Cooperative breeding

Cooperative breeding is a social system characterized by alloparental care: offspring receive care not only from their parents, but also from additional group members, often called helpers.^[1] Cooperative breeding encompasses a wide variety of group structures, from a breeding pair with helpers that are offspring from a previous season,^[2] to groups with multiple breeding males and females (polygynandry) and helpers that are the adult offspring of some but not all of the breeders in the group,^[3] to groups in which helpers sometimes achieve co-breeding status by producing their own offspring as part of the group's brood.^[4] Cooperative breeding occurs across taxonomic groups including birds,^[5] mammals,^[6] fish,^[7] and insects.^[8]

Costs for helpers include a fitness reduction, increased territory defense, offspring guarding and an increased cost of growth. Benefits for helpers include a reduced chance of predation, increased foraging time, territory inheritance, increased environmental conditions and an inclusive fitness. Inclusive fitness is the sum of all direct and indirect fitness, where direct fitness is defined as the amount of fitness gained through producing offspring. Indirect fitness is defined as the amount of fitness gained through aiding the offspring of related individuals, that is, relatives are able to indirectly pass on their genes through increasing the fitness of related offspring.^[9] This is also called kin selection.^[10]

For the breeding pair, costs include increased mate guarding and suppression of subordinate mating. Breeders receive benefits as reductions in offspring care and territory maintenance. Their primary benefit is an increased reproductive rate and survival.

Cooperative breeding causes the reproductive success of all sexually mature adults to be skewed towards one mating pair. This means the reproductive fitness of the group is held within a select few breeding members and helpers have little to no reproductive fitness.^[11] With this system, breeders gain an increased reproductive fitness, while helpers gain an increased inclusive fitness.^[11]