Reproductive cessation and post-reproductive lifespan in Asian elephants and pre-industrial humans

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.</p

Nearby grandmother enhances calf survival and reproduction in Asian elephants

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

Differences in age-specific mortality between wild-caught and captive-born Asian elephants

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

Parasite-associated mortality in a long-lived mammal: Variation with host age, sex, and reproduction.

Parasites can cause severe host morbidity and threaten survival. As parasites are generally aggregated within certain host demographics, they are likely to affect a small proportion of the entire population, with specific hosts being at particular risk. However, little is known as to whether increased host mortality from parasitic causes is experienced by specific host demographics. Outside of theoretical studies, there is a paucity of literature concerning dynamics of parasite-associated host mortality. Empirical evidence mainly focuses on short-lived hosts or model systems, with data lacking from long-lived wild or semi-wild vertebrate populations. We investigated parasite-associated mortality utilizing a multigenerational database of mortality, health, and reproductive data for over 4,000 semi-captive timber elephants (Elephas maximus), with known causes of death for mortality events. We determined variation in mortality according to a number of host traits that are commonly associated with variation in parasitism within mammals: age, sex, and reproductive investment in females. We found that potentially parasite-associated mortality varied significantly across elephant ages, with individuals at extremes of lifespan (young and old) at highest risk. Mortality probability was significantly higher for males across all ages. Female reproducers experienced a lower probability of potentially parasite-associated mortality than females who did not reproduce at any investigated time frame. Our results demonstrate increased potentially parasite-associated mortality within particular demographic groups. These groups (males, juveniles, elderly adults) have been identified in other studies as susceptible to parasitism, stressing the need for further work investigating links between infection and mortality. Furthermore, we show variation between reproductive and non-reproductive females, with mothers being less at risk of potentially parasite mortality than nonreproducers

Elephants born in the high stress season have faster reproductive ageing

Senescent declines in reproduction and survival are found across the tree of life, but little is known of the factors causing individual variation in reproductive ageing rates. One contributor may be variation in early developmental conditions, but only a few studies quantify the effects of early environment on reproductive ageing and none concern comparably long-lived species to humans. We determine the effects of &lsquo;stressful&rsquo; birth conditions on lifetime reproduction in a large semi-captive population of Asian elephants (Elephas maximus). We categorise birth month into stressful vs. not-stressful periods based on longitudinal measures of glucocorticoid metabolites in reproductive-aged females, which peak during heavy workload and the start of the monsoon in June-August. Females born in these months exhibit faster reproductive senescence in adulthood and have significantly reduced lifetime reproductive success than their counterparts born at other times of year. Improving developmental conditions could therefore delay reproductive ageing in species as long-lived as humans

Compromised survivorship in zoo elephants

Keeping elephants in zoos is extremely costly, yet does not yield self-sustaining 16 populations. In Europe, which holds c. half the global zoo elephant population, a long17 term decline of c.10% per year is expected in both species, if reliant on zoo-bred animals 18 under historically prevailing conditions. Fitness in zoos is compromised in several ways. 19 Compared with protected in situ populations (Burmese working Asians; Kenyan free20 living Africans), zoo elephants show premature reproductive senescence and -- despite 21 improving adult survivorship for Africans -- die earlier in adulthood than expected. In 22 Asian elephants, infant survivorship in zoos is also greatly reduced relative to Burmese 23 elephants, and furthermore, zoo-born animals die earlier in adulthood than wild-caught 24 conspecifics kept in zoos, via effects ‘programmed’ peri-natally. In this species, being 25 transferred between zoos also increases mortality rates. Both survival and fecundity 26 would need to improve to attain self-sustaining zoo populations. Our findings 27 demonstrate deficits in zoo elephant management, particularly for Asians, and implicate 28 stress and obesity as likely problems

High levels of mitochondrial genetic diversity in Asian elephants (Elephas maximus) from Myanmar

L

Changes in age-structure over four decades were a key determinant of population growth rate in a long-lived mammal

A changing environment directly influences birth and mortality rates, and thus population growth rates. However, population growth rates in the short term are also influenced by population age-structure. Despite its importance, the contribution of age-structure to population growth rates has rarely been explored empirically in wildlife populations with long-term demographic data. Here we assessed how changes in age-structure influenced short-term population dynamics in a semi-captive population of Asian elephantsElephas maximus. We addressed this question using a demographic dataset of female Asian elephants from timber camps in Myanmar spanning 45 years (1970-2014). First, we explored temporal variation in age-structure. Then, using annual matrix population models, we used a retrospective approach to assess the contributions of age-structure and vital rates to short-term population growth rates with respect to the average environment. Age-structure was highly variable over the study period, with large proportions of juveniles in the years 1970 and 1985, and made a substantial contribution to annual population growth rate deviations. High adult birth rates between 1970 and 1980 would have resulted in large positive population growth rates, but these were prevented by a low proportion of reproductive-aged females. We highlight that an understanding of both age-specific vital rates and age-structure is needed to assess short-term population dynamics. Furthermore, this example from a human-managed system suggests that the importance of age-structure may be accentuated in populations experiencing human disturbance where age-structure is unstable, such as those in captivity or for endangered species. Ultimately, changes to the environment drive population dynamics by influencing birth and mortality rates, but understanding demographic structure is crucial for assessing population growth

Parasite-associated mortality in a long-lived mammal: Variation with host age, sex, and reproduction

Parasites can cause severe host morbidity and threaten survival. As parasites are generally aggregated within certain host demographics, they are likely to affect a small proportion of the entire population, with specific hosts being at particular risk. However, little is known as to whether increased host mortality from parasitic causes is experienced by specific host demographics. Outside of theoretical studies, there is a paucity of literature concerning dynamics of parasite-associated host mortality. Empirical evidence mainly focuses on short-lived hosts or model systems, with data lacking from long-lived wild or semi-wild vertebrate populations. We investigated parasite-associated mortality utilizing a multigenerational database of mortality, health, and reproductive data for over 4,000 semi-captive timber elephants (Elephas maximus), with known causes of death for mortality events. We determined variation in mortality according to a number of host traits that are commonly associated with variation in parasitism within mammals: age, sex, and reproductive investment in females. We found that potentially parasite-associated mortality varied significantly across elephant ages, with individuals at extremes of lifespan (young and old) at highest risk. Mortality probability was significantly higher for males across all ages. Female reproducers experienced a lower probability of potentially parasite-associated mortality than females who did not reproduce at any investigated time frame. Our results demonstrate increased potentially parasite-associated mortality within particular demographic groups. These groups (males, juveniles, elderly adults) have been identified in other studies as susceptible to parasitism, stressing the need for further work investigating links between infection and mortality. Furthermore, we show variation between reproductive and non-reproductive females, with mothers being less at risk of potentially parasite mortality than nonreproducers.</p