The meaning of weaning in wild Phayre's leaf monkeys: last nipple contact, survival, and independence.

In primates and other mammals, weaning is an equivocal concept, as is reflected in the numerous ways it is measured: a) first intake of solid food, b) conflict over access to the nipple, c) ability to survive without mother, d) maternal resumption of cycling, or e) the cessation of nipple contact. The lack of a consistent definition means that weaning age, although it falls between gestation (fetal growth) and age at first reproduction (most energy diverted from growth), is currently not a reliable life history variable capturing offspring independence. Using data for wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at Phu Khieo Wildlife Sanctuary, Thailand (51 offspring, four groups), we asked whether the end of nipple contact indicates offspring independence as measured by survival to 3 years. To establish a baseline for the onset of independence, we assessed the youngest age at which individuals were orphaned (15-17 months) but then survived to 3 years. Next we determined that offspring age at last nipple contact (19.0 months) was comparable to two other independently calculated measures: offspring age at mother's first postpartum ovulation (11.5 months), and age at mother's re-conception (15.6 months). Using these separate "starting points," we arrived at similar ages for nipple contact cessation (18.4 and 19.2 months, respectively). Overall, in wild (but not in provisioned) Asian colobines, age at last nipple contact was allometrically related to adult female body mass, supporting its designation as a life history variable. Future comparisons need to show if this holds for other taxa.