Early reproductive investment, senescence and lifetime reproductive success in female Asian elephants

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. © 2014 The Authors. Journal of Evolutionary Biology published by John Wiley & Sons Ltd on behalf of European Society for Evolutionary Biology

Limits to Fitness Benefits of Prolonged Post-reproductive Lifespan in Women

Recent advances in medicine and life-expectancy gains have fueled multidisciplinary research into the limits of human lifespan [1-3]. Ultimately, how long humans can live for may depend on selection favoring extended longevity in our evolutionary past [4]. Human females have an unusually extended post-reproductive lifespan, which has been explained by the fitness benefits provided from helping to raise grandchildren following menopause [5, 6]. However, formal tests of whether such grandmothering benefits wane with grandmother age and explain the observed length of post-reproductive lifespan are missing. This is critical for understanding prevailing selection pressures on longevity but to date has been overlooked as a possible mechanism driving the evolution of lifespan. Here, we use extensive data from pre-industrial humans to show that fitness gains from grandmothering are dependent on grandmother age, affecting selection on the length of post-reproductive lifespan. We find both opportunities and ability to help grandchildren declined with age, while the hazard of death of women increased greatly in their late 60s and 70s compared to menopausal ages, together implying waning selection on subsequent longevity. The presence of maternal grand-mothers aged 50-75 increased grandchild survival after weaning, confirming the fitness advantage of post-reproductive lifespan. However, co-residence with paternal grand-mothers aged 75+ was detrimental to grandchild survival, with those grandmothers close to death and presumably in poorer health particularly associated with lower grandchild survival. The age limitations of gaining inclusive fitness from grand-mothering suggests that grandmothering can select for post-reproductive longevity only up to a certain point

Evaluating the reliability of non-specialist observers in the behavioural assessment of semi-captive Asian elephant welfare

Recognising stress is an important component in maintaining the welfare of captive animal populations, and behavioural observation provides a rapid and non-invasive method to do this. Despite substantial testing in zoo elephants, there has been relatively little interest in the application of behavioural assessments to the much larger working populations of Asian elephants across Southeast Asia, which are managed by workers possessing a broad range of behavioural knowledge. Here, we developed a new ethogram of potential stress- and work-related behaviour for a semi-captive population of Asian elephants. We then used this to collect observations from video footage of over 100 elephants and evaluated the reliability of behavioural welfare assessments carried out by non-specialist observers. From observations carried out by different raters with no prior experience of elephant research or management, we tested the reliability of observations between-observers, to assess the general inter-observer agreement, and within-observers, to assess the consistency in behaviour identification. The majority of ethogram behaviours were highly reliable both between- and within-observers, suggesting that overall, behaviour was highly objective and could represent easily recognisable markers for behavioural assessments. Finally, we analysed the repeatability of individual elephant behaviour across behavioural contexts, demonstrating the importance of incorporating a personality element in welfare assessments. Our findings highlight the potential of non-expert observers to contribute to the reliable monitoring of Asian elephant welfare across large captive working populations, which may help to both improve elephant wellbeing and safeguard human workers

Asian elephants exhibit post-reproductive lifespans

Background The existence of extended post-reproductive lifespan is an evolutionary puzzle, and its taxonomic prevalence is debated. One way of measuring post-reproductive life is with post-reproductive representation, the proportion of adult years lived by females after cessation of reproduction. Analyses of post-reproductive representation in mammals have claimed that only humans and some toothed whale species exhibit extended post-reproductive life, but there are suggestions of a post-reproductive stage for false killer whales and Asian elephants. Here, we investigate the presence of post-reproductive lifespan in Asian elephants using an extended demographic dataset collected from semi-captive timber elephants in Myanmar. Furthermore, we investigate the sensitivity of post-reproductive representation values to availability of long-term data over 50 years. Results We find support for the presence of an extended post-reproductive stage in Asian elephants, and that post-reproductive representation and its underlying demographic rates depend on the length of study period in a long-lived animal. Conclusions The extended post-reproductive lifespan is unlikely due to physiological reproductive cessation, and may instead be driven by mating preferences or condition-dependent fertility. Our results also show that it is crucial to revisit such population measures in long-lived species as more data is collected, and if the typical lifespan of the species exceeds the initial study period

