, Volume 24, Issue 2, pp 125–135 | Cite as

Animal Play and the Evolution of Morality: An Ethological Approach

  • Colin AllenEmail author
  • Marc Bekoff


In this paper we argue that there is much to learn about “wild justice” and the evolutionary origins of morality – behaving fairly – by studying social play behavior in group-living mammals. Because of its relatively wide distribution among the mammals, ethological investigation of play, informed by interdisciplinary cooperation, can provide a comparative perspective on the evolution of ethical behavior that is broader than is provided by the usual focus on primate sociality. Careful analysis of social play reveals rules of engagement that guide animals in their social encounters. Because of its significance in development, play may provide a foundation of fairness for other forms of cooperation that are advantageous to group living. Questions about the evolutionary roots of cooperation, fairness, trust, forgiveness, and morality are best answered by attention to the details of what animals do when they engage in social play – how they negotiate agreements to cooperate, to forgive, to behave fairly, and to develop trust. We consider questions such as why play fairly? Why did play evolve as it has? Does “being fair” mean being more fit? Do individual variations in play influence an individual’s reproductive fitness? Can we use information about the foundations of moral behavior in animals to help us understand ourselves? We conclude that there is likely to be strong selection for cooperative fair play because there are mutual benefits when individuals adopt this strategy and group stability may also be fostered. Numerous mechanisms have evolved to facilitate the initiation and maintenance of social play, to keep others engaged, so that agreeing to play fairly and the resulting benefits of doing so can be readily achieved.


