A New Approach to Understanding the Evolutionary Conditions Leading to Colonial Breeding: A Comparison of Colonial Breeding with Solitary Territorial Breeding and Solitary Non-Territorial Breeding

Despite many decades of study, scientists cannot answer why colonial breeding has evolved. This paper shows that the evolution of colonial breeding cannot be sufficiently explained by the evolutionary conditions proposed in previous studies. Until now, studies have compared colonial breeding with only solitary non-territorial breeding (two-state comparison) and have discussed factors that facilitate evolution of the former. Here, I propose a new approach, a three-state comparison, which compares colonial breeding with not only solitary non-territorial breeding but also solitary territorial breeding. This three-state comparison indicates that (i) merely determining the factors that make colonial breeding advantageous over non-territorial solitary breeding is not sufficient to explain the evolution of colonial breeding, (ii) the disadvantages of solitary territorial breeding can promote the evolution of colonial breeding, and (iii) the evolution of colonial breeding is far more influenced by different selection pressures than has been predicted by previous studies. Thus, the three-state comparison provides a more precise description of the conditions that lead to the evolution of colonial breeding.