. However most of the earlier theoretical studies have considered communities with only one interaction type (either antagonistic, competitive or mutualistic). Recently, it has been proposed that multiple interaction types might stabilize ecosystems and that, in this hybrid case, increasing complexity increases stability. Here we show that these results depend on ad hoc hypothesis that the authors used in their model and we highlight the need to disentangle the role of multiple interaction types and constant interaction effort allocation on community stability. Indeed, we find that mixing of mutualistic and predator–prey interaction types does not stabilize the community dynamics and we demonstrate that a positive correlation between complexity and stability is observed only if a constant effort allocation is imposed in the ecological interactions. Synthesis In recent years a sparkling research has been devoted to the search of new theoretical mechanisms to explain way ecosystems may persist despite their complexity. Here we show that, contrary to what recently suggested (Mougi et al. 2012), the mismatch between theoretical results and empirical evidences on the stability of ecological community is still there also for communities with both mutualistic and antagonistic interactions, and the ‘complexity‐stability’ paradox is still alive. Indeed, we demonstrate that their results arise as an artifact of the peculiar rescaling of the interaction strengths they imposed. Our study suggests that further theoretical studies and experimental evidences are still needed to better understand the role of interaction strengths in real ecological communities.     

Spatial covariance of herbivorous and predatory guilds of forest canopy arthropods along a latitudinal gradient

In arthropod community ecology, species richness studies tend to be prioritised over those investigating patterns of abundance. Consequently, the biotic and abiotic drivers of arboreal arthropod abundance are still relatively poorly known. In this cross‐continental study, we employ a theoretical framework in order to examine patterns of covariance among herbivorous and predatory arthropod guilds. Leaf‐chewing and leaf‐mining herbivores, and predatory ants and spiders, were censused on > 1000 trees in nine 0.1 ha forest plots. After controlling for tree size and season, we found no negative pairwise correlations between guild abundances per plot, suggestive of weak signals of both inter‐guild competition and top‐down regulation of herbivores by predators. Inter‐guild interaction strengths did not vary with mean annual temperature, thus opposing the hypothesis that biotic interactions intensify towards the equator. We find evidence for the bottom‐up limitation of arthropod abundances via resources and abiotic factors, rather than for competition and predation.     

Landscape diversity promotes stable food-web architectures in large rivers

Uncovering relationships between landscape diversity and species interactions is crucial for predicting how ongoing land-use change and homogenization will impact the stability and persistence of communities. However, such connections have rarely been quantified in nature. We coupled high-resolution river sonar imaging with annualized energetic food webs to quantify relationships among habitat diversity, energy flux, and trophic interaction strengths in large-river food-web modules that support the endangered Pallid Sturgeon. Our results demonstrate a clear relationship between habitat diversity and species interaction strengths, with more diverse foraging landscapes containing higher production of prey and a greater proportion of weak and potentially stabilizing interactions. Additionally, rare patches of large and relatively stable river sediments intensified these effects and further reduced interaction strengths by increasing prey diversity. Our findings highlight the importance of landscape characteristics in promoting stabilizing food-web architectures and provide direct relevance for future management of imperilled species in a simplified and rapidly changing world.     

Network structural properties mediate the stability of mutualistic communities

Key advances are being made on the structures of predator–prey food webs and competitive communities that enhance their stability, but little attention has been given to such complexity–stability relationships for mutualistic communities. We show, by way of theoretical analyses with empirically informed parameters, that structural properties can alter the stability of mutualistic communities characterized by nonlinear functional responses among the interacting species. Specifically, community resilience is enhanced by increasing community size (species diversity) and the number of species interactions (connectivity), and through strong, symmetric interaction strengths of highly nested networks. As a result, mutualistic communities show largely positive complexity–stability relationships, in opposition to the standard paradox. Thus, contrary to the commonly-held belief that mutualism's positive feedback destabilizes food webs, our results suggest that interplay between the structure and function of ecological networks in general, and consideration of mutualistic interactions in particular, may be key to understanding complexity–stability relationships of biological communities as a whole.     

Competitive interactions between forest trees are driven by species' trait hierarchy, not phylogenetic or functional similarity: implications for forest community assembly

The relative importance of competition vs. environmental filtering in the assembly of communities is commonly inferred from their functional and phylogenetic structure, on the grounds that similar species compete most strongly for resources and are therefore less likely to coexist locally. This approach ignores the possibility that competitive effects can be determined by relative positions of species on a hierarchy of competitive ability. Using growth data, we estimated 275 interaction coefficients between tree species in the French mountains. We show that interaction strengths are mainly driven by trait hierarchy and not by functional or phylogenetic similarity. On the basis of this result, we thus propose that functional and phylogenetic convergence in local tree community might be due to competition-sorting species with different competitive abilities and not only environmental filtering as commonly assumed. We then show a functional and phylogenetic convergence of forest structure with increasing plot age, which supports this view.     

Nonmanipulative determination of plant community dynamics

Knowledge of the strengths of interactions between species in plant communities is of fundamental importance to our understanding of how communities are structured, although they are notoriously difficult to quantify. Techniques have recently been developed that allow the detailed enumeration of the strength of interactions between plant species within unmanipulated multispecies communities. Nonlinear regression analysis is used to fit competition models to long-term census data using natural variations in plant densities in lieu of manipulation. The models generated have been used to infer the intensity and importance of interactions as well as to analyse the effects of spatial and temporal variability. Theoretical work has begun to look at how different techniques for measuring competition perform in a range of systems, highlighting the importance of spatial scale. The lessons learned from applying these methods will enable improved estimation of the strength of competition in natural communities.     

Predator diversity and environmental change modify the strengths of trophic and nontrophic interactions

Understanding the dependence of species interaction strengths on environmental factors and species diversity is crucial to predict community dynamics and persistence in a rapidly changing world. Nontrophic (e.g. predator interference) and trophic components together determine species interaction strengths, but the effects of environmental factors on these two components remain largely unknown. This impedes our ability to fully understand the links between environmental drivers and species interactions. Here, we used a dynamical modelling framework based on measured predator functional responses to investigate the effects of predator diversity, prey density, and temperature on trophic and nontrophic interaction strengths within a freshwater food web. We found that (i) species interaction strengths cannot be predicted from trophic interactions alone, (ii) nontrophic interaction strengths vary strongly among predator assemblages, (iii) temperature has opposite effects on trophic and nontrophic interaction strengths, and (iv) trophic interaction strengths decrease with prey density, whereas the dependence of nontrophic interaction strengths on prey density is concave up. Interestingly, the qualitative impacts of temperature and prey density on the strengths of trophic and nontrophic interactions were independent of predator identity, suggesting a general pattern. Our results indicate that taking multiple environmental factors and the nonlinearity of density‐dependent species interactions into account is an important step towards a better understanding of the effects of environmental variations on complex ecological communities. The functional response approach used in this study opens new avenues for (i) the quantification of the relative importance of the trophic and nontrophic components in species interactions and (ii) a better understanding how environmental factors affect these interactions and the dynamics of ecological communities.     

Community assembly, stability and signatures of dynamical constraints on food web structure

To understand the dynamics of natural species communities, a major challenge is to quantify the relationship between their assembly, stability, and underlying food web structure. To this end, two complementary aspects of food web structure can be related to community stability: sign structure, which refers to the distributions of trophic links irrespective of interaction strengths, and interaction strength structure, which refers to the distributions of interaction strengths with or without consideration of sign structure. In this paper, using data from a set of relatively well documented community food webs, I show that natural communities generally exhibit a sign structure that renders their stability sensitive to interaction strengths. Using a Lotka-Volterra type population dynamical model, I then show that in such communities, individual consumer species with high values of a measure of their total biomass acquisition rate, which I term “weighted generality”, tend to undermine community stability. Thus consumer species’ trophic modules (a species and all its resource links) should be “selected” through repeated immigrations and extinctions during assembly into configurations that increase the probability of stable coexistence within the constraints of the community's trophic sign structure. The presence of such constraints can be detected by the incidence and strength of certain non-random structural characteristics. These structural signatures of dynamical constraints are readily measurable, and can be used to gauge the importance of interaction-driven dynamical constraints on communities during and after assembly in natural communities.     

