In a given area, plant-animal mutualistic interactions form complex networks that often display nestedness, a particular type of asymmetry in interactions. Simple ecological and evolutionary factors have been hypothesized to lead to nested networks. Therefore, nestedness is expected to occur in other types of mutualisms as well. We tested the above prediction with the network structure of interactions in cleaning symbiosis at three reef assemblages. In this type of interaction, shrimps and fishes forage on ectoparasites and injured tissues from the body surface of fish species. Cleaning networks show strong patterns of nestedness. In fact, after controlling for species richness, cleaning networks are even more nested than plant-animal mutualisms. Our results support the notion that mutualisms evolve to a predictable community-level structure, be it in terrestrial or marine communities.

Synopsis Fish species in many families and different trophic levels forage by following fishes and other animals. This interspecific foraging association was examined at an oceanic archipelago in the tropical West Atlantic. We recorded 27 reef fish species, two invertebrate species, and one turtle species playing the nuclear role, and 26 reef fish species acting as followers. The puddingwife wrasse following the spotted goatfish was the commonest foraging association recorded. The spotted goatfish was the nuclear fish that attracted the largest number of follower species (68% of the total number of follower species). The coney and the Noronha wrasse were the follower species that associated with the largest number of nuclear species (63 and 55% of the total number). About 20% of the reef fish species recorded in the archipelago engages in interspecific foraging associations. Substratum disturbance is a strong predictor for a fish displaying the nuclear role in the association, whereas the follower role may be predicted by carnivory. Nuclear species are diverse both in morphology and behaviour, and the nuclear role may be played either by fishes or other marine animals from invertebrates to turtles. Followers, on the other hand, comprise fishes only, which tend to display a more uniform feeding behaviour.

Mutualisms often form networks of interacting species, characterized by the existence of a central core of species that potentially drive the ecology and the evolution of the whole community. Centrality measures allow quantification of how central or peripheral a species is within a network, thus informing about the role of each species in network organization, dynamics, and stability. In the present study we addressed the question whether the structural position of species in the network (i.e. their topological importance) relates to their ecological traits. We studied interactions between cleaner and client reef fishes to identify central and peripheral species within a mutualistic network, and investigated five ecological correlates. We used three measures to estimate the level of centrality of a species for distinct structural patterns, such as the number of interactions and the structural proximity to other species. Through the use of a principal component analysis (PCA) we observed that the centrality measures were highly correlated (92.5%) in the studied network, which indicates that the same species plays a similar role for the different structural patterns. Three cleaner and ten client species had positive values of centrality, which suggests that these species are modulating ecological and evolutionary dynamics within the network. Higher centralities were related to higher abundances and feeding habits for client fishes, but not for cleaners. The high correlation between centrality measures in the present study is likely related to the nested structure of the cleaning network. The cleaner species’ set, by having central species that are not necessarily the most abundant ones, bears potentially more vulnerable points for network cohesiveness. Additionally, the present study generalizes previous findings for plant–animal mutualisms, as it shows that the structure of marine mutualisms is also related to a complex interplay between abundance and niche‐related features.

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