Nahallage, C. A. D., Leca, J.-B., Huffman, M. A. 2016. Stone handling, an object play behaviour in macaques: Welfare and neurological health implications of a bio-culturally driven tradition. Behaviour, 153(6-7), 845-869.

Object play in primates is viewed as generally having no immediate functional purpose, limited for the most part to immature individuals. At the proximate level, the occurrence of object play in immatures is regarded as being intrinsically self-rewarding, with the ultimate function of supporting motoneuronal development and the acquisition of skills necessary to prepare them for survival as adults. Stone handling (SH), a solitary object play behaviour occurs, and has been studied, in multiple free-ranging and captive troops of provisioned Japanese macaques, as well as rhesus and long-tailed macaques for over 35 years now. A review of our combined findings from these observations reveal that infants acquire SH in the first 3-4 months of life and exhibit increasingly more complex and varied behavioural patterns with age. The longitudinal data shows that many individuals maintain this activity throughout life, practicing it under relaxed ecological and social conditions. The ultimate function may be bimodal, promoting motor development in young and neural maintenance and regeneration in adult and aging individuals.