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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. [ABSTRACT FROM AUTHOR]

Animals often spend less time vigilant and more time feeding when foraging in larger groups. This group-size effect does not, however, consider if larger groups differ systematically from smaller ones: Large groups could form in different habitats than small groups or be composed of a different mix of ages or classes than small groups. We examined how habitat differences and flock size and composition explain feeding and vigilance rates in common cranes Grus grus, wintering in holm oak Quercus ilex dehesas of Spain. Flock size and composition were related to habitat type in cranes: flocks formed in areas sown with cereal crops were larger than flocks formed in set aside areas. Vigilance rate depended on habitat but decreased with increasing flock size in a similar way across all habitats. Juveniles were less vigilant than adults and showed little change in vigilance with flock size. Vigilance increased and feeding time decreased over months from November through February. Our results show that vigilance is affected by habitat but that the group size effect on vigilance is not the product of differences between habitats in group size or composition. [ABSTRACT FROM AUTHOR]