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Body size has long been recognized to play a key role in shaping species interactions. For example, while small species thrive in a diversity of environments, they typically lose aggressive contests for resources with larger species. However, numerous examples exist of smaller species dominating larger species during aggressive interactions, suggesting that the evolution of traits can allow species to overcome the competitive disadvantage of small size. If these traits accumulate as lineages diverge, then the advantage of large size in interspecific aggressive interactions should decline with increased evolutionary distance. We tested this hypothesis using data on the outcomes of 23,362 aggressive interactions among 246 bird species pairs involving vultures at carcasses, hummingbirds at nectar sources, and antbirds and woodcreepers at army ant swarms. We found the advantage of large size declined as species became more evolutionarily divergent, and smaller species were more likely to dominate aggressive contests when interacting with more distantly-related species. These results appear to be caused by both the evolution of traits in smaller species that enhanced their abilities in aggressive contests, and the evolution of traits in larger species that were adaptive for other functions, but compromised their abilities to compete aggressively. Specific traits that may provide advantages to small species in aggressive interactions included well-developed leg musculature and talons, enhanced flight acceleration and maneuverability, novel fighting behaviors, and traits associated with aggression, such as testosterone and muscle development. Traits that may have hindered larger species in aggressive interactions included the evolution of morphologies for tree trunk foraging that compromised performance in aggressive contests away from trunks, and the evolution of migration. Overall, our results suggest that fundamental trade-offs, such as those associated with body size, are more likely to break down over evolutionary time, changing the rules that govern species interactions and structure ecological communities.

Here, we test this hypothesis using data from 23,362 interactions among 246 species pairs, representing three phylogenetically and ecologically distinct groups of birds that have been studied extensively with respect to aggressive contests for shared food resources. These three groups are: 1) New World vultures (Accipitriformes: Cathartidae) and Old World vultures (Accipitriformes: Accipitridae) interacting at carcasses, 2) hummingbirds (Apodiformes: Trochilidae) interacting at nectar sources, and 3) antbirds (Passeriformes: Thamnophilidae) and woodcreepers (Passeriformes: Dendrocolaptidae) competing for invertebrates and small vertebrates fleeing from army ant swarms (Hymenoptera: Formicidae) [13].

Lines connect species pairs for which we include data on aggressive interactions in this study. Dominant species (left column) were defined as species winning the majority of aggressive interactions with the subordinate species (right column).

The number of aggressive interactions over shared resources are expected to decline as species diverge over evolutionary time because more distantly-related species, on average, are ecologically more distinct [8], [28]. Small species could win more interactions with larger species when interactions are rare if they can bluff or if species are more inclined to back down when the other species is poorly known. We did not include the number of interactions among species pairs in our main linear mixed-effects model because the number of interactions was already incorporated into the dependent variable. We did, however, include the total number of interactions among species pairs in our generalized linear mixed model.

The larger species won the majority of aggressive interactions with the smaller species (i.e., was behaviorally dominant) in 201 of 246 species pairs (81.7%). Smaller species were behaviorally dominant in 43 species pairs (17.5%) involving 32 different small, dominant species, and 33 different large, subordinate species (Figure 4). Wins and losses were equal in only two species pairs. In some cases where smaller species were behaviorally dominant, differences in weight were small and perhaps insignificant. In 32 species pairs, however, the larger species was>5% heavier than the smaller, dominant species, while 25 species pairs involved larger species that were>10% heavier than the smaller, dominant species.

The advantage of large size in aggressive contests was greatest among closely-related species and declined with increasing genetic distance (Figure 5; Tables 1,2; see Figure S1 for a 3-dimensional plot that includes difference in mass, genetic distance and dominance asymmetry). Similarly, smaller species were more likely to be dominant in aggressive interactions with distantly-related species (Figure 6; Tables 3,4). For example, among interacting species in the same genus, the smaller species was dominant in only 2 of 27 species pairs (7.4%), whereas among interacting species in different taxonomic families, the smaller species was dominant in 18 of 62 species pairs (29.0%). The decline in the importance of large size in aggressive interactions with increased genetic distance was evident in each of the three independent groups and did not differ significantly between them (Tables 5,6).

The decline in the advantage of large size with increased genetic distance was not caused by variation in the number of interactions among species pairs (overall number of interactions: Tables 3,4; number of interactions within a study: Tables S3,S4). The number of interactions among species (overall, or within a study) also did not predict when small bird species would be dominant in aggressive interactions (overall number of interactions: Tables 3,4; number of interactions within a study: Tables S3,S4). 2351a5e196