Milk Composition of Asian Elephants (Elephas maximus) in a Natural Environment in Myanmar during Late Lactation

The nutritional content of milk from free-living Asian elephants has not previously been reported, despite being vital for better management of captive populations. This study analyzed both milk composition and consumed plant species of Asian elephants managed in their natural environment in Myanmar. Longitudinal samples (n = 36) were obtained during both the wet and the dry season from six mature females in mid to late lactation in 2016 and 2017. Milk composition averaged 82.44% water, with 17.56% total solids containing 5.23% protein, 15.10% fat, 0.87% ash, and 0.18 µg/mL vitamin E. Solids and protein increased with lactation month. Total protein in milk was higher during the wet vs. the dry season. Observed factors linked with maternal (age, parity, size and origin) and calf traits (sex) had significant associations with milk nutrient levels. Primary forages consumed contained moderate protein and fiber. Higher dietary protein during the wet season (11–25%) compared to the dry season (6–19%) may be linked with increased milk protein observed. Our results call for further field studies of milk and diet composition, over entire seasons/lactation periods, and across maternal and calf traits, to improve feeding management, with an overall goal of maximized health and survival

Middleborns disadvantaged? testing birth-order effects on fitness in pre-industrial finns

Parental investment is a limited resource for which offspring compete in order to increase their own survival and reproductive success. However, parents might be selected to influence the outcome of sibling competition through differential investment. While evidence for this is widespread in egg-laying species, whether or not this may also be the case in viviparous species is more difficult to determine. We use pre-industrial Finns as our model system and an equal investment model as our null hypothesis, which predicts that (all else being equal) middleborns should be disadvantaged through competition. We found no overall evidence to suggest that middleborns in a family are disadvantaged in terms of their survival, age at first reproduction or lifetime reproductive success. However, when considering birth-order only among same-sexed siblings, first-, middle-and lastborn sons significantly differed in the number of offspring they were able to rear to adulthood, although there was no similar effect among females. Middleborn sons appeared to produce significantly less offspring than first-or lastborn sons, but they did not significantly differ from lastborn sons in the number of offspring reared to adulthood. Our results thus show that taking sex differences into account is important when modelling birth-order effects. We found clear evidence of firstborn sons being advantaged over other sons in the family, and over firstborn daughters. Therefore, our results suggest that parents invest differentially in their offspring in order to both preferentially favour particular offspring or reduce offspring inequalities arising from sibling competition

Is bigger better? The relationship between size and reproduction in female Asian elephants

The limited availability of resources is predicted to impose trade-offs between growth, reproduction and self-maintenance in animals. However, although some studies have shown that early reproduction suppresses growth, reproduction positively correlates with size in others. We use detailed records from a large population of semi-captive elephants in Myanmar to assess the relationships between size (height and weight), reproduction and survival in female Asian elephants, a species characterized by slow, costly life history. Although female height gain during the growth period overlapped little with reproductive onset in the population, there was large variation in age at first reproduction and only 81% of final weight had been reached by peak age of reproduction at the population level (19 years). Those females beginning reproduction early tended to be taller and lighter later in life, although these trends were not significant. We found that taller females were more likely to have reproduced by a given age, but such effects diminished with age, suggesting there may be a size threshold to reproduction which is especially important in young females. Because size was not linked with female survival during reproductive ages, the diminishing effect of height on reproduction with age is unlikely to be due to biased survival of larger females. We conclude that although reproduction may not always impose significant costs on growth, height may be a limiting factor to reproduction in young female Asian elephants, which could have important implications considering their birth rates are low and peak reproduction is young – 19 years in this population. © 2017 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2017 European Society For Evolutionary Biology</p