All together now: Assessing variation in maternal and nonmaternal handling of wild Colobus vellerosus infants.

Primatologists have a long-standing interest in the study of maternal care and nonmaternal handling (NH) of infants stemming from recognition that early social relationships can have enduring consequences. Though maternal care and NH often include expression of similar behaviors, they are regularly studied in isolation from each other with nonoverlapping terminology, thereby overlooking possible interplay between them and obfuscating potential developmental ramifications that ensue from trade-offs made between maternal (MH) and NH during infancy. To that end, identifying how MH and NH patterns interact and contribute to the total handling (TH) infants receive is a critical first step. We present durational handling data collected from 25 wild Colobus vellerosus infants from 2016 to 2017 and assess the relationships between TH, MH, and NH. Patterns of social affiliation are shaped in part by surrounding context, and therefore, we also assess whether NH and TH differ in their responsivity to various infant and social group characteristics. Ninety-four percent of observed handling was MH, while just 5.5% was NH. Young infants who received more MH (excluding nursing) also received more NH; there was no relationship between the two in older infants. Infants in larger groups participated in more handling of all types. Additionally, NH time was associated with infant sex and group stability. Non-nursing TH time was associated with group stability and infant cohort size. Though NH variation likely confers social-networking advantage, in this population NH is not a major contributor to TH and would not effectively replace reduced MH. The positive association between MH and NH during early infancy suggests that colobus mothers may play a mediating role in shaping infant socialization. This is a first step in elucidating how different forms of handling relate to one another in wild primates and in identifying the impact of handling on infant socialization.