Downscaling pollen-transport networks to the level of individuals.

Most plant-pollinator network studies are conducted at species level, whereas little is known about network patterns at the individual level. In fact, nodes in traditional species-based interaction networks are aggregates of individuals establishing the actual links observed in nature. Thus, emergent properties of interaction networks might be the result of mechanisms acting at the individual level. Pollen loads carried by insect flower visitors from two mountain communities were studied to construct pollen-transport networks. For the first time, these community-wide pollen-transport networks were downscaled from species-species (sp-sp) to individuals-species (i-sp) in order to explore specialization, network patterns and niche variation at both interacting levels. We used a null model approach to account for network size differences inherent to the downscaling process. Specifically, our objectives were (i) to investigate whether network structure changes with downscaling, (ii) to evaluate the incidence and magnitude of individual specialization in pollen use and (iii) to identify potential ecological factors influencing the observed degree of individual specialization. Network downscaling revealed a high specialization of pollinator individuals, which was masked and unexplored in sp-sp networks. The average number of interactions per node, connectance, interaction diversity and degree of nestedness decreased in i-sp networks, because generalized pollinator species were composed of specialized and idiosyncratic conspecific individuals. An analysis with 21 pollinator species representative of two communities showed that mean individual pollen resource niche was only c. 46% of the total species niche. The degree of individual specialization was associated with inter- and intraspecific overlap in pollen use, and it was higher for abundant than for rare species. Such niche heterogeneity depends on individual differences in foraging behaviour and likely has implications for community dynamics and species stability. Our findings highlight the importance of taking interindividual variation into account when studying higher-order structures such as interaction networks. We argue that exploring individual-based networks will improve our understanding of species-based networks and will enhance the link between network analysis, foraging theory and evolutionary biology.

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