title: The role of menopause and reproductive senescence in a long-lived social mammal.
abstract: BACKGROUND
  Menopause is a seemingly maladaptive life-history trait that is found in many long-lived mammals. There are two competing evolutionary hypotheses for this phenomenon; in the adaptive view of menopause, the cessation of reproduction may increase the fitness of older females; in the non-adaptive view, menopause may be explained by physiological deterioration with age. The decline and eventual cessation of reproduction has been documented in a number of mammalian species, however the evolutionary cause of this trait is unknown.
  
  
  RESULTS
  We examined a unique 30-year time series of killer whales, tracking the reproductive performance of individuals through time. Killer whales are extremely long-lived, and may have the longest documented post-reproductive lifespan of any mammal, including humans. We found no strong support for either of the adaptive hypotheses of menopause; there was little support for the presence of post-reproductive females benefitting their daughter's reproductive performance (interbirth interval and reproductive lifespan of daughters), or the number of mature recruits to the population. Oldest mothers (> 35) did appear to have a small positive impact on calf survival, suggesting that females may gain experience with age. There was mixed support for the grandmother hypothesis - grandoffspring survival probabilities were not influenced by living grandmothers, but grandmothers may positively influence survival of juveniles at a critical life stage.
  
  
  CONCLUSIONS
  Although existing data do not allow us to examine evolutionary tradeoffs between survival and reproduction for this species, we were able to examine the effect of maternal age on offspring survival. Our results are consistent with similar studies of other mammals - oldest mothers appear to be better mothers, producing calves with higher survival rates. Studies of juvenile survival in humans have reported positive benefits of grandmothers on newly weaned infants; our results indicate that 3-year old killer whales may experience a positive benefit from helpful grandmothers. While our research provides little support for menopause evolving to provide fitness benefits to mothers or grandmothers, our work supports previous research showing that menopause and long post-reproductive lifespans are not a human phenomenon.
authors.affiliation: Northwest Fisheries Science Center, 2725 Montlake Blvd E, Seattle, WA 98112, USA. eric.ward@noaa.gov.
authors.forename: Eric J
authors.initials: EJ
authors.lastname: Ward
authors.forename: Kim
authors.initials: K
authors.lastname: Parsons
authors.forename: Elizabeth E
authors.initials: EE
authors.lastname: Holmes
authors.forename: Ken C
authors.initials: KC
authors.lastname: Balcomb
authors.forename: John Kb
authors.initials: JK
authors.lastname: Ford