Food web models: a plea for groups

The concept of a group is ubiquitous in biology. It underlies classifications in evolution and ecology, including those used to describe phylogenetic levels, the habitat and functional roles of organisms in ecosystems. Surprisingly, this concept is not explicitly included in simple models for the structure of food webs, the ecological networks formed by consumer–resource interactions. We present here the simplest possible model based on groups, and show that it performs substantially better than current models at predicting the structure of large food webs. Our group-based model can be applied to different types of biological and non-biological networks, and for the first time merges in the same framework two important notions in network theory: that of compartments (sets of highly interacting nodes) and that of roles (sets of nodes that have similar interaction patterns). This model provides a basis to examine the significance of groups in biological networks and to develop more accurate models for ecological network structure. It is especially relevant at a time when a new generation of empirical data is providing increasingly large food webs.     

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.     

Phenology drives mutualistic network structure and diversity

Several network properties have been identified as determinants of the stability and complexity of mutualistic networks. However, it is unclear which mechanisms give rise to these network properties. Phenology seems important, because it shapes the topology of mutualistic networks, but its effects on the dynamics of mutualistic networks have scarcely been studied. Here, we study these effects with a general dynamical model of mutualistic and competitive interactions where the interaction strength depends on the temporal overlap between species resulting from their phenologies. We find a negative complexity-stability relationship, where phenologies maximising mutualistic interactions and minimising intraguild competitive interactions generate speciose, nested and poorly connected networks with moderate asymmetry and low resilience. Moreover, lengthening the season increases diversity and resilience. This highlights the fragility of real mutualistic communities with short seasons (e.g. Arctic environments) to drastic environmental changes.     

Parasites reduce food web robustness because they are sensitive to secondary extinction as illustrated by an invasive estuarine snail

A robust food web is one in which few secondary extinctions occur after removing species. We investigated how parasites affected the robustness of the Carpinteria Salt Marsh food web by conducting random species removals and a hypothetical, but plausible, species invasion. Parasites were much more likely than free-living species to suffer secondary extinctions following the removal of a free-living species from the food web. For this reason, the food web was less robust with the inclusion of parasites. Removal of the horn snail, Cerithidea californica, resulted in a disproportionate number of secondary parasite extinctions. The exotic Japanese mud snail, Batillaria attramentaria, is the ecological analogue of the native California horn snail and can completely replace it following invasion. Owing to the similarities between the two snail species, the invasion had no effect on predator–prey interactions. However, because the native snail is host for 17 host-specific parasites, and the invader is host to only one, comparison of a food web that includes parasites showed significant effects of invasion on the native community. The hypothetical invasion also significantly reduced the connectance of the web because the loss of 17 native trematode species eliminated many links.     

Do bipartite binary antagonistic and mutualistic networks have different responses to the taxonomic resolution of nodes?

1. Bipartite network analyses are increasingly being used to better understand mutualistic and antagonistic plant–insect interactions at the community level. As a result of taxonomic limitations, it is usually very difficult to identify all nodes of a network down to the species level and many studies leave some specimens identified as lower resolution taxa. Accordingly, we do not know how much a lower resolution taxonomic representation changes the network structure compared with a representation with all nodes at species level. 2. The present study aimed to test whether insect–plant networks built using different combinations of taxonomic levels can still preserve the same basic structure of networks built only with species. 3. In total, 73 bipartite published interaction networks (mutualistic and antagonistic) were selected, which were turned into binary networks and reconstructed using the nodes classified as species, genus, family or order (representing different levels of classification difficulty). The network structures were compared using their binary representations mainly using connectance, NODF (Nestedness metric based on Overlap and Decreasing Fill) and modularity. 4. The mutualistic network structure was strongly linearly related to the original network structures if all nodes were grouped up to genus level. In antagonistic networks, the structure was related to the original network only if nodes were only grouped at the species level. 5. The findings of the present study are especially helpful for comparative network studies, such as those assessing the effects of environmental gradients. For mutualistic networks, Citizen Science programmes can provide useful ecological indicators, even with its taxonomic limitations.     

基于结构洞理论的产业生态群落关联度赋值方法

产业生态系统稳定性的度量是工业生态学研究领域的热点和难点问题.在生态群落关联度的基础上,借鉴社会网络理论中的"结构洞"思想,构造了考虑权重的产业生态群落关联度.对卡伦堡和安阳高新区生态工业园区的案例研究表明,与不考虑权重的关联度指标比较,考虑权重的关联度能够更好地体现节点和链接重要性的不同,由此可以为生态工业园区规划提供了更好的定量化指标.     

