When David beats Goliath: the advantage of large size in interspecific aggressive contests declines over evolutionary time.

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.