Topological properties of food webs: from real data to community assembly models

We explore patterns of trophic connections between species in the largest and highest-quality empirical food webs to date, introducing a new topological property called the link distribution frequency (i.e. degree distribution), defined as the frequency of species S _L with L links. Non-trivial differences are shown in link distribution frequencies between species-rich and species-poor communities, which might have important consequences for the responses of ecosystems to disturbances. Coarse-grained topological properties observed, as species richness-connectance and number of links-species richness relationships, provide no support for the theory of links-species scaling law or constant connectance across empirical food webs investigated. We further explore these observations by means of simulated food webs resulting from multitrophic assembly models using different functional responses between species. Species richness-connectance and links-species richness relationships of empirical food webs are reproduced by our models, but degree distributions are not properly predicted, suggesting the need of new theoretical approximations to food web assembly. The best agreement between empirical and simulated webs occurs for low values of interaction strength between species, corroborating previous empirical and theoretical findings where weak interactions govern food web dynamics.     

Threshold facilitations of interacting species

The dynamics of species interactions are of central importance for the understanding of ecological coexistence, community structure and the effects of biological invasions. Using bark beetles that colonize the same habitat as an example, we explore species interactions in a resource-based model system with positive feedback between insect abundance and resource availability. The net interspecies interaction was found to be highly dynamic and may alternate in time between competition and mutualism. When both bark beetle species were able to kill trees (“aggressive”), our simulations showed strong facilitations between beetle species. This may lead to escape from control by competition, and increase the frequency of outbreaks of tree-killing. The frequency of net positive interactions varied with interaction strengths and the relative aggressiveness of the species and was highest when both species were strongly aggressive; which predicts disastrous outbreaks if, e.g., the European spruce bark beetle Ips typographus and the North American spruce beetle Dendroctonus rufipennis should become interacting species due to introductions. In imbalanced pairs, the relatively less aggressive species was facilitated more often than the aggressive species. Net positive interactions did not occur for strongly inferior species, but their survival was an increasing function of interaction strength with aggressive species and availability of resources. The benefits for the inferior species in the model are consistent with the structure of one aggressive and several less aggressive or non-aggressive species, which is common in bark beetle communities in many parts of the world.     

Examining the effects of species richness on community stability: an assembly model approach

We build dynamic models of community assembly by starting with one species in our model ecosystem and adding colonists. We find that the number of species present first increases, then fluctuates about some level. We ask: how large are these fluctuations and how can we characterize them statistically? As in Robert May's work, communities with weaker interspecific interactions permit a greater number of species to coexist on average. We find that as this average increases, however, the relative variation in the number of species and return times to mean community levels decreases. In addition, the relative frequency of large extinction events to small extinction events decreases as mean community size increases. While the model reproduces several of May's results, it also provides theoretical support for Charles Elton's idea that diverse communities such as those found in the tropics should be less variable than depauperate communities such as those found in arctic or agricultural settings.     

Novel communities from climate change

Climate change is generating novel communities composed of new combinations of species. These result from different degrees of species adaptations to changing biotic and abiotic conditions, and from differential range shifts of species. To determine whether the responses of organisms are determined by particular species traits and how species interactions and community dynamics are likely to be disrupted is a challenge. Here, we focus on two key traits: body size and ecological specialization. We present theoretical expectations and empirical evidence on how climate change affects these traits within communities. We then explore how these traits predispose species to shift or expand their distribution ranges, and associated changes on community size structure, food web organization and dynamics. We identify three major broad changes: (i) Shift in the distribution of body sizes towards smaller sizes, (ii) dominance of generalized interactions and the loss of specialized interactions, and (iii) changes in the balance of strong and weak interaction strengths in the short term. We finally identify two major uncertainties: (i) whether large-bodied species tend to preferentially shift their ranges more than small-bodied ones, and (ii) how interaction strengths will change in the long term and in the case of newly interacting species.     