Flower-heads, herbivores, and their parasitoids: food web structure along a fertility gradient

Abstract. 1. The ways in which a soil fertility gradient affects three trophic level food webs defined by plants of the family Asteraceae, flower‐head herbivores, and their parasitoids was investigated. It was tested how the fertility gradient alters: (i) the abundance and richness of plants, herbivores, and their parasitoids, (ii) the herbivore–plant ratio, and (iii) the connectance of the plant–herbivore community matrix. 2. From April to May 2000, plants and insects were sampled in 16 Brazilian Cerrado (sensu stricto) sites along a physiognomic gradient varying from open shrublands (cerrado) to closed woodlands (cerradão). Sites were objectively positioned along the physiognomic gradient by a single index, tree density. Sixty‐seven per cent of the variation in tree density among sites was correlated to two principal components of a PCA, representing gradients of soil fertility. 3. Asteraceae abundance, richness, and flower‐head availability were negatively related to tree density due to their preference for sunny environments, despite the surplus of soil nutrients. The abundance and richness of Diptera and Lepidoptera, the most important flower‐head herbivores, were also negatively related to tree density. Parasitoid abundance decreased with tree density; however, the number of parasitoids per hosts was lower in cerrado, suggesting that top‐down forces are not getting stronger in more productive sites, as could be expected. 4. Community allometry analyses showed that the herbivore to plant ratio was independent of community richness and did not respond to tree density. 5. Connectance of the plant–herbivore matrix was dependent on the community matrix size. Proportionally, species‐rich cerrado sites had fewer interactions than their species‐poor counterparts. Nevertheless, after removing the effect of the matrix size, connectance was not related to tree density. 6. Soil fertility, as the primary cause of the cerrado–cerradão physiognomic gradient, strongly affected the abundance and richness of plants, herbivores and their parasitoids; however, it had little effect on important community attributes, such as the herbivore–plant ratio and the connectance of the plant–herbivore matrix.     

Maturation of integrated functions during development. I. Modifications of the regulatory network during transition periods in Sorghum bicolor

Gibberellin (GA) and cytokinin (CK) were exogenously supplied at different periods of the vegetative development in Sorghum bicolor. Growth response to these hormonal treatments differed according to the developmental stage. This reveals the existence of discrete phenophases, each one characterized by a specific sensitivity to plant growth regulations (PGRs). Developmental changes in sensitivity were less accentuated in plants grown in optimal conditions than in plants exposed to 150 mM NaCl. Variations in organ connectance (the level of coordination in growth of the shoot, adventitious roots and seminal root) were analysed during vegetative growth of salt‐treated plants. This analysis shows a temporary decrease in connectance during the transition period between phenophases. From the effect of hormonal treatments on connectance, it was concluded that (i) the transition period coincides with a partial dismantling of the initial regulatory network followed by the emergence of a new network coordinating growth of the different organs; (ii) GA is involved in the process of emergence of a transition period; and (iii) duration of the transition period is considerably enlarged for plants exposed to NaCl stress. This dynamics of alternance of phenophase and transition periods enables the integration of the two modes of action of the PGRs (dose–response and change in sensitivity) within a unified framework.     

Food-web structure in two shallow salt lakes in Los Monegros (NE Spain): energetic vs dynamic constraints

Energetic and dynamic constraints have been proposed as rival factors in determining food-web structure. Food-web length might be controlled either by the amount of energy entering the web (energetic constraints) or by time span between consecutive disturbances relative to time needed to build up a population (dynamic constraints). Dynamic constraints are identified with processes functioning at a regional scale such as climate, lithology and hydrogeology. Energetic constraints are related with processes operating both at a regional and a local scale. We studied the contribution of energetic constraints to food-web organization in two temporary saline lakes with similar dynamic constraints. Lakes were sampled fortnightly during two hydroperiods (1994/1995 and 1995/1996). Differences in energetic constraints between lakes result in divergent assemblages of primary producers. Consumer assemblages in both lakes, however, are similar in species composition although differ in total biomass and species abundances. Food-webs are short with a high proportion of omnivores. To simulate an increase in the energy input entering to these systems, an addition of nutrients (to a final concentration of 100 gl^–1 P-PO₄ ^3-) was done in mesocosms placed within the lakes in order to obtain an increase in the phytoplankton biomass. No significant response to nutrient enrichment was found in food-web structure (composition, density or biomass).