title: Reproductive cessation and post-reproductive lifespan in Asian elephants and pre-industrial humans.
abstract: BACKGROUND
  Short post-reproductive lifespan is widespread across species, but prolonged post-reproductive life-stages of potential adaptive significance have been reported only in few mammals with extreme longevity. Long post-reproductive lifespan contradicts classical evolutionary predictions of simultaneous senescence in survival and reproduction, and raises the question of whether extreme longevity in mammals promotes such a life-history. Among terrestrial mammals, elephants share the features with great apes and humans, of having long lifespan and offspring with long dependency. However, little data exists on the frequency of post-reproductive lifespan in elephants. Here we use extensive demographic records on semi-captive Asian elephants (n = 1040) and genealogical data on pre-industrial women (n = 5336) to provide the first comparisons of age-specific reproduction, survival and post-reproductive lifespan in both of these long-lived species.
  
  
  RESULTS
  We found that fertility decreased after age 50 in elephants, but the pattern differed from a total loss of fertility in menopausal women with many elephants continuing to reproduce at least until the age of 65 years. The probability of entering a non-reproductive state increased steadily in elephants from the earliest age of reproduction until age 65, with the longer living elephants continuing to reproduce until older ages, in contrast to humans whose termination probability increased rapidly after age 35 and reached 1 at 56 years, but did not depend on longevity. Post-reproductive lifespan reached 11-17 years in elephants and 26-27 years in humans living until old age (depending on method), but whereas half of human adult lifespan (of those reproductive females surviving to the age of 5% fecundity) was spent as post-reproductive, only one eighth was in elephants. Consequently, although some elephants have long post-reproductive lifespans, relatively few individuals reach such a phase and the decline in fertility generally parallels declines in survivorship in contrast to humans with a decoupling of senescence in somatic and reproductive functions.
  
  
  CONCLUSIONS
  Our results show that the reproductive and survival patterns of Asian elephants differ from other long-lived animals exhibiting menopause, such as humans, and extreme longevity alone does not promote the evolution of menopause or post-reproductive lifespan, adding weight to the unusual kin-selected benefits suggested to favour such traits in humans and killer whales.
authors.affiliation: Section of Ecology, Department of Biology, University of Turku, Turku FIN-20014, Finland.
authors.forename: Mirkka
authors.initials: M
authors.lastname: Lahdenperä
authors.affiliation: Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK.
authors.forename: Khyne U
authors.initials: KU
authors.lastname: Mar
authors.affiliation: Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK.
authors.forename: Virpi
authors.initials: V
authors.lastname: Lummaa
keywords: Age-specific fertility; Ageing; Reproduction; Senescence; Survival