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.

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).

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.

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.

Handler familiarity helps to improve working performance during novel situations in semi-captive Asian elephants.

Working animals spend hours each day in close contact with humans and require training to understand commands and fulfil specific tasks. However, factors driving cooperation between humans and animals are still unclear, and novel situations may present challenges that have been little-studied to-date. We investigated factors driving cooperation between humans and animals in a working context through behavioural experiments with 52 working semi-captive Asian elephants. Human-managed Asian elephants constitute approximately a third of the remaining Asian elephants in the world, the majority of which live in their range countries working alongside traditional handlers. We investigated how the familiarity and experience of the handler as well as the elephant's age and sex affected their responses when asked to perform a basic task and to cross a novel surface. The results highlighted that when novelty is involved in a working context, an elephant's relationship length with their handler can affect their cooperation: elephants who had worked with their handler for over a year were more willing to cross the novel surface than those who had a shorter relationship with their handler. Older animals also tended to refuse to walk on the novel surface more but the sex did not affect their responses. Our study contributes much needed knowledge on human-working animal relationships which should be considered when adjusting training methods and working habits.

Within-litter covariance of allele-specific MHC heterozygosity, coccidian endoparasite load and growth is modulated by sibling differences in starting mass.

Although littermates in altricial mammals usually experience highly similar environmental conditions during early life, considerable differences in growth and health can emerge among them. In a study on subadults of a European rabbit (Oryctolagus cuniculus) population with low MHC polymorphism, we tested whether litter-sibling differences in endoparasitic coccidia load and body mass at the end of the vegetation period were associated with within-litter differences in starting body mass (measured around 2 weeks prior to weaning) and in immune-genetic (MHC class II DRB) constitution. We hypothesized that siblings with a lighter starting mass might be more susceptible to endoparasite infections and thus, negative effects of a more unfavourable MHC constitution might be particularly pronounced in such individuals. Within-litter comparisons revealed that animals with a lighter starting mass reached a relatively lower body mass in autumn. Furthermore, there were indications for an allele-specific heterozygote advantage, as animals with heterozygous combinations of the allele Orcu-DRB*4 had relatively lower hepatic coccidia loads than their littermates with certain homozygous allele combinations. Consistent with our hypothesis, significantly higher hepatic coccidia loads and tendentially lower autumn body masses in homozygous compared to heterozygous individuals for the allele Orcu-DRB*4 were evident in initially lighter but not in heavier siblings, suggesting synergistic effects between an unfavourable MHC constitution and a light starting mass. Taken together, these effects might lead to notable differences in fitness among litter siblings, as a low body mass and a high endoparasite burden are key factors limiting young rabbits' survival during winter.

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.

Faecal Glucocorticoid Metabolites and H/L Ratio are Related Markers of Stress in Semi-Captive Asian Timber Elephants.

Animals are kept in captivity for various reasons, but species with a slower pace of life may adapt to captive environments less easily, leading to welfare concerns and the need to assess stress reliably in order to develop effective interventions. Our aim was to assess welfare of semi-captive timber elephants from Myanmar by investigating the relationship between two physiological markers of stress commonly used as proxies for welfare, faecal glucocorticoid metabolite concentrations (FGM) and heterophil/lymphocyte ratios (H/L), and link these measures to changes in body condition (determined by body weight). We further assessed how robustly these two markers of stress performed in animals of different age or sex, or in different ecological contexts. We measured FGM concentrations and H/L ratios between 2016 and 2018 from 316 samples of 75 females and 49 males ranging in age from 4 to 68. We found a positive and consistent link between FGMs and H/L ratios in Asian elephants, irrespective of their sex, age, or ecological context. Our results will help to inform managers of (semi-) captive elephants about using heterophil/lymphocyte ratio data from blood smears on site as a potentially cheaper and faster alternative to determining stress than measuring faecal glucocorticoid metabolite concentrations in the laboratory.

Future Directions for Personality Research: Contributing New Insights to the Understanding of Animal Behavior.

As part of the European Conference on Behavioral Biology 2018, we organized a symposium entitled, "Animal personality: providing new insights into behavior?" The aims of this symposium were to address current research in the personality field, spanning both behavioral ecology and psychology, to highlight the future directions for this research, and to consider whether differential approaches to studying behavior contribute something new to the understanding of animal behavior. In this paper, we discuss the study of endocrinology and ontogeny in understanding how behavioral variation is generated and maintained, despite selection pressures assumed to reduce this variation. We consider the potential mechanisms that could link certain traits to fitness outcomes through longevity and cognition. We also address the role of individual differences in stress coping, mortality, and health risk, and how the study of these relationships could be applied to improve animal welfare. From the insights provided by these topics, we assert that studying individual differences through the lens of personality has provided new directions in behavioral research, and we encourage further research in these directions, across this interdisciplinary field.

