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Aggression and hierarchical steepness inhibit social play in adult wolves

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In canids, play dynamics seem to be more affected by dominance hierarchy rather than cooperative social bonds. To test this hypothesis we studied a colony of grey wolves (Canis lupus lupus). We quantified the dynamics of aggression and hierarchical changes in two periods (Sample 1 and Sample 2). Sample 2 was characterized by higher level of aggressiveness and by a more strict and steep linear hierarchy. The negative correlation between rank distance and play frequency characterizing both periods and the higher play asymmetry in Sample 2 suggest that rank rules dictate play rules thus highlighting the competitive side of wolf play behaviour. The overall affiliation rates showed no variation between the two samples. Yet, play performance was modified. In Sample 2, wolves reduced playful activity, limited the number of players per session and avoided playing during high competition contexts. Our findings support the hypothesis that wolf play is modulated by dominance relationships more than by cooperative social bonds.

Affiliations: 1: aNatural History Museum, University of Pisa, Calci, Pisa, Italy ; 2: bInstitute of Cognitive Sciences and Technologies, Unit of Cognitive Primatology and Primate Center, CNR, Rome, Italy

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1. Aldis O. (1975). Play fighting. — Academic Press, New York, NY.
2. Altmann J. (1974). "Observational study of behaviour sampling methods". — Behaviour Vol 49: 227-265.
3. Altmann S.A. (1962). "A field study of the sociobiology of rhesus monkeys, Macaca mulatta ". — Ann. N.Y. Acad. Sci. Vol 102: 338-435.
4. Bauer E.B. , Smuts B.B. (2007). "Cooperation and competition during dyadic play in domestic dogs, Canis familiaris ". — Anim. Behav. Vol 73: 489-499.
5. Bekoff M. (1974). "Social play and play-soliciting by infant canids". — Am. Zool. Vol 14: 323-340.
6. Bekoff M. (1995). "Play signals as punctuation: the structure of social play in canids". — Behaviour Vol 132: 419-429.
7. Bekoff M. (2001a). "Social play behaviour: cooperation, fairness, trust, and the evolution of morality". — J. Consc. Stud. Vol 8: 81-90.
8. Bekoff M. (2001b). "The evolution of animal play, emotions, and social morality: on science, theology, spirituality, personhood, and love". — Zygon Vol 36: 615-654.
9. Bekoff M. , Wells M.C. (1986). "Social behavior and ecology of coyotes". — Adv. Stud. Behav. Vol 16: 251-338.
10. Ciani F. , Dall’Olio S. , Stanyon R. , Palagi E. (2012). "Social tolerance and adult play in macaque societies: a comparison with different human cultures". — Anim. Behav. Vol 84: 1313-1322.
11. Cohen J. , Cohen P. (1983). Applied multiple regression/correlation analysis for the behavioral sciences, 2nd edn.Erlbaum, Hillsdale, NJ.
12. Cordoni G. (2009). "Social play in captive wolves (Canis lupus): not only an immature affair". — Behaviour Vol 146: 1363-1385.
13. Cordoni G. , Palagi E. (2008). "Reconciliation in wolves (Canis lupus): new evidence for a comparative perspective". — Ethology Vol 114: 298-308.
14. Cordoni G. , Palagi E. (2011). "Ontogenetic trajectories of chimpanzee social play: similarities with humans". — PLoS ONE Vol 6: e27344.
15. de Oliveira C.R. , Ruiz-Miranda C.R. , Kleiman D.G. , Beck B.B. (2003). "Play behavior in juvenile golden lion tamarins (Callitrichidae: Primates): organization in relation to costs". — Ethology Vol 109: 593-612.
16. de Vries H. , Stevens J.M.G. , Vervaecke H. (2006). "Measuring and testing steepness of dominance hierarchies". — Anim. Behav. Vol 71: 585-592.
17. Fagen R. (1981). Animal play behavior. — Oxford University Press, New York, NY.
18. Feddersen-Petersen D.U. (1991). "The ontogeny of social play and agonistic behaviour in selected canid species". — Bonn. Zool. Beitr. Vol 42: 97-114.
