Play fighting in Visayan warty pigs (Sus cebifrons): insights on restraint and reciprocity in the maintenance of play

in Behaviour
No Access
Get Access to Full Text
Rent on DeepDyve

Have an Access Token?

Enter your access token to activate and access content online.

Please login and go to your personal user account to enter your access token.


Have Institutional Access?

Access content through your institution. Any other coaching guidance?


Restraint is thought to be essential to enable the reciprocity needed for play fighting to remain playful. Descriptions of playing in pigs suggest that they do not exhibit restraint. Analysis of videotaped sequences of play fighting in captive family groups of warty pigs was used to test three hypotheses about restraint and reciprocity. Hypothesis 1 asserts that the lack of restraint arises from neither participant handicapping its actions in favour of its opponent: this was supported. Hypothesis 2 asserts that the winner of a contest will show restraint by not prosecuting further attack: this was not supported. However, the winner did refrain from attacking if the loser signalled submission. Hypothesis 3 asserts that restraint by the winner will allow reciprocal attacks by the loser — this was supported. The dissociation of restraint and reciprocity evident in the pigs offers some new insights into the evolution of play fighting.

Play fighting in Visayan warty pigs (Sus cebifrons): insights on restraint and reciprocity in the maintenance of play

in Behaviour



AldisO. (1975). Play fighting. — Academic PressNew York, NY.

AltmannS.A. (1962). Social behavior of anthropoid primates: analysis of recent concepts. — In: Roots of behavior ( BlissE.L. ed.). Harper-CollinsNew York, NY p.  277-285.

BarretteC. (1986). Fighting behavior of wild Sus scrofa. — J. Mammal. 67: 177-179.

BatesonG. (1955). A theory of play and fantasy. — Psychol. Res. Rep. 2: 39-51.

BauerE.B.SmutsB.B. (2007). Cooperation and competition during dyadic play in domestic dogs, Canis familiaris. — Anim. Behav. 73: 489-499.

BeebeB. (2006). Co-constructing mother-infant distress in face-to-face interactions: contributions of microanalysis. — Infant Observ. 9: 151-164.

BekoffM. (1975). The communication of play intention: are play signals functional?Semiotica 15: 231-239.

BekoffM. (2002). Minding animals. — Oxford University PressNew York, NY.

BibenM. (1998). Squirrel monkey play fighting: making the case for a cognitive training function for play. — In: Animal play: evolutionary comparative and ecological perspectives ( BekoffM.ByersJ.A. eds). Cambridge University PressCambridge p.  161-162.

BlackshawJ.K.SwainA.J.BlackshawA.W.ThomasF.J.M.GilliesK.J. (1997). The development of playful behaviour in piglets from birth to weaning in three farrowing enviroments. — Appl. Anim. Behav. Sci. 55: 37-49.

BlanchardR.J.BlanchardD.C. (1994). Environmental targets and sensorimotor systems in aggression and defense. — In: Ethology and psychopharmacology ( CooperS.J.HendrieC.A. eds). WileyNew York, NY p.  133-157.

BlumsteinD.T.ChungL.K.SmithJ.E. (2013). Early play may predict later dominance relationships in yellow-bellied marmots (Marmota flaviventris). — Proc. Roy. Soc. Lond. B: Biol. Sci. 280: 20130485.

BradburyJ.W.VehrencampS.L. (2011). Principles of animal communication2nd edn.Sinauer AssociatesSunderland, MA.

BrownS.M.KlaffenböckM.NevisonI.M.LawrenceA.B. (2015). Evidence for litter differences in play behaviour in pre-weaned pigs. — Appl. Anim. Behav. Sci. 172: 17-25.

BurghardtG.M. (2005). The genesis of animal play: testing the limits. — MIT PressCambridge, MA.

BurghardtG.M.BurghardtL.S. (1972). Notes on the behavioral development of two female black bear cubs: the first eight months. — In: Bears: their biology and managementVol. 2 ( HerreroS.M. ed.). IUCNMorges p.  207-220.

ByersJ.A.WalkerC. (1995). Refining the motor training hypothesis for the evolution of play. — Am. Nat. 146: 25-41.

CamerlinkI.TurnerS.P. (2013). The pig’s nose and its role in dominance relationships and harmful behaviour. — Appl. Anim. Behav. Sci. 145: 84-91.

ChaloupkováH.IllmannG.BartošL.ŠpinkaM. (2007). The effect of pre-weaning housing on the play and agonistic behaviour of domestic pigs. — Appl. Anim. Behav. Sci. 103: 25-34.

ChanceM.R.A. (1962). An interpretation of some agonistic postures: the role of “cut-off” acts and postures. — Symp. Zool. Soc. Lond. 8: 71-89.

