Cooperative breeding occurs in several major animal phyla, predominantly in arthropods and chordates. A number of comparative analyses have focused on understanding the evolution of cooperative breeding, yielding mixed, inconclusive, and often phyla-specific findings. We argue that much of this ambiguity results from an erroneous classification of social systems into noncooperatively and cooperatively breeding species. The shortcomings of this assumption are apparent among birds where noncooperative species constitute a heterogeneous group: some species are clearly non-family living, with offspring dispersing at or shortly after nutritional independency, whereas other species form persistent family groups through offspring delaying their dispersal substantially beyond independency. Here, we propose an objective, life history-based criterion classifying noncooperative bird species into non-family living and family living species. We demonstrate that by using the family time (the time offspring remain with its parent/s beyond independence) and body size-scaled reproductive investment, we are able to differentiate 2 groups with contrasting life histories. Our classification matches seasonal environmental variation experienced by different species: family living species postpone dispersal beyond the onset of less favorable autumn conditions. We discuss the consequences of this new social system classification for evolutionary and ecological research, potentially allowing solutions to some of the most intriguing riddles in the evolutionary history of birds-and cooperative behavior itself.

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Life history theory is an essential framework to understand the evolution of reproductive allocation. It predicts that individuals of long-lived species favour their own survival over current reproduction, leading individuals to refrain from reproducing under harsh conditions. Here we test this prediction in a long-lived bird species, the Siberian jay Perisoreus infaustus. Long-term data revealed that females rarely refrain from breeding, but lay smaller clutches in unfavourable years. Neither offspring body size, female survival nor offspring survival until the next year was influenced by annual condition, habitat quality, clutch size, female age or female phenotype. Given that many nests failed due to nest predation, the variance in the number of fledglings was higher than the variance in the number of eggs and female survival. An experimental challenge with a novel pathogen before egg laying largely replicated these patterns in two consecutive years with contrasting conditions. Challenged females refrained from breeding only in the unfavourable year, but no downstream effects were found in either year. Taken together, these findings demonstrate that condition-dependent reproductive allocation may serve to maintain female survival and offspring quality, supporting patterns found in long-lived mammals. We discuss avenues to develop life history theory concerning strategies to offset reproductive costs.

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