Asian elephant calf physiology and mahout perspectives during taming in Myanmar.

A quarter of Asian elephants are captive, with greater than 90% of these tamed and cared for by handlers (mahouts) in Asia. Although taming is a much-discussed welfare issue, no studies to our knowledge have empirically assessed its impact on calves, and dialogue surrounding taming often lacks perspectives of those involved. Here, we interviewed mahouts involved in taming and monitored five physiological measures (faecal glucocorticoid metabolites (FGMs), serum cortisol, glucose, creatine kinase (CK) and heterophil:lymphocyte (H:L)) over the first 10 days of taming and following six months in 41 calves undergoing taming and 16 control individuals. These measures assess the duration and intensity of stress during taming. Interviews suggested mahouts had major concerns for their safety when discussing changing taming practices, an important consideration for future management. Calf physiological measures were elevated by 50-70% (FGMs/cortisol/glucose), 135% (H:L) and greater than 500% (CK) over the first few days of taming, indicative of elevated stress, not seen to the same extent in control adults. Some measures stabilized sooner (glucose/cortisol/CK/FGM: 7-10 days) than others (H:L: one-two months), indicating mostly acute stress. Our findings inform the welfare of approximately 15 000 captive elephants around the world. Future studies should compare taming in different populations and consider calf and mahout welfare.

Grandmother effects over the Finnish demographic transition.

Demographic transitions are defining events for human societies, marking shifts from natural mortality and fertility rates to the low rates seen in industrialised populations. These transitions can affect trait evolution through altering the direction and strength of selection when variance in fertility and mortality decline. One key feature of human evolution is the evolution of extended post-reproductive life through indirect fitness benefits from grandmothering. Although studies in pre- and post-transition societies have documented beneficial grandmother presence, it remains unknown whether these associations changed before, during, or after the transition. Here, we use genealogical data from eighteenth- to twientieth-century Finland to show grandmother-associated changes of two measures of evolutionary fitness (grandchild survival and birth rate) over the transition. We find that grandmothers had greater opportunity to help as the transition progressed, but their effect on grandchild survival declined alongside general mortality rates, implying that selection on lifespan from grandmothering declined too. Whilst grandmother presence was still associated with reduced birth intervals and hence more grandchildren born post-transition, the nature of this relationship changed greatly. This suggests that although potential for intergenerational interactions increased over the demographic transition, the (hypothesised) evolutionary importance of these interactions declined, which reduced selection for extended post-reproductive lifespan.

Seasonal variation in molecular and physiological stress markers in Asian elephants.

Free-living species exhibit seasonal variation in various life history traits, including vital rates such as birth and death patterns. Different physiological mechanisms are thought to underlie the expression of life history traits that contribute to lifetime fitness. However, although the broad impacts of seasonality on life history traits and trade-offs is well established in many systems, the exact physiological mechanisms responsible for driving differences within and between individuals are poorly understood. Among them, molecular and physiological stress pathways, such as stress hypothalamic-pituitary-adrenal axis and oxidative stress, have potential to mediate relationships between individual survival, reproduction and environmental seasonality. Here, we determine how different physiological markers of stress including faecal cortisol metabolites (FCMs), heterophils/lymphocytes (H/L) ratio, two markers indicating oxidative balance including a marker of oxidative damage (reactive oxygen metabolites, ROM) and a marker of antioxidant defences (superoxide dismutase, SOD) and body weight vary in a large semi-captive population of Asian elephants (Elephas maximus) exposed to extreme seasonality (e.g. elevated temperatures). Individuals showed higher FCM levels and H/L ratios during cold season, indicating increased stress, and the lowest FCM levels during monsoon season and H/L ratios during hot and dry season, but we found no pattern in oxidative stress (ROM and SOD) levels. Hot season also associated with a decline in body weight. The present study shows how different physiological parameters (FCM levels and H/L ratio), molecular (oxidative stress) and body condition vary with seasonal changes, and how these parameters might allow individuals to adapt to such variations. Our results on an endangered long-lived species are crucial in indicating the most productive timing for conservation efforts, predicting how individuals cope with environmental changes, and allow for a more accurate representation of how animal physiology operates in nature.

Sex, age, and family structure influence dispersal behaviour after a forced migration.

Dispersal does not only mean moving from one environment to another, but can also refer to shifting from one social group to another. Individual characteristics such as sex, age and family structure might influence an individual's propensity to disperse. In this study, we use a unique dataset of an evacuated World War II Finnish population, to test how sex, age, number of siblings and birth order influence an individual's dispersal away from their own social group at a time when society was rapidly changing. We found that young women dispersed more than young men, but the difference decreased with age. This suggests that young men might benefit more from staying near a familiar social group, whereas young women could benefit more from moving elsewhere to find work or spouses. We also found that having more younger brothers increased the propensity for firstborns to disperse more than for laterborns, indicating that younger brothers might pressure firstborn individuals into leaving. However, sisters did not have the same effect as brothers. Overall, the results show that individual characteristics are important in understanding dispersal behaviour, but environmental properties such as social structure and the period of flux after World War II might upend the standard predictions concerning residence and dispersal. Social media summary: Individual characteristics influence dispersal away from social group after a forced migration in a Finnish population.

First report of reindeer (Rangifer tarandus tarandus) response to human-given cues.

Many argue that the animal understanding of human referential communication is a by-product of domestication. However, the domestication hypothesis is not unanimously supported as some nondomesticated species such as sea lions, dolphins, or African elephants perform well in the understanding of human pointing gesture. There is a need to study species with different levels of domestication across different taxa to understand the emerging communicative sociocognitive skills in animals that provide them with the ability to comprehend human-given cues. We conducted a pilot study to assess the performance of eight sledging reindeer following a commonly used human-given cue (a pointing gesture associated with gaze at the target and local enhancement) in a two-way choice task. Domestic reindeer are considered semicaptive, because of their husbandry system in free-ranging conditions, with limited control of their reproduction. We observed that the willingness of the reindeer to participate in the test was age-related, with the younger individuals which lack experience with humans being reluctant to approach the experimenters. Within the more experienced working sledging reindeer, two individuals showed excellent skills and followed the human-given cues 9 out of 10 times. Reindeer show great potential in following a human indication to locate hidden food with minimal training when properly tamed. The effect of previous experience with humans requires further investigation. This is the first demonstration in cervids of an ability to make use of experimenter-given cues in an object-choice task. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Grandmother presence improved grandchild survival against childhood infections but not vaccination coverage in historical Finns.

Grandmother presence can improve the number and survival of their grandchildren, but what grandmothers protect against and how they achieve it remains poorly known. Before modern medical care, infections were leading causes of childhood mortality, alleviated from the nineteenth century onwards by vaccinations, among other things. Here, we combine two individual-based datasets on the genealogy, cause-specific mortality and vaccination status of eighteenth- and nineteenth-century Finns to investigate two questions. First, we tested whether there were cause-specific benefits of grandmother presence on grandchild survival from highly lethal infections (smallpox, measles, pulmonary and diarrhoeal infections) and/or accidents. We show that grandmothers decreased all-cause mortality, an effect which was mediated through smallpox, pulmonary and diarrhoeal infections, but not via measles or accidents. Second, since grandmothers have been suggested to increase vaccination coverage, we tested whether the grandmother effect on smallpox survival was mediated through increased or earlier vaccination, but we found no evidence for such effects. Our findings that the beneficial effects of grandmothers are in part driven by increased survival from some (but not all) childhood infections, and are not mediated via vaccination, have implications for public health, societal development and human life-history evolution.

Grandparental co-residence and grandchild survival: the role of resource competition in a pre-industrial population.

Although grandparents are and have been important alloparents to their grandchildren, they are not necessarily only beneficial but can also compete with grandchildren over limited resources. Competition over parental care or other resources may exist especially if grandparents live in the same household with grandchildren and it can be dependent on grandchild age. By utilizing demographic data collected from historic population registers in Finland between 1761 and 1895 (study sample n = 4041) we investigate whether grandparents living in the same household with grandchildren are detrimental or beneficial for grandchild survival. Having a living but not co-residing grandmother or grandfather were both associated with better survival whereas having a co-resident grandfather was associated with lower chance to survive for infants (age < 1 year). Separating the effect between maternal and paternal grandparents and grandmothers and grandfathers revealed no differences in the effects between lineages. Negative effect of having a co-residing grandfather was not significant when grandfathers were separated for lineage specific models. These results implicate that accounting for the co-residence status and child's age, grandparents were mostly beneficial when not co-residing with very young children and that having a co-residing grandfather at that age could be associated with lower chances to survive. Predictions made by grandmother hypothesis and resource competition both received support. The results presented here also offered comparison points to preindustrial and contemporary three-generational families.

Sibling competition, dispersal and fitness outcomes in humans.