CordoniG.PalagiE. (2011). Ontogenetic trajectories of chimpanzee social play: similarities with humans. — PLoS ONE 6: e27344.

DapportoL.TurillazziS.PalagiE. (2006). Dominance interactions in young adult paper wasp (Polistes dominulus) foundresses: a playlike behavior?J. Comp. Psychol. 120: 394-400.

D’EathR.B.PickupH.E. (2002). Behaviour of young growing pigs in a resident-intruder test designed to measure aggressiveness. — Aggress. Behav. 28: 401-415.

DonaldsonT.M.NewberryR.C.ŠpinkaM.CloutierS. (2002). Effects of early experience on play behaviour of piglets after weaning. — Appl. Anim. Behav. Sci. 79: 221-231.

DugatkinL.A.BekoffM. (2003). Play and the evolution of fairness: a game theory model. — Behav. Process. 60: 209-214.

EshkolN.WachmannA. (1958). Movement notation. — Weidenfeld & NicholsonLondon.

EstesR.D. (1993). The safari companion. A guide to watching African mammals. — Chelsea GreenPost Mills, VT.

FagenR. (1981). Animal play behavior. — Oxford University PressNew York, NY.

FoxM.W. (1969). The anatomy of aggression and its ritualization in Canidae: a developmental and comparative study. — Behaviour. 35: 242-258.

FrädrichH. (1974). A comparison of behaviour in the Suidae. — In: The behaviour of ungulates and its relation to management ( GeistV.WaltherF. eds). IUCNMorges p.  133-143.

FryD.P. (2005). Rough and tumble social play in humans. — In: The nature of play ( PellegriniA.D.SmithP.K. eds). Guilford PressNew York, NY p.  54-85.

GeistV. (1978). On weapons, combat and ecology. — In: Advances in the study of communication and affectVol. 4 Aggression dominance and individual spacing ( KramesL.PlinerP.AllowayT. eds). Plenum PressNew York, NY p.  1-30.

GolaniI. (1976). Homeostatic motor processes in mammalian interactions: a choreography of display. — In: Perspectives in ethologyVol. 2 ( BatesonP.P.G.KlopferP.H. eds). Plenum PressNew York, NY p.  69-134.

HimmlerB.T.BellH.C.HorwoodL.HarkerA.KolbB.PellisS.M. (2014). The role of the medial prefrontal cortex in regulating inter-animal coordination of movements. — Behav. Neurosci. 128: 603-613.

HimmlerS.M.HimmlerB.T.PellisV.C.PellisS.M. (2016). Play, variation in play and the development of socially competent rats. — Behaviourin press. (BEH 3307)

HorbackK. (2014). Nosing around: play in pigs. — Anim. Behav. Cog. 1: 186-196.

JohnsonR.E. (1985). Communication. — In: The hamster ( SiegelH.I. ed.). Plenum PressNew York, NY p.  121-154.

LeRescheL.A. (1976). Dyadic play in hamadryas baboons. — Behaviour 57: 290-305.

LeyhausenP. (1979). Cat behavior. The predatory and social behavior of domestic and wild cats. — Garland STPM PressNew York, NY.

McGloneJ.J. (1985). A quantitative ethogram of aggressive and submissive behaviors in recently regrouped pigs. — J. Anim. Sci. 61: 559-565.

MillerS. (1973). Ends, means and galumphing: some leitmotifs of play. — Am. Anthropol. 75: 87-98.

MoynihanM. (1955). Some aspects of reproductive behavior in the black-headed gull (Larus ridibundus ridibundus L.), and related species. — Behaviour Suppl. 4: 1-201.

NewberryR.C.Wood-GushD.G.M.HallJ.W. (1988). Playful behaviour of piglets. — Behav. Process. 17: 205-216.

PalagiE. (2008). Sharing the motivation to play: the role of signals in adult bonobos. — Anim. Behav. 75: 887-896.

PellisS.M. (1988). Agonistic versus amicable targets of attack and defense: consequences for the origin, function, and descriptive classification of play-fighting. — Aggress. Behav. 14: 85-104.

PellisS.M. (1997). Targets and tactics: the analysis of moment-to-moment decision making in animal combat. — Aggress. Behav. 23: 107-129.

PellisS.M. (2002). Keeping in touch: play fighting and social knowledge. — In: The cognitive animal: empirical and theoretical perspectives on animal cognition ( BekoffM.AllenC.BurghardtG.M. eds). MIT PressCambridge, MA p.  421-427.

PellisS.M.PellisV.C. (1996). On knowing it’s only play: the role of play signals in play fighting. — Aggress. Violent Behav. 1: 249-268.

PellisS.M.PellisV.C. (1998). The structure-function interface in the analysis of play fighting. — In: Play behavior: comparative evolutionary and ecological aspects ( BekoffM.ByersJ.A. eds). Cambridge University PressCambridge p.  115-140.