Investigating changes within the handling system of the largest semi-captive population of Asian elephants.

The current extinction crisis leaves us increasingly reliant on captive populations to maintain vulnerable species. Approximately one third of Asian elephants (Elephas maximus) are living in semi-captive conditions in range countries. Their relationship with humans stretches back millennia, yet elephants have never been fully domesticated. We rely on the expertise of traditional handlers (mahouts) to manage these essentially wild animals, yet this profession may be threatened in the modern day. Here, we study the handling system of semi-captive timber elephants in Myanmar; the largest global semi-captive population (~5 000). We investigate how recent changes in Myanmar may have affected the keeping system and mahout-elephant interactions. Structured interviews investigated changes to mahout attitude and experience over the last two decades, as perceived by those who had worked in the industry for at least 10 years (n = 23) and as evaluated in current mahouts (n = 210), finding mahouts today are younger (median age 22yrs), less experienced (median experience 3yrs), and change elephants frequently, threatening traditional knowledge transfer. Mahout-elephant interactions manifested as 5 components ('job appreciation'; 'experience is necessary'; 'human-elephant interaction'; 'own knowledge'; 'elephant relationship'), according to Principal Components Analysis. Experienced mahouts and mahouts of bulls and younger elephants were more likely to agree that 'experience is necessary' to be a mahout. Mahouts with difficult elephants scored lower on 'human-elephant interaction' and a mahout's perception of their 'own knowledge' increased with more experience. Our finding of change in terms of mahout experience, age and commitment in the largest semi-captive elephant population suggests need for formal training and assessment of impacts on elephant welfare; these are findings applicable to thousands of elephants under similar management.

Males have more aggressive and less sociable personalities than females in semi-captive Asian elephants.

Personality, i.e. consistent between-individual differences in behaviour, has been documented in many species. Yet little is known about how males and females of long-lived, highly social species differ in their measures of personality structure. We investigated sex differences in the mean, variance, and covariance of three previously reported personality traits (Attentiveness, Sociability, Aggressiveness) in 150 female and 107 male Asian elephants (Elephas maximus) from a semi-captive population in Myanmar. These three personality traits were obtained by performing exploratory factor analysis on 28 behavioural items that had been rated by experienced elephant handlers. We found that males scored significantly higher on Aggressiveness and tended to score lower on Sociability than females. However, no sex difference was found in the mean scores of Attentiveness. Variances for the three personality traits did not differ between the sexes, suggesting that male and female elephants share the same range of personality variation. Likewise, trait covariances were similar between the sexes. While both sexes show complex sociality in the wild, female Asian elephants typically live in highly social family units, whereas male elephants' social bonds are weaker. Males usually form dominance ranks by aggressive interactions, especially during musth. Our results on a large sample of individuals living in their natural environment are thus in agreement with elephant life-histories and parallel the findings of sex differences in other long-lived highly social species with similar life-histories.

Evaluating the personality structure of semi-captive Asian elephants living in their natural habitat.

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.

Context dependency of baseline glucocorticoids as indicators of individual quality in a capital breeder.

Identifying markers of individual quality is a central goal of life-history theory and conservation biology. The 'corticosterone (CORT)-fitness hypothesis' postulates that low fitness signals impaired ability to cope with the environment, resulting in elevated baseline CORT levels. CORT can, however, be negatively, positively or neutrally related to fitness, depending on the context. In order to clarify this controversial issue, we elucidate the utility of using baseline CORT as a correlate of individual fitness in incubating female eiders across variable environments. An increase in serum CORT with decreasing body condition was evident in older, more experienced breeders, while increased clutch mass was associated with elevated serum CORT in females breeding late in the season. For faecal CORT, the expected negative association with body condition was observed only in early breeders. We found a strong increase in faecal CORT with increasing baseline body temperature, indicating the utility of body temperature as a complementary stress indicator. Females in good body condition had a lower baseline body temperature, but this effect was only observed on open islands, a harsher breeding habitat less buffered against weather variability. Females with higher reproductive investment also maintained a lower baseline body temperature. Nest success strongly decreased with increasing serum and faecal CORT concentrations, and individual stress hormone and body temperature profiles were repeatable over years. Although our data support the tenet that baseline CORT is negatively related to fitness, