Distinctions among reciprocal altruism,kin selection,and cooperation and a model for the initial evolution of beneficent behavior |
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| Institution: | 1. 10527A Skyline Drive, Corning, NY 14830, USA;2. CESCO, UMR7204 Sorbonne Universités-MNHN-CNRS-UPMC, CP51, 55 rue Buffon, 75005 Paris, France;3. Centre d’Ecologie Fonctionnelle et Evolutive UMR 5175, Campus CNRS, 1919 route de Mende, 34293 Montpellier Cedex 5, France;4. Instituto de Investigacion en Recursos Cinegeticos (IREC) (CSIC-UCLM-JCCM), Ronda de Toledo s/n, 13005 Ciudad, Real, Spain;5. Institute of Avian Research, “Vogelwarte Helgoland”, An der Vogelwarte 21 D26386 Wilhelmshaven, Germany;6. Department of Animal and Plant Sciences, University of Sheffield, Sheffield, UK;7. Institute of Biological and Environmental Sciences, University of Aberdeen, Aberdeen, UK;8. Department of Ecology and Evolutionary Biology, University of California, 621 Young Drive South, Los Angeles, CA 90095-1606, USA;9. CNRS, Université Lyon 1, Université de Lyon, UMR 5558, Laboratoire Biométrie et Biologie Évolutive, 43 boulevard du 11 Novembre 1918, F-69622 Villeurbanne Cedex, France;10. Department of Biological Sciences, University of Alberta, Edmonton, AB T6G 2E9, Canada;11. Department of Vertebrate Zoology, Faculty of Biology, Lomonosov Moscow State University, Leninskie Gory 1/12, 119234 Moscow, Russia;12. UMR 5174 EDB Laboratoire Évolution et Diversité Biologique, CNRS, ENFA, Université Toulouse 3 Paul Sabatier, Toulouse 31062 Cedex 9, France;13. Department of Evolution, Ecology, and Genetics, Research School of Biology, The Australian National University, Canberra, Australia;14. Département de Biologie and Centre d’Études Nordiques, Université Laval, 1045 avenue de la Médecine, QC G1V 0A6, Canada;15. 29St Mary''s Close, Shincliffe, Durham DH1 2ND, UK;16. Centre for Ecology and Hydrology, Bush Estate, Penicuik EH26 0QB, UK;17. Behavioural Ecology, Department of Biology, Ludwig Maximilian University of Munich, Planegg-Martinsried, Germany;18. Evolutionary Ecology of Variation Research Group, Max Planck Institute for Ornithology, Seewiesen, Germany;19. Departamento de Ecología Evolutiva, Instituto de Ecología, Universidad Nacional Autónoma de México, AP 70-275, México DF 04510, México;20. Technical Resource Branch, Saskatchewan Ministry of Environment, 3211 Albert Street, Regina, SK S4S 5W6, Canada;21. Département de Biologie, Université de Sherbrooke, 2500 boulevard de L’Université, Sherbrooke, QC J1K 2R1, Canada;22. School of Biomedical Sciences and Institute of Health and Biomedical Innovation, Queensland University of Technology, Kelvin Grove, QLD 4059, Australia;23. Cornell Laboratory of Ornithology, 159 Sapsucker Woods Road, Ithaca, NY 14850, USA;24. Graham Kerr Building, University of Glasgow, Glasgow G12 8QQ, UK;25. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544-1003, USA;26. Anthropological Institute and Museum, University of Zürich, Zürich, Switzerland;27. Department of Animal Ecology, Evolutionary Biology Center, Uppsala University, Uppsala, Sweden;28. Department of Biology, Lund University, Ecology Building, 223 62 Lund, Sweden;29. Grimso Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences (SLU), SE-73091 Riddarhyttan, Sweden;30. Section of Ecology, Department of Biology, University of Turku, FI-20014 Turku, Finland;31. Department of Zoology, University of British Columbia, Vancouver, BC V6T 1Z4, Canada;32. Terrestrial Ecology Unit, Department of Biology, Ghent University, Ledeganckstraat 35, B-9000 Gent, Belgium;33. Norwegian Institute for Nature Research, PO Box 5685 Sluppen, N-7485 Trondheim, Norway;34. Department of Ecology, Swedish University of Agricultural Sciences, 75007 Uppsala, Sweden;35. Department of Integrative Biology, University of Guelph, Guelph, ON N1G 2W1, Canada;36. Faculty of Life Sciences and Engineering, University of Lleida, E-25198 Lleida, Spain;37. Ecological Genetics Research Unit, Department of Biosciences, PO Box 65 (Biocenter 3, Viikinkaari 1), FIN-00014 University of Helsinki, Finland;38. Laboratoire Ecologie, Systématique, et Evolution, Equipe Diversité, Ecologie et Evolution Microbiennes, Bâtiment 362, 91405 Orsay Cedex, France;39. Evolution and Ecology Research Centre and School of Biological, Earth, and Environmental Sciences, University