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IntroductionShort 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.ResultsWe 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.ConclusionsOur 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.

Juvenile mortality is a key factor influencing population growth rate in density-independent, predation-free, well-managed captive populations. Currently at least a quarter of all Asian elephants live in captivity, but both the wild and captive populations are unsustainable with the present fertility and calf mortality rates. Despite the need for detailed data on calf mortality to manage effectively populations and to minimize the need for capture from the wild, very little is known of the causes and correlates of calf mortality in Asian elephants. Here we use the world's largest multigenerational demographic dataset on a semi-captive population of Asian elephants compiled from timber camps in Myanmar to investigate the survival of calves (n = 1020) to age five born to captive-born mothers (n = 391) between 1960 and 1999. Mortality risk varied significantly across different ages and was higher for males at any age. Maternal reproductive history was associated with large differences in both stillbirth and liveborn mortality risk: first-time mothers had a higher risk of calf loss as did mothers producing another calf soon (<3.7 years) after a previous birth, and when giving birth at older age. Stillbirth (4%) and pre-weaning mortality (25.6%) were considerably lower than those reported for zoo elephants and used in published population viability analyses. A large proportion of deaths were caused by accidents and lack of maternal milk/calf weakness which both might be partly preventable by supplementary feeding of mothers and calves and work reduction of high-risk mothers. Our results on Myanmar timber elephants with an extensive keeping system provide an important comparison to compromised survivorship reported in zoo elephants. They have implications for improving captive working elephant management systems in range countries and for refining population viability analyses with realistic parameter values in order to predict future population size of the Asian elephant.

Human menopause is ubiquitous among women and is uninfluenced by modernity. In addition, it remains an evolutionary puzzle: studies have largely failed to account for diminishing selection on reproduction beyond 50 years. Using a 200-year dataset on pre-industrial Finns, we show that an important component is between-generation reproductive conflict among unrelated women. Simultaneous reproduction by successive generations of in-laws was associated with declines in offspring survivorship of up to 66%. An inclusive fitness model revealed that incorporation of the fitness consequences of simultaneous intergenerational reproduction between in-laws, with those of grandmothering and risks of dying in childbirth, were sufficient to generate selection against continued reproduction beyond 51 years. Decomposition of model estimates suggested that the former two were most influential in generating selection against continued reproduction. We propose that menopause evolved, in part, because of age-specific increases in opportunities for intergenerational cooperation and reproductive competition under ecological scarcity.

The evolutionary theory of senescence posits that as the probability of extrinsic mortality increases with age, selection should favour early-life over late-life reproduction. Studies on natural vertebrate populations show early reproduction may impair later-life performance, but the consequences for lifetime fitness have rarely been determined, and little is known of whether similar patterns apply to mammals which typically live for several decades. We used a longitudinal dataset on Asian elephants (Elephas maximus) to investigate associations between early-life reproduction and female age-specific survival, fecundity and offspring survival to independence, as well as lifetime breeding success (lifetime number of calves produced). Females showed low fecundity following sexual maturity, followed by a rapid increase to a peak at age 19 and a subsequent decline. High early life reproductive output (before the peak of performance) was positively associated with subsequent age-specific fecundity and offspring survival, but significantly impaired a female's own later-life survival. Despite the negative effects of early reproduction on late-life survival, early reproduction is under positive selection through a positive association with lifetime breeding success. Our results suggest a trade-off between early reproduction and later survival which is maintained by strong selection for high early fecundity, and thus support the prediction from life history theory that high investment in reproductive success in early life is favoured by selection through lifetime fitness despite costs to later-life survival. That maternal survival in elephants depends on previous reproductive investment also has implications for the success of (semi-)captive breeding programmes of this endangered species.

Usually animals reproduce into old age, but a few species such as humans and killer whales can live decades after their last reproduction. The grandmother hypothesis proposes that such life-history evolved through older females switching to invest in their existing (grand)offspring, thereby increasing their inclusive fitness and selection for post-reproductive lifespan. However, positive grandmother effects are also found in non-menopausal taxa, but evidence of their associated fitness effects is rare and only a few tests of the hypothesis in such species exist. Here we investigate the grandmother effects in Asian elephants. Using a multigenerational demographic dataset on semi-captive elephants in Myanmar, we found that grandcalves from young mothers (<20 years) had 8 times lower mortality risk if the grandmother resided with her grandcalf compared to grandmothers residing elsewhere. Resident grandmothers also decreased their daughters’ inter-birth intervals by one year. In contrast to the hypothesis predictions, the grandmother’s own reproductive status did not modify such grandmother benefits. That elephant grandmothers increased their inclusive fitness by enhancing their daughter’s reproductive rate and success irrespective of their own reproductive status suggests that fitness-enhancing grandmaternal effects are widespread, and challenge the view that grandmother effects alone select for menopause coupled with long post-reproductive lifespan.

Wild-capture of numerous species is common for diverse purposes, including medical experiments, conservation, veterinary interventions and research, but little objective data exists on its consequences. We use exceptional demographic records on Asian elephants from timber camps in Myanmar to investigate the long-term consequences of wild-capture during 1951–2000 on their mortality (N = 5150). We show that captured elephants have increased mortality compared to captive-born elephants, regardless of their capture method. These detrimental effects of capture are similar for both sexes but differ substantially according to age. Elephants captured and tamed at older ages show a higher increase in mortality after capture than elephants captured and tamed young. Moreover, the increased mortality risk following capture and taming is still perceived several years after capture. Our results are timely given the continued capture of elephants and other wild animals to supplement captive populations despite the alarming declines of wild populations globally.

Data on personality for long-lived, highly social wild mammals with high cognitive abilities are rare. We investigated the personality structure of Asian elephants (Elephas maximus) by using a large sample of semi-captive timber elephants in Myanmar. Data were collected during 2014–2017 using questionnaires, for which elephant riders (mahouts) rated 28 behavioural adjectives of elephants. Repeated questionnaires were obtained for each elephant from several raters whenever possible, resulting in 690 ratings of 150 female and 107 male elephants. We started by performing a confirmatory factor analysis to compare the fit of our data to a previously published captive elephant personality structure. Owing to a poor fit of this model to our data, we proceeded by performing explanatory factor analysis to determine the personality structure in our study population. This model suggested that personality in these elephants was manifested as three factors that we labelled as Attentiveness, Sociability and Aggressiveness. This structure did not differ between the sexes. These results provide the basis for future research on the link between personality and reproductive success in this endangered species and more generally, help to resolve the selective pressures on personalities in long-lived, highly social species.