PellisS.M.PellisV.C.HimmlerB.T. (2014). How play makes for a more adaptable brain: a comparative and neural perspective. — Am. J. Play 7: 73-98.

PellisS.M.PellisV.C.ReinhartC.J. (2010). The evolution of social play. — In: Formative experiences: the interaction of caregiving culture and developmental psychobiology ( WorthmanC.PlotskyP.SchechterD.CummingsC. eds). Cambridge University PressCambridge p.  404-431.

PellisS.M.BlundellM.A.BellH.C.PellisV.C.KrakauerA.H.PatricelliG.L. (2013). Drawn into the vortex: the facing-past encounter and combat in lekking male greater sage-grouse (Centrocercus urophasianus). — Behaviour 150: 1567-1599.

PerrettD.I.MistlinA.J. (1990). Perception of facial characteristics by monkeys. — In: Comparative perception — complex signals ( StebbinsW.C.BerkleyM.A. eds). WileyNew York, NY p.  187-215.

RushenJ.PajorE. (1987). Offence and defence in fighting between young pigs (Sus scrofa). — Aggress. Behav. 13: 329-346.

SharpeL.L. (2005). Play fighting does not affect subsequent fighting success in wild meerkats. — Anim. Behav. 69: 1023-1029.

SiegelS.CastellanN.J.J. (1988). Nonparametric statistics for the behavioral sciences. — McGraw-HillNew York, NY.

ŠilerováJ.ŠpinkaM.ŠárováR.AlgersB. (2010). Playing and fighting by piglets around weaning on farms, employing individual or group housing of lactating sows. — Appl. Anim. Behav. Sci. 124: 83-89.

SiviyS.M.PankseppJ. (2011). In search of the neurobiological substrates for social playfulness in mammalian brains. — Neurosci. BioBehav. Rev. 35: 1821-1830.

SmithP.K. (1982). Does play matter? Functional and evolutionary play. — Behav. Brain Sci. 5: 139-184.

SmithP.K. (1997). Play fighting and real fighting. Perspectives on their relationship. — In: New aspects of human ethology ( SchmittA.AtzwangerK.GrammarK.SchäferK. eds). Plenum PressNew York, NY p.  47-64.

SmutsB. (2014). Social behavior among companion dogs with an emphasis on play. — In: The social dog. Behavior and cognition ( KaminskiJ.Marshall-PesciniS. eds). ElsevierLondon p.  105-130.

SuomiS.J. (2005). Genetic and environmental factors influencing the expression of impulsive aggression and serotonergic functioning in rhesus monkeys. — In: Developmental origins of aggression ( TremblayR.E.HartupW.W.ArcherJ. eds). Guilford PressNew York, NY p.  63-82.

SymonsD. (1978). Play and aggression. A study of rhesus monkeys. — Columbia University PressNew York, NY.

ThompsonK.V. (1998). Self assessment in juvenile play. — In: Animal play: evolutionary comparative and ecological perspectives ( BekoffM.ByersJ.A. eds). Cambridge University PressCambridge p.  183-204.

VanderschurenL.J.M.J. (2010). How the brain makes play fun. — Am. J. Play 2: 315-337.

VanderschurenL.J.M.J.TrezzaV. (2014). What the laboratory rat has taught us about social play behavior: role in behavioral development and neural mechanisms. — Curr. Top. Behav. Neurosci. 16: 189-212.

WaltherF.R. (1984). Communication and expression in hoofed mammals. — Indiana University PressBloomington, IN.

WilmerA.H. (1991). Behavioral deficiencies of aggressive 8–9 year-old boys: an observational study. — Aggress. Behav. 17: 135-154.


  • View in gallery

    A photograph of a piglet, with dark lines superimposed on the underlying leg and body segments that were notated. The white lines with arrowheads on either end show the overall axis between the piglet’s feet and its shoulder and pelvis, respectively. Shifting the body axis forward (i.e., when pushing against the partner) would orient the shoulder and pelvis in front of where its feet press on the ground, and in this way, the upper arrowheads would point forward. Conversely, in shifting the body axis backwards, the piglet’s shoulder and pelvis would be directly above or a little behind where its feet press on the ground, and in this way, the upper arrowheads would point backwards. This figure is published in colour in the online edition of this journal, which can be accessed via

  • View in gallery

    The percentage distribution of the three main categories by which play fights were terminated is shown for both sample 1 (N=60) and sample 2 (N=40).


Content Metrics

Content Metrics

All Time Past Year Past 30 Days
Abstract Views 61 61 15
Full Text Views 169 169 67
PDF Downloads 9 9 2
EPUB Downloads 0 0 0