Determination of temperate bird-flower interactions as entangled mutualistic and antagonistic sub-networks: characterization at the network and species levels.

Most network studies on biological interactions consider only a single interaction type. However, individual species are simultaneously positioned in various types of interactions. The ways in which different network types are merged and entangled, and the variations in network structures between different sympatric networks, require full elucidation. Incorporating interaction types and disentangling complex networks is crucial, because the integration of various network architectures has the potential to alter the stability and co-evolutionary dynamics of the whole network. To reveal how different types of interaction networks are entangled, we focused on the interaction between birds and flowers of temperate plants in Japan, where flower-feeding birds are mainly generalist passerines, acting as pollinators and predators of flowers. Using long-term monitoring data, we investigated the flower-feeding episodes of birds. We constructed the whole network (WN) between birds and plants, separating the network into mutualistic and antagonistic sub-networks (MS and AS, respectively). We investigated structural properties of the three quantified networks and species-level characteristics of the main bird species. For bird species, we evaluated dietary similarity, dietary specialization and shifts of feeding behaviour relative to plant traits. Our results indicate that WN comprises entangled MS and AS, sharing considerable proportions of bird and plant assemblages. We observed distinctive differences in the network structural properties between the two sub-networks. In comparison with AS, MS had lower numbers of bird and plant species, showed lower specialization and modularity and exhibited higher nestedness. At the species level, the Japanese white-eye acted as pollinator, while the brown-eared bulbul acted as both pollinator and predator for large numbers of flowers, based on its behavioural plasticity. Overall, the pattern of avian feeding behaviour was influenced by flower size and plant origin. Birds showed nectarivory for plants with medium-sized flowers and exotic origins. Our results highlight the complex patterns of interactions between birds and the flowers of plants in temperate regions. They also indicate that understanding the interaction type for each species pair and consideration of the behavioural plasticity of animal species are important for elucidating integrated network structures.