Community assembly time and the relationship between local and regional species richness

Many previous studies have assumed that a linear relationship between local and regional species richness indicates that communities are limited by regional processes, while a saturating relationship suggests that species interactions restrict local richness. We show theoretically that the relationship between local and regional richness changes in a consistent fashion with assembly time in interacting communities. Communities show saturation in their early assembly stages because only a subset of the regional pool may colonize a locality. At intermediate assembly times, communities will appear unsaturated until significant competitive exclusion occurs. Finally, when communities reach equilibrium, we found saturation as a result of resource competition resulting in the dominance of a limited number of species. We show that habitat size and species fecundity are important in determining the time needed for the community to reach equilibrium and thus affect the relationship between local and regional species richness. Our results suggest the number of coexisting species is a function of local and regional processes whose relative influences might vary over time and that research using the relationship between local and regional species richness to infer mechanisms limiting species richness must have knowledge of the assembly time of the community.     

Competitive interactions change the pattern of species co‐occurrences under neutral dispersal

Non‐random patterns of species segregation and aggregation within ecological communities are often interpreted as evidence for interspecific interactions. However, it is unclear whether theoretical models can predict such patterns and how environmental factors may modify the effects of species interactions on species co‐occurrence. Here we extend a spatially explicit neutral model by including competitive effects on birth and death probabilities to assess whether competition alone is able to produce non‐random patterns of species co‐occurrence. We show that transitive and intransitive competitive hierarchies alone (in the absence of environmental heterogeneity) are indeed able to generate non‐random patterns with commonly used metrics and null models. Moreover, even weak levels of intransitive competition can increase local species richness. However, there is no simple rule or consistent directional change towards aggregation or segregation caused by competitive interactions. Instead, the spatial pattern depends on both the type of species interaction and the strength of dispersal. We conclude that co‐occurrence analysis alone may not able to identify the underlying processes that generate the patterns.     

Positive interactions among competitors can produce species-rich communities

Although positive interactions between species are well documented, most ecological theory for investigating multispecies coexistence remains rooted in antagonistic interactions such as competition and predation. Standard resource-competition models from this theory predict that the number of coexisting species should not exceed the number of factors that limit population growth. Here I show that positive interactions among resource competitors can produce species-rich model communities supported by a single limiting resource. Simulations show that when resource competitors reduce each others' per capita mortality rate (e.g. by ameliorating an abiotic stress), stable multispecies coexistence with a single resource may be common, even while the net interspecific interaction remains negative. These results demonstrate that positive interactions may provide an important mechanism for generating species-rich communities in nature. They also show that focusing on the net interaction between species may conceal important coexistence mechanisms when species simultaneously engage in both antagonistic and positive interactions.     

Do extrafloral nectar resources, species abundances, and body sizes contribute to the structure of ant–plant mutualistic networks?

Recent research has shown that many mutualistic communities display non-random structures. While our understanding of the structural properties of mutualistic communities continues to improve, we know little of the biological variables resulting in them. Mutualistic communities include those formed between ants and extrafloral (EF) nectar-bearing plants. In this study, we examined the contributions of plant and ant abundance, plant and ant size, and plant EF nectar resources to the network structures of nestedness and interaction frequency of ant–plant networks across five sites within one geographic locality in the Sonoran Desert. Interactions between ant and plant species were largely symmetric. That is, ant and plant species exerted nearly equivalent quantitative interaction effects on one another, as measured by their frequency of interaction. The mutualistic ant–plant networks also showed nested patterns of structure, in which there was a central core of generalist ant and plant species interacting with one another and few specialist–specialist interactions. Abundance and plant size and ant body size were the best predictors of symmetric interactions between plants and ants, as well as nestedness. Despite interactions in these communities being ultimately mediated by EF nectar resources, the number of EF nectaries had a relatively weak ability to explain variation in symmetric interactions and nestedness. These results suggest that different mechanisms may contribute to structure of bipartite networks. Moreover, our results for ant–plant mutualistic networks support the general importance of species abundances for the structure of species interactions within biological communities.