Variable social organization is ubiquitous in Artiodactyla and probably evolved from pair-living ancestors

Previous studies to understand the evolution of specific variation in mammalian social organization (SO; composition of social units) produced inconsistent results, possibly by ignoring specific variation. Here we present systematic data on SO in artiodactyl populations, coding SO as solitary, pair-living, group-living, sex-specific or variable (different kinds of SOs in the same population). We found that 62% of 245 populations and 83% of species (83/100) exhibited variable SO. Using Bayesian phylogenetic mixed-effects models, we simultaneously tested whether research effort, habitat, sexual dimorphism, breeding seasonality or body size predicted the likelihood of different SOs and inferred the ancestral SO. Body size and sexual dimorphism were strongly associated with different SOs. Contingent on the small body size (737 g) and putative sexual monomorphism of the earliest fossil artiodactyl, the ancestral SO was most likely to be pair-living (probability = 0.76, 95% CI = 0-1), followed by variable ( = 0.19, 95% CI = 0-0.99). However, at body size values typical of extant species, variable SO becomes the dominant form ( = 0.74, 95% CI = 0.18-1.00). Distinguishing different kinds of 'variable' highlights transitions from SOs involving pair-living to SOs involving solitary and/or group-living with increasing body size and dimorphism. Our results support the assumption that ancestral artiodactyl was pair-living and highlight the ubiquity of intraspecific variation in SO.