The evolution of coloniality in birds in relation to food, habitat, predation, and life-history traits: A comparative analysis

Coloniality in birds has been intensively studied under the cost and benefit approach, but no general conclusion can be given concerning its evolutionary function. Here, we report on a comparative analysis carried out on 320 species of birds using the general method of comparative analysis for discrete variables and the contrast method to analyze the evolution of coloniality. Showing a mean of 23 convergences and 10 reversals, coloniality appears to be a rather labile trait. Colonial breeding appears strongly correlated with the absence of feeding territory, the aquatic habitat, and nest exposure to predators but was not correlated with changes in life-history traits (body mass and clutch size). The correlation of coloniality with the aquatic habitat is in fact explained by a strong correlation with the marine habitat. Unexpectedly, we found that the evolution toward a marine habitat in birds was contingent on coloniality and that coloniality evolved before the passage to a marine life. These results-along with the lack of transitions from the nonmarine to marine habitat in solitary species and the precedence of the loss of feeding territoriality on the passage to a marine life-contradict most of the hypotheses classically accepted to explain coloniality and suggest that we use a different framework to study this evolutionary enigma.