Determining how sibling interactions alter the fitness outcomes of dispersal is pivotal for the understanding of family living, but such studies are currently scarce. Using a large demographic dataset on pre-industrial humans from Finland, we studied dispersal consequences on different indicators of lifetime reproductive success according to sex-specific birth rank (a strong determinant of dispersal in our population). Contrary to the predictions of the leading hypotheses, we found no support for differential fitness benefits of dispersal for either males or females undergoing low vs. high sibling competition. Our results are inconsistent with both hypotheses that family members could have different fitness maximizing strategies depending on birth rank, and that dispersal could be mainly driven by indirect fitness benefits for philopatric family members. Our study stresses the need for studying the relative outcomes of dispersal at the family level in order to understand the evolution of family living and dispersal behaviour.

The long-lasting legacy of reproduction: lifetime reproductive success shapes expected genetic contributions of humans after 10 generations.

An individual's lifetime reproductive success (LRS) measures its realized genetic contributions to the next generation, but how well does it predict this over longer periods? Here we use human genealogical data to estimate expected individual genetic contributions (IGC) and quantify the degree to which LRS, relative to other fitness proxies, predicts IGC over longer periods. This allows an identification of the life-history stages that are most important in shaping variation in IGC. We use historical genealogical data from two non-isolated local populations in Switzerland to estimate the stabilized IGC for 2230 individuals approximately 10 generations after they were born. We find that LRS explains 30% less variation in IGC than the best predictor of IGC, the number of grandoffspring. However, albeit less precise than the number of grandoffspring, we show that LRS does provide an unbiased prediction of IGC. Furthermore, it predicts IGC better than lifespan, and accounting for offspring survival to adulthood does not improve the explanatory power. Overall, our findings demonstrate the value of human genealogical data to evolutionary biology and suggest that reproduction-more than lifespan or offspring survival-impacts the long-term genetic contributions of historic humans, even in a population with appreciable migration.

What drives horse success at following human-given cues? An investigation of handler familiarity and living conditions.

Cues such as the human pointing gesture, gaze or proximity to an object are widely used in behavioural studies to evaluate animals' abilities to follow human-given cues. Many domestic mammals, such as horses, can follow human cues; however, factors influencing their responses are still unclear. We assessed the performance of 57 horses at a two-way choice task testing their ability to follow cues of either a familiar (N = 28) or an unfamiliar informant (N = 29). We investigated the effects of the length of the relationship between the horse and a familiar person (main caregiver), their social environment (living alone, in dyads, or in groups) and their physical environment (living in stalls/paddocks, alternating between paddocks and pastures, or living full time in pastures). We also controlled for the effects of horses' age and sex. Our results showed that horses' success rate at the task was not affected by the familiarity of the informant and did not improve with the relationship length with the familiar informant but did increase with the age of the horses. Horses living in groups had better success than the ones kept either in dyads or alone. Finally, horses housed in small paddocks had lower success than those living on pasture. These results indicate that with age, horses get better at following human-given indications regardless of who the human informant is and that an appropriate living and social environment could contribute to the development of socio-cognitive skills towards humans. Therefore, such aspects should be considered in studies evaluating animal behaviour.

Correction to: Female-biased sex ratios in urban centers create a "fertility trap" in post-war Finland.

[This corrects the article DOI: 10.1093/beheco/arab007.].

Investigating associations between nematode infection and three measures of sociality in Asian elephants.

Abstract: Frequent social interactions, proximity to conspecifics, and group density are main drivers of infections and parasite transmissions. However, recent theoretical and empirical studies suggest that the health benefits of sociality and group living can outweigh the costs of infection and help social individuals fight infections or increase their infection-related tolerance level. Here, we combine the advantage of studying artificially created social work groups with different demographic compositions with free-range feeding and social behaviours in semi-captive Asian elephants (Elephas maximus), employed in timber logging in Myanmar. We examine the link between gastro-intestinal nematode load (strongyles and Strongyloides spp.), estimated by faecal egg counts, and three different aspects of an elephant's social world: individual solitary behaviour, work group size, and work group sex ratio. Controlling for sex, age, origin, time since last deworming treatment, year, human sampler bias, and individual identity, we found that infection by nematodes ranged from 0 to 2720 eggs/g between and within 26 male and 45 female elephants over the 4-year study period. However, such variation was not linked to any investigated measures of sociality in either males or females. Our findings highlight the need for finer-scale studies, establishing how sociality is limited by, mitigates, or protects against infection in different ecological contexts, to fully understand the mechanisms underlying these pathways. Significance statement: Being social involves not only benefits, such as improved health, but also costs, including increased risk of parasitism and infectious disease. We studied the relationship between and three different sociality measures-solitary behaviour, group size, and the proportion of females to males within a group-and infection by gut nematodes (roundworms), using a unique study system of semi-captive working Asian elephants. Our system allows for observing how infection is linked to sociality measures across different social frameworks. We found that none of our social measures was associated with nematode infection in the studied elephants. Our results therefore suggest that here infection is not a large cost to group living, that it can be alleviated by the benefits of increased sociality, or that there are weak infection-sociality associations present which could not be captured and thus require finer-scale measures than those studied here. Overall, more studies are needed from a diverse range of systems that investigate specific aspects of social infection dynamics. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-022-03192-8.