19. Feddersen-Petersen D.U. (2007). "Social behaviour of dogs and related canids". — In: The behavioural biology of dogs ( Jensen P. , ed.). CAB International, Wallingford, p.  105-119.
20. Fox M.W. (1971). "Socio-ecological implications of individual differences in wolf litters: a developmental and evolutionary perspective". — Behaviour Vol 41: 298-313.
21. Fox M.W. , Andrews R.V. (1973). "Physiological and biochemical correlates of individual differences in behaviour of wolf cubs". — Behaviour Vol 46: 129-140.
22. Hare B. , Melis A.P. , Woods V. , Hastings S. , Wrangham R.W. (2007). "Tolerance allows bonobos to outperform chimpanzees on a cooperative task". — Curr. Biol. Vol 17: 619-623.
23. Hayaki H. (1985). "Social play of juvenile and adolescent chimpanzees in the Mahale Mountains National Park, Tanzania". — Primates Vol 26: 343-360.
24. Kaufman A.B. , Rosenthal R. (2009). "Can you believe my eyes? The importance of inter-observer reliability statistics in observations of animal behaviour". — Anim. Behav. Vol 78: 1487-1491.
25. Leiva D. , de Vries H. (2011). Steepness: testing steepeness of dominance hierarchies. R package version 0.2. — Available online at .
26. Loranca A. , Torrero C. , Salas M. (1999). "Development of play behavior in neonatally undernourished rats". — Physiol. Behav. Vol 66: 3-10.
27. Mech D. (1970). The wolf: the ecology and behaviour of an endangered species. — Natural History Press, Garden City, NY.
28. Mech D. (1999). "Alpha status, dominance, and division of labor in wolf packs". — Can. J. Zool. Vol 77: 1196-1203.
29. Mech L.D. , Boitani L. (2003). "Wolf social ecology". — In: Behavior, ecology, and conservation ( Mech L.D. , Boitani L. , eds). University of Chicago Press, Chicago, IL, p.  1-34.
30. Miklósi A. (1999). "The ethological analysis of imitation". — Biol. Rev. Vol 74: 347-374.
31. Mills M.G.L. (1990). Kalahari hyaena. Comparative behavioural biology of two species. — Unwin Hyman, London.
32. Mundry R. , Fischer J. (1998). "Use of statistical programs for nonparametric tests of small samples often lead to incorrect P values: examples from Animal Behaviour". — Anim. Behav. Vol 56: 256-259.
33. Packard J.M. (2003). "Wolf behaviour: reproductive, social, and intelligent". — In: Wolves: behaviour, ecology, and conservation ( Mech L.D. , Boitani L. , eds). University of Chicago Press, Chicago, IL, p.  35-65.
34. Packard J.M. (2012). "Wolf social intelligence". — In: Wolves: biology, behavior and conservation ( Maia A.P. , Crussi H.F. , eds). Nova Science Publishers, Hauppauge, NY, p.  1-48.
35. Palagi E. (2008). "Sharing the motivation to play: the use of signals in adult bonobos". — Anim. Behav. Vol 75: 887-896.
36. Palagi E. (2009). "Adult play fighting and potential role of tail signals in ringtailed lemurs (Lemur catta)". — J. Comp. Psychol. Vol 123: 1-9.
37. Palagi E. , Cordoni G. (2009). "Postconflict third-party affiliation in Canis lupus: do wolves share similarities with the great apes?" — Anim. Behav. Vol 78: 979-986.
38. Palagi E. , Cordoni G. (2012). "The right time to happen: play developmental divergence in the two Pan species". — PLoS ONE Vol 7(12): e52767.
39. Palagi E. , Cordoni G. , Borgognini Tarli S.M. (2004). "Immediate and delayed benefits of play behaviour: new evidence from chimpanzees (Pan troglodytes)". — Ethology Vol 110: 949-962.
40. Palagi E. , Paoli T. , Borgognini Tarli S.M. (2006). "Short-term benefits of play behaviour: conflict prevention in captive bonobos (Pan paniscus)". — Int. J. Primatol. Vol 27: 1257-1270.