Moral Behavior Nonhuman Animal Social Play Evolutionarily Stable Strategy Play Signal 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Adams, E.R., Burnett, G.W. 1991‘Scientific Vocabulary Divergence Among Female Primatologists Working in East Africa’Social Studies of Science21547560CrossRefGoogle Scholar
  2. Aldis, O. 1975Play FightingAcademic PressNew York, NYGoogle Scholar
  3. Allen, C., Bekoff, M. 1997Species of Mind: The Philosophy and Biology of Cognitive EthologyMIT PressCambridge, MAGoogle Scholar
  4. Axelrod, R. 1984The Evolution of CooperationBasic BooksNew York, NYGoogle Scholar
  5. Bateson, G. 1955‘A Theory of Play and Fantasy’Psychiatric Research Reports23951Google Scholar
  6. Bauer E. B., Smuts, B. B. (2002). ‘Role Reversal and Self-handicapping During Playfighting in Domestic Dogs, Canis familiaris’. Paper presented at the meetings of the Animal Behavior Society, Indiana University: IndianaGoogle Scholar
  7. Bekoff, M. 1972‘The Development of Social Interaction, Play, and Metacommunication in Mammals: An Ethological Perspective’Quarterly Review of Biology47412434CrossRefGoogle Scholar
  8. Bekoff, M. 1975‘The Communication of Play Intention: Are Play Signals Functional?’Semiotica15231239CrossRefGoogle Scholar
  9. Bekoff, M. 1977a‘Social Communication in Canids: Evidence for the Evolution of a Stereotyped Mammalian Display’Science19710971099CrossRefGoogle Scholar
  10. Bekoff, M. 1977b‘Mammalian Dispersal and the Ontogeny of Individual Behavioral Phenotypes’American Naturalist111715732CrossRefGoogle Scholar
  11. Bekoff, M. 1995‘Play Signals as Punctuation: The Structure of Social Play in Canids’Behaviour132419429CrossRefGoogle Scholar
  12. Bekoff, M. 2002Minding Animals: Awareness, Emotions, and HeartOxford University PressNew York, NYGoogle Scholar
  13. Bekoff, M. 2004‘Wild Justice and Fair Play: Cooperation, Forgiveness, and Morality in Animals’Biology and Philosophy19489520CrossRefGoogle Scholar
  14. Bekoff, M., Allen, C. 1992‘Intentional Icons: Towards an Evolutionary Cognitive Ethology’Ethology91116CrossRefGoogle Scholar
  15. Bekoff, M., Allen, C. 1998‘Intentional Communication and Social Play: How and Why Animals Negotiate and Agree to Play’Bekoff, M.Byers, J.A. eds. Animal Play: Evolutionary, Comparative, and Ecological Perspectives.Cambridge University PressCambridge and New York97114CrossRefGoogle Scholar
  16. Bekoff, M.Allen, C.Burghardt, G.M. eds. 2002The Cognitive AnimalMIT PressCambridge, MAGoogle Scholar
  17. Bekoff, M.Byers, J.A. eds. 1998Animal Play: Evolutionary, Comparative, and Ecological ApproachesCambridge University PressNew York, NYGoogle Scholar
  18. Bekoff, M., Wells, M.C. 1986‘Social Behavior and Ecology of Coyotes’Advances in the Study of Behavior16251338CrossRefGoogle Scholar
  19. Bernstein I.S. (2000). ‘The Law of Parsimony Prevails: Missing Premises Allow Any Conclusion’. in L. D. Katz (ed.). op. cit., pp. 31–34Google Scholar
  20. Boehm, C. 1999Hierarchy in the Forest: The Evolution of Egalitarian BehaviorHarvard University PressCambridge, MAGoogle Scholar
  21. Brosnan, S.F., Waal, F.B.M. 2003‘Monkeys Reject Unequal Pay’Nature425297299CrossRefGoogle Scholar
  22. Brosnan, S.F., Waal, F.B.M. 2004‘Reply to “Fair Refusal by Capuchin Monkeys”’Nature428140CrossRefGoogle Scholar
  23. Burghardt, G.M. 1997‘Amending Tinbergen: A Fifth Aim for Ethology’Mitchell, R.W.Thompson, N.S.Miles, H.L. eds. Anthropomorphism, Anecdotes and Animals.SUNY PressAlbany, NY354476Google Scholar
  24. Burghardt, G.M. 2005The Genesis of PlayMIT PressCambridge, MAGoogle Scholar
  25. Dennett, D.C. 1983‘Intentional Systems in Cognitive Ethology: The “Panglossian Paradigm” Defended’Behavioral and Brain Sciences6343390CrossRefGoogle Scholar
  26. Dugatkin, L.A. 1997Cooperation Among Animals: An Evolutionary PerspectiveOxford University PressNew York, NYGoogle Scholar
  27. Dugatkin, L.A., Bekoff, M. 2003‘Play and the Evolution of Fairness: A Game Theory Model’Behavioural Processes60209214CrossRefGoogle Scholar
  28. Fagen, R. 1993‘Primate Juveniles and Primate Play’Pereira, M.E.Fairbanks, L.A. eds. Juvenile Primates: Life History, Development, and Behavior.Oxford University PressNew York, NY183196Google Scholar
  29. Flack J.C., de Waal F. (2000). ‘Any Animal Whatever: Darwinian Building Blocks of Morality in Monkeys and Apes’. In: Katz L.D. (ed). op. cit., pp. 1–29Google Scholar
  30. Flack, J.C., Jeannotte, L.A., Waal, F. 2004‘Play Signaling and the Perception of Social Rules by Juvenile Chimpanzees (Pan Troglodytes)’Journal of Comparative Psychology118149159CrossRefGoogle Scholar
