A systematic review of sex differences in rough and tumble play across non-human mammals

It is widely believed that juvenile male mammals typically engage in higher rates of rough and tumble play (RTP) than do females, in preparation for adult roles involving intense physical competition between males. The consistency of this sex difference across diverse mammalian species has, however, not yet been systematically investigated, limiting our current understanding of its possible adaptive function. This review uses narrative synthesis to (i) evaluate the ubiquity of male-biased RTP across non-human mammals, (ii) identify patterns of variation within and between taxonomic groups, and (iii) propose possible predictors of variation in these differences, including methodological and socio-ecological factors, for investigation by future studies. We find that most species studied do exhibit higher rates or RTP in males than females, while female-biased RTP is rare. Sex differences are smaller and less consistent than expected, with many studies finding similar rates of RTP in males and females. We identify multiple potential socio-ecological predictors of variation in sex differences in RTP, such as intrasexual competition and dietary niche. However, variation is not strongly phylogenetically patterned, suggesting that methodological and environmental factors, such as sample size and play partner availability, are important to consider in future comparative analyses. Significance statement Rough and tumble play (RTP) is thought to be vital for developing physical skills necessary for aggressive competition in adulthood, explaining an apparently widespread sex difference in RTP in mammals whereby immature males are more likely to engage in this behaviour than females. However, no prior study has systematically investigated the extent to which a male bias in RTP is consistent across diverse mammalian species. We find that although RTP is commonly male biased, findings were highly variable both within- and between-species, and equal participation in RTP by males and females is more common than widely assumed. Our review suggests several potential predictors of variation in sex differences in RTP, particularly levels of intrasexual competition in both males and females. However, our findings also suggest the importance of considering methodological in addition to socio-ecological factors for future research.