Modularity is a recurrent and important property of bipartite ecological networks. Although well-resolved ecological networks describe interaction frequencies between species pairs, modularity of bipartite networks has been analysed only on the basis of binary presence-absence data. We employ a new algorithm to detect modularity in weighted bipartite networks in a global analysis of avian seed-dispersal networks. We define roles of species, such as connector values, for weighted and binary networks and associate them with avian species traits and phylogeny. The weighted, but not binary, analysis identified a positive relationship between climatic seasonality and modularity, whereas past climate stability and phylogenetic signal were only weakly related to modularity. Connector values were associated with foraging behaviour and were phylogenetically conserved. The weighted modularity analysis demonstrates the dominating impact of ecological factors on the structure of seed-dispersal networks, but also underscores the relevance of evolutionary history in shaping species roles in ecological communities.

Interactions between resource and consumer species are organized in ecological networks. Species interactions in these networks are influenced by the functional traits of the interacting partners, but the generality of trait-based interaction rules and the relationship between functional traits and a species' specialization on specific interaction partners are not yet understood. Here we combine data on eight interaction networks between fleshy-fruited plants and frugivorous birds sampled across the tropical and subtropical Andean range. We test which combinations of morphological plant and animal traits determine trait matching between resource and consumer species in these networks. In addition, we test which of the morphological traits influence functional specialization of plant and bird species. In a meta-analysis across network-specific fourth-corner analyses, we found that plant-animal trait pairs related to size matching (fruit size-beak size) and avian foraging behavior (plant height-wing shape and crop mass-body mass) were positively related in these networks. The degree of functional specialization on specific interaction partners was positively related to crop mass in plants and to the pointedness of the wing in birds. Our findings show that morphological trait matching between fleshy-fruited plants and frugivorous birds is a general phenomenon in plant-frugivore networks across the Andes and that specific plant and bird traits can be used to approximate the degree of functional specialization. These insights into the generality of interaction rules are the base for predictions of species interactions in ecological networks, for instance in novel communities in the future, and can be applied to identify plant and animal species that fulfill specialized functional roles in ecological communities.

Although seed-dispersal networks are increasingly used to infer the functioning of ecosystems, few studies have investigated the link between the properties of these networks and the ecosystem function of seed dispersal by animals. We investigate how frugivore communities and seed dispersal change with habitat disturbance and test whether relationships between morphological traits and functional roles of seed dispersers change in response to human-induced forest edges. We recorded interaction frequencies between fleshy fruited plants and frugivorous bird species in tropical montane forests in the Bolivian Andes and recorded functional bird traits (body mass, gape width and wing tip length) associated with quantitative (seed-removal rate) and qualitative (seed-deposition pattern) components of seed-dispersal effectiveness. We found that the abundance and richness of frugivorous birds were higher at forest edges. More fruits were removed and dispersed seeds were less clustered at edges than in the interior. Additionally, functional and interaction diversity were higher at edges than in the interior, but functional and interaction evenness did not differ. Interaction strength of bird species increased with body mass, gape width and wing tip length in the forest interior, but was not related to bird morphologies at forest edges. Our study suggests that increases in functional and interaction diversity and an even distribution of interaction strength across bird morphologies lead to enhanced quantity and tentatively enhanced quality of seed dispersal. It also suggests that the effects of species traits on ecosystem functions can vary along small-scale gradients of human disturbance.