  31. Greene, J.D., Sommerville, R.B., Nystrom, L.E., Darley, J.M., Cohen, J.D. 2001‘An fMRI Investigation of Emotional Engagement in Moral Judgment’Science29321052108CrossRefGoogle Scholar
  32. Griffin, D.R. 1978‘Prospects for a Cognitive Ethology’Behavioral and Brain Sciences4527538CrossRefGoogle Scholar
  33. Gruen, L. (2002). ‘The Morals of Animal Minds’. in M. Bekoff et al. (eds.). op. cit., pp. 437–442Google Scholar
  34. Hauser, M. 2000Wild MindsHenry HoltNew York, NYGoogle Scholar
  35. Hauser, M.D., Marler, P. 1993‘Food Associated Calls in Rhesus Macaques (Macaca mulatta). II. Costs and Benefits of Call Production and Suppression’Behavioral Ecology4206212CrossRefGoogle Scholar
  36. Horowitz, A.C. 2002‘The Behaviors of Theories of Mind, and a Case Study of Dogs at Play’. Ph.D. dissertationUniversity of CaliforniaSan DiegoGoogle Scholar
  37. Jamieson, D., Bekoff, M. 1993‘On Aims and Methods of Cognitive Ethology’Philosophy of Science Association2110124Google Scholar
  38. Katz, L. D. (ed.): (2000). Evolutionary Origins of Morality: Interdisciplinary Perspectives. Exeter, UK: Imprint Academic (reprint of Journal of Consciousness Studies 7)Google Scholar
  39. Markowitz, H. 1982Behavioral Enrichment in the ZooVan Reinhold CompanyNew York, NYGoogle Scholar
  40. Millikan, R.G. 1984Language, Thought, and Other Biological CategoriesMIT PressCambridge, MAGoogle Scholar
  41. Panksepp, J. 1998Affective NeuroscienceOxford University PressNew York, NYGoogle Scholar
  42. Panksepp, J. 2000‘The Rat Will Play’Bekoff, M. eds. The Smile of a Dolphin: Remarkable Accounts of Animal Emotions.Random House/Discovery BooksNew York146147Google Scholar
  43. Pellis S. (2002). ‘Keeping in Touch: Play Fighting and Social Knowledge’. In: Bekoff M., Allen C., Burghardt G.M. (ed). op. cit., pp. 421–427Google Scholar
  44. Power, T.G. 2000Play and Exploration in Children and AnimalsLawrence Erlbaum AssociatesHillsdalem, NJGoogle Scholar
  45. Ridley, M. 1996The Origins of Virtue: Human Instincts and the Evolution of CooperationVikingNew York, NYGoogle Scholar
  46. Scott, J.P., Fuller, J.L. 1965Genetics and the Social Behavior of the DogsUniversity of Chicago PressChicago, ILGoogle Scholar
  47. Singer, P. 1999A Darwinian Left: Politics, Evolution, and CooperationYale University PressNew HavenGoogle Scholar
  48. Siviy, S. 1998‘Neurobiological Substrates of Play Behavior: Glimpses into the Structure and Function of Mammalian Playfulness’Bekoff, M.Byers, J.A. eds. Animal Play: Evolutionary, Comparative, and Ecological Perspectives.Cambridge University PressNew York, NY221242CrossRefGoogle Scholar
  49. Skyrms B. (2000). ‘Game Theory, Rationality and Evolution of the Social Contract’. in L. D. Katz (ed.). op. cit., pp. 269–284Google Scholar
  50. Smuts, B.B. 2001‘Encounters with Animal Minds’Journal of Consciousness Studies8293309Google Scholar
  51. Sober, E., Wilson, D.S. 1998Unto Others: The Evolution and Psychology of Unselfish BehaviorHarvard University PressCambridge, MAGoogle Scholar
  52. Sober, E., Wilson, D. S. (2000). ‘Summary of “Unto Others: The Evolution and Psychology of Unselfish Behavior”’. in L. D. Katz (ed.). op. cit., pp. 185–206Google Scholar
  53. Solomon, R. 1995A Passion for Justice: Emotions and the Origins of the Social ContractRowman & Littlefield PublishersLanham, Maryland MDGoogle Scholar
  54. Spinka, M., Newberry, R.C., Bekoff, M. 2001‘Mammalian Play: Training for the Unexpected’Quarterly Review of Biology76141168CrossRefGoogle Scholar
  55. Sutton-Smith, B. 1997The Ambiguity of PlayCambridge University PressNew York, NYGoogle Scholar
  56. Tinbergen, N. 1963‘On Aims and Methods of Ethology’Zeitschrift für Tierpsychologie20410433CrossRefGoogle Scholar
  57. Waal, F. 1996Good-Natured: The Origins of Right and Wrong in Humans and Other AnimalsHarvard University PressCambridge, MAGoogle Scholar
  58. Watson, D.M., Croft, D.B. 1996‘Age-related Differences in Playfighting Strategies of Captive Male Red-necked Wallabies (Macropus Rufogriseus Banksianus)’Ethology10233346Google Scholar
  59. Wilson, D.S. 2002Darwin’s Cathedral: Evolution, Religion, and the Nature of SocietyUniversity of Chicago PressChicago, ILGoogle Scholar

Copyright information

© Springer 2005

Authors and Affiliations

  1. 1.Department of History and Philosophy of Science and Program in Cognitive ScienceIndiana UniversityBloomingtonUSA
  2. 2.Department of Ecology and Evolutionary BiologyUniversity of ColoradoBoulderUSA

Personalised recommendations