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]
^Lukas, D.; Clutton-Brock, T. (2012). "Life histories and the evolution of cooperative breeding in mammals". Proceedings of the Royal Society B: Biological Sciences. 279 (1744): 4065–70. doi:10.1098/rspb.2012.1433. PMC 3427589. PMID 22874752.
^Dickinson, Janis L.; Koenig, Walter D.; Pitelka, Frank A. (1996-06-20). "Fitness consequences of helping behavior in the western bluebird". Behavioral Ecology. 7 (2): 168–177. doi:10.1093/beheco/7.2.168. ISSN 1045-2249.
^Haydock, J.; Koenig, W. D.; Stanback, M. T. (2001-06-01). "Shared parentage and incest avoidance in the cooperatively breeding acorn woodpecker". Molecular Ecology. 10 (6): 1515–1525. Bibcode:2001MolEc..10.1515H. doi:10.1046/j.1365-294X.2001.01286.x. ISSN 1365-294X. PMID 11412372. S2CID 21904045.
^Richardson, David S.; Burke, Terry; Komdeur, Jan; Dunn, P. (2002-11-01). "Direct benefits and the evolution of female-biased cooperative breeding in seychelles warblers". Evolution. 56 (11): 2313–2321. doi:10.1554/0014-3820(2002)056[2313:DBATEO]2.0.CO;2. ISSN 0014-3820. PMID 12487360. S2CID 198157808.
^Cockburn, Andrew (1998-01-01). "Evolution of Helping Behavior in Cooperatively Breeding Birds". Annual Review of Ecology and Systematics. 29: 141–177. doi:10.1146/annurev.ecolsys.29.1.141. JSTOR 221705.
^Jennions, M (1994-01-01). "Cooperative breeding in mammals". Trends in Ecology & Evolution. 9 (3): 89–93. doi:10.1016/0169-5347(94)90202-x. PMID 21236784.
^Wong, Marian; Balshine, Sigal (2011-05-01). "The evolution of cooperative breeding in the African cichlid fish, Neolamprologus pulcher". Biological Reviews. 86 (2): 511–530. doi:10.1111/j.1469-185X.2010.00158.x. ISSN 1469-185X. PMID 20849492. S2CID 39910620.
^Bourke, Andrew F. G.; Heinze, Jurgen (1994-09-30). "The Ecology of Communal Breeding: The Case of Multiple-Queen Leptothoracine Ants". Philosophical Transactions of the Royal Society of London B: Biological Sciences. 345 (1314): 359–372. Bibcode:1994RSPTB.345..359B. doi:10.1098/rstb.1994.0115. ISSN 0962-8436.
^Nicholas B. Davies, John R. Krebs, S. A. W. An Introduction to Behavioural Ecology.pdf. 522 (2012).
^West, Stuart (2007). "Evolutionary explanations for cooperation". Current Biology. 17 (16): R661–R672. Bibcode:2007CBio...17.R661W. doi:10.1016/j.cub.2007.06.004. PMID 17714660. S2CID 14869430.
^ abGerlach, Gabriele; Bartmann, Susann (2002-05-01). "Reproductive skew, costs, and benefits of cooperative breeding in female wood mice (Apodemus sylvaticus)". Behavioral Ecology. 13 (3): 408–418. doi:10.1093/beheco/13.3.408.
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