41. Palagi E. , Antonacci D. , Cordoni G. (2007). "Fine-tuning of social play in juvenile lowland gorillas (Gorilla gorilla gorilla)". — Dev. Psychobiol. Vol 49: 433-445.
42. Palagi E. , Norscia I. , Spada G. (2014). "Relaxed open mouth as a playful signal in wild ring-tailed lemurs". — Am. J. Primatol. Vol 76: 1074-1083.
43. Palagi E. , Burghardt G.M. , Smuts B. , Cordoni G. , Dall’Olio S. , Fouts H.N. , Řeháková-Petrů M. , Siviy S.M. , Pellis S.M. (2015). "Rough-and-tumble play as a window on animal communication". — Biol. Rev. DOI:.
44. Pedersen J.M. , Glickman S.E. , Frank L.G. , Beach F.A. (1990). "Sex differences in the play behavior of immature spotted hyenas, Crocuta crocuta ". — Horm. Behav. Vol 24: 403-420.
45. Pellis S.M. (1991). "How motivationally distinct is play? A preliminary case study". — Anim. Behav. Vol 42: 851-853.
46. Pellis S.M. , Iwaniuk A.N. (2000). "Comparative analyses of the role of postnatal development on the expression of play fighting". — Dev. Psychobiol. Vol 36: 136-147.<136::AID-DEV5>3.0.CO;2-V
47. Pellis S.M. , Pellis V.C. (2009). The playful brain: venturing to the limits of neuroscience. — Oneworld Publications, Oxford.
48. Pellis S.M. , Pellis V.C. , Reinhart C.J. (2010). "The evolution of social play". — In: Formative experiences: the interaction of caregiving, culture, and developmental psychobiology ( Worthman C. , Plotsky P. , Schechter D. , Cummings C. , eds). Cambridge University Press, Cambridge, p.  404-431.
49. Power T.G. (2000). Play and exploration in children and animals. — Lawrence Erlbaum Associates, Mahwah, NJ.
50. Siegel S. , Castellan N.J.J. (1988). Nonparametric statistics for the behavioral sciences. — McGraw Hill, New York, NY.
51. Smuts B. (2014). "Social behaviour among companion dogs with an emphasis on play". — In: The social dog: behavior and cognition ( Kaminski J. , Marshall-Pescini S. , eds). Academic Press, San Diego, CA, p.  105-130.
52. Špinka M. , Newberry R.C. , Bekoff M. (2001). "Mammalian play: training for the unexpected". — Q. Rev. Biol. Vol 76: 141-167.
53. Stahler D.R. , Smith D.W. , Landis R. (2002). "The acceptance of a new breeding male into a wild wolf pack". — Can. J. Zool. Vol 80: 360-365.
54. Ward C. , Bauer E.B. , Smuts B.B. (2008). "Partner preferences and asymmetries in social play among domestic dog, Canis lupus familiaris, littermates". — Anim. Behav. Vol 76: 1187-1199.
55. Ward C. , Trisko R. , Smuts B.B. (2009). "Third-party interventions in dyadic play between littermates of domestic dogs, Canis lupus familiaris ". — Anim. Behav. Vol 78: 1153-1160.
56. Wilson E.O. (2000). Sociobiology — the new synthesis. — The Belknap Press of Harvard University Press, Cambridge, MA.
57. Wrangham R.W. (1993). "The evolution of sexuality in chimpanzees and bonobos". — Hum. Nat. Vol 4: 47-79.
58. Yachmennikova A.A. (2012). "Agonistic behavior inter-connections in hidden patterns in wolf pups group during the juvenile hierarchy stabilizing period". — In: Wolves: biology, behavior and conservation ( Maya A.P. , Crussi H.F. , eds). Nova Science Publishers, Hauppauge, NY, p.  109-132.
59. Zimen E. (1975). "Social dynamics of the wolf pack". — In: The wild canids: their systematics, behavioural ecology and evolution ( Fox M.W. , ed.). Van Nostrand Reinhold, New York, NY, p.  336-368.
60. Zimen E. (1976). "On the regulation of pack size in wolves". — Z. Tierpsychol. Vol 40: 300-341.
61. Zimen E. (1981). The wolf. — Souvenir Press, London.

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