of New South Wales, Sydney, Australia;40. ICT Nisbet & Company, 150 Alder Lane, North Falmouth, MA 02556, USA;41. Department of Animal Ecology, Netherlands Institute of Ecology (NIOO-KNAW), PO Box 50, 6700 AB Wageningen, The Netherlands;42. Institut Mediterrani d’Estudis Avançats IMEDEA (CSIC-UIB), Miquel Marques 21, 07190 Esporles Mallorca, Spain;43. Departamento de Ecologia Evolutiva, Estación Biológica de Doñana-CSIC, Av. Américo Vespucio s/n, 41092 Seville, Spain;44. Département des Sciences Biologiques, Université du Québec à Montréal, CP 8888-succursale Centre-Ville, Montreal, QC H3C 3P8, Canada;45. Vertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA;46. Institut Pluridisciplinaire Hubert Curien, CNRS UMR7178, 23 rue Becquerel, 67087 Strasbourg, France;47. Department of Ecology and Evolution, University of Lausanne, Lausanne, Switzerland;48. Department of Natural Resources and Environmental Science, University of Nevada Reno, Reno, NV 89512, USA;49. Department of Ecology and Natural Resource Management, Norwegian University of Life Sciences, PO Box 5003, NO-1432 Ås, Norway and Norwegian Institute for Nature Research, PO Box 5685 Sluppen, N-7485 Trondheim, Norway;50. Department of Biology, Center for Ecology, Evolution, and Behavior, University of Kentucky, Lexington, KY, USA;51. Centre for Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Cornwall Campus, Penryn TR10 9EZ, UK;52. Faculty of Arts and Sciences, Department of Environmental and Health Studies, Telemark University College, N-3800 Bø i Telemark, Norway;1. Department of Anthropology, University of Wisconsin-Madison, U.S.A.;2. ABEERU, University of South Africa, Pretoria, South Africa;3. School of Biological Sciences, University of Aberdeen, U.K.;4. Department of Psychology, University of Lethbridge, Canada;5. Department of Experimental and Applied Psychology, VU University Amsterdam, The Netherlands |
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| Abstract: | Reciprocal altruism is usually regarded as distinct from kin selection. However, because reciprocators are likely to establish long-term relations and to deliver most of their aid to other individuals genetically predisposed to reciprocation, most acts of reciprocal altruism should involve indirect increments to inclusive fitness, at least as regards alleles for reciprocation. Thus, as usually defined, reciprocal altruism is not clearly distinct from kin selection because both involve indirect increments to inclusive fitness. We propose a new definition for reciprocal altruism that makes the phenomenon distinct from kin selection and allows for reciprocation between nonrelatives in which current costs exceed future benefits returned to the reciprocal altruist. Cooperation and reciprocal altruism are often considered synonymous or different only in the timing of donating and receiving aid. We show, however, that there are other critical differences between reciprocal altruism and other forms of cooperation, most importantly, the latter often involve no clearly identifiable aid. We propose a four-category system to encompass the range of cooperative and beneficent behaviors that occur in nature (reciprocal altruism, pseudoreciprocity, simultaneous cooperation and by-product beneficence). Reciprocal altruism must involve aid that is returned to an original donor as a result of behavior that has a net cost to an original recipient. Our simplest category of cooperative/beneficent behavior, “by-product beneficence,” occurs when a selfish act also benefits another individual and requires no prior or subsequent interactions between the individuals involved. By-product beneficence may be the primitive state from which more complicated types of cooperative/beneficent behavior evolved. We show via simple models that by-product beneficence can allow for the initial increase of helping behavior in a completely unstructured population although the individuals showing such behavior pay all the costs while sharing the benefits with other individuals. Previous models that attempted to explain the initial increase of cooperative/beneficent behavior were much more complex and were based on the prisoner's dilemma, which does not accurately reflect most forms of cooperation and beneficence that occur in nature. |
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