Short-term and delayed effects of mother death on calf mortality in Asian elephants.

Long-lived, highly social species with prolonged offspring dependency can show long postreproductive periods. The Mother hypothesis proposes that a need for extended maternal care of offspring together with increased maternal mortality risk associated with old age select for such postreproductive survival, but tests in species with long postreproductive periods, other than humans and marine mammals, are lacking. Here, we investigate the Mother hypothesis with longitudinal data on Asian elephants from timber camps of Myanmar 1) to determine the costs of reproduction on female age-specific mortality risk within 1 year after calving and 2) to quantify the effects of mother loss on calf survival across development. We found that older females did not show an increased immediate mortality risk after calving. Calves had a 10-fold higher mortality risk in their first year if they lost their mother, but this decreased with age to only a 1.1-fold higher risk in the fifth year. We also detected delayed effects of maternal death: calves losing their mother during early ages still suffered from increased mortality risk at ages 3-4 and during adolescence but such effects were weaker in magnitude. Consequently, the Mother hypothesis could account for the first 5 years of postreproductive survival, but there were no costs of continued reproduction on the immediate maternal mortality risk. However, the observed postreproductive lifespan of females surviving to old age commonly exceeds 5 years in Asian elephants, and further studies are thus needed to determine selection for (postreproductive) lifespan in elephants and other comparably long-lived species.