Mothers with higher twinning propensity had lower fertility in pre-industrial Europe.

Historically, mothers producing twins gave birth, on average, more often than non-twinners. This observation has been interpreted as twinners having higher intrinsic fertility - a tendency to conceive easily irrespective of age and other factors - which has shaped both hypotheses about why twinning persists and varies across populations, and the design of medical studies on female fertility. Here we show in >20k pre-industrial European mothers that this interpretation results from an ecological fallacy: twinners had more births not due to higher intrinsic fertility, but because mothers that gave birth more accumulated more opportunities to produce twins. Controlling for variation in the exposure to the risk of twinning reveals that mothers with higher twinning propensity - a physiological predisposition to producing twins - had fewer births, and when twin mortality was high, fewer offspring reaching adulthood. Twinning rates may thus be driven by variation in its mortality costs, rather than variation in intrinsic fertility.

Sex-specific body mass ageing trajectories in adult Asian elephants.

In species with marked sexual dimorphism, the classic prediction is that the sex which undergoes stronger intrasexual competition ages earlier or quicker. However, more recently, alternative hypotheses have been put forward, showing that this association can be disrupted. Here, we utilize a unique, longitudinal data set of a semi-captive population of Asian elephants (Elephas maximus), a species with marked male-biased intrasexual competition, with males being larger and having shorter lifespans, and investigate whether males show earlier and/or faster body mass ageing than females. We found evidence of sex-specific body mass ageing trajectories: adult males gained weight up to the age of 48 years old, followed by a decrease in body mass until natural death. In contrast, adult females gained body mass with age until a body mass decline in the last year of life. Our study shows sex-specific ageing patterns, with an earlier onset of body mass declines in males than females, which is consistent with the predictions of the classical theory of ageing.

The Long-Term Success of Mandatory Vaccination Laws After Implementing the First Vaccination Campaign in 19th Century Rural Finland.

In high-income countries, childhood infections are on the rise, a phenomenon attributed in part to persistent hesitancy toward vaccines. To combat vaccine hesitancy, several countries recently made vaccinating children mandatory, but the effect of such vaccination laws on vaccination coverage remains debated, and the long-term consequences are unknown. Here we quantified the consequences of vaccination laws on vaccination coverage, monitoring for a period of 63 years (1837-1899) rural Finland's first vaccination campaign against the highly lethal childhood infection smallpox. We found that annual vaccination campaigns were focused on children up to 1 year old and that their vaccination coverage was low and declined over time until the implementation of the vaccination law, which stopped the declining trend and was associated with an abrupt coverage increase, of 20%, to cover >80% of all children. Our results indicate that vaccination laws can have a long-term beneficial effect of increasing the vaccination coverage and will help public health practitioners to make informed decisions on how to act against vaccine hesitancy and optimize the impact of vaccination programs.

Neighborhood Disadvantage, Greenness, and Population Density as Predictors of Breastfeeding Practices: A Population Cohort Study from Finland.

BACKGROUND: Many environmental factors are known to hinder breastfeeding, yet the role of the family living environment in this regard is still poorly understood. OBJECTIVES: We used data from a large cohort to identify associations between neighborhood characteristics and breastfeeding behavior. METHODS: Our observational study included 11,038 children (0-2 years) from the Southwest Finland Birth Cohort. Participant information was obtained from the Medical Birth Register and municipal follow-up clinics. Neighborhood socioeconomic disadvantage, greenness, and population density were measured for a period of 5 years prior to childbirth within the residential neighborhood on a 250 × 250-m grid. Any breastfeeding and breastfeeding at 6 months were the primary outcomes. Binary logistic regression models were adjusted for maternal health and socioeconomic factors. RESULTS: Adjusted analyses suggest that mothers living in less populated areas were less likely to display any breastfeeding (OR: 0.46; 95% CI: 0.36, 0.59) and breastfeeding at 6 months (OR: 0.37; 95% CI: 0.34, 0.40). Mothers living in highly disadvantaged neighborhoods were less likely to display any breastfeeding if the neighborhood was less populated (OR: 0.54; 95% CI: 0.30, 0.95) but more likely to breastfeed at 6 months if the neighborhood was highly populated (OR: 3.74; 95% CI: 1.92, 7.29). Low greenness was associated with higher likelihood of any breastfeeding (OR: 3.82; 95% CI: 1.53, 9.55) and breastfeeding at 6 months (OR: 4.41; 95% CI: 3.44, 5). CONCLUSIONS: Our results suggest that neighborhood characteristics are associated with breastfeeding behavior in Finland. Unravelling breastfeeding decisions linked to the living environment could help identify interventions that will allow the appropriate support for all mothers and infants across different environmental challenges.

Sex-specific links between the social landscape and faecal glucocorticoid metabolites in semi-captive Asian elephants.

Although social behaviour is common in group-living mammals, our understanding of its mechanisms in long-lived animals is largely based on studies in human and non-human primates. There are health and fitness benefits associated with strong social ties, including increased life span, reproductive success, and lower disease risk, which are attributed to the proximate effects of lowered circulating glucocorticoid hormones. However, to deepen our understanding of health-social dynamics, we must explore species beyond the primate order. Here, using Asian elephants as a model species, we combine social data generated from semi-captive timber elephants in Myanmar with measurements of faecal glucocorticoid metabolite (FGM) concentrations. These data enable a "natural experiment" because individuals live in work groups with different demographic compositions. We examine sex-specific FGM concentrations for four different aspects of an individuals' social world: general sociality, work group size, sex ratio and the presence of immatures (<5 years) within the work group. Males experienced lower FGM concentrations when engaged in more social behaviours and residing in female-biased work groups. Surprisingly, females only exhibited lower FGM concentrations when residing with calves. Together, our findings highlight the importance of sociality on individual physiological function among elephants, which may have broad implications for the benefits of social interactions among mammals.

Intention to have a second child, family support and actual fertility behavior in current China: An evolutionary perspective.

OBJECTIVES: This study provides an evolutionary perspective to a classic topic in demography, that is, the discrepancy between reproductive intention and subsequent behavior, in the context of China's two-child policy. METHODS: We conduct an event history analysis of longitudinal data from the 2015 and 2018 waves of the Xi'an Fertility Survey (sample size = 321 followed one-child mothers) to test the hypotheses of how within-family support/conflict affects women's fertility behavior. RESULTS: Only 50% of positive intentions (i.e., intending to have a second child) led to another (live) birth within the 3-year interval; meanwhile, 15% of uncertain intentions and 5% of negative intentions resulted in a birth. Husband's and the firstborn's emotional support raised the hazard of second childbirth along maternal life course, which cannot be fully mediated by mother's fertility intention and thus, contributed to an intention-behavior gap. Husband's sibship size had dual effects on female childbearing behavior: A positive indirect effect mediated by fertility intention, but a negative direct effect presumably due to sibling competition for intergenerational support. Finally, after controlling for fertility intention, having a firstborn son was still associated significantly with a lower second-childbirth hazard, presumably due to son preference as well as concern over parental investment. CONCLUSIONS: Our study identifies a discrepancy between maternal fertility intention and realized childbearing, which was partly explained by (lack of) support from other (multiple) stakeholders in family reproduction.

Age related variation of health markers in Asian elephants.

Although senescence is often observed in the wild, its underlying mechanistic causes can rarely be studied alongside its consequences, because data on health, molecular and physiological measures of senescence are rare. Documenting how different age-related changes in health accelerate ageing at a mechanistic level is key if we are to better understand the ageing process. Nevertheless, very few studies, particularly on natural populations of long-lived animals, have investigated age-related variation in biological markers of health and sex differences therein. Using blood samples collected from semi-captive Asian elephants, we show that pronounced differences in haematology, blood chemistry, immune, and liver functions among age classes are also evident under natural conditions in this extremely long-lived mammal. We provide strong support that overall health declined with age, with progressive declines in immune and liver functions similarly in both males and females. These changes parallel those mainly observed to-date in humans and laboratory mammals, and suggest a certain ubiquity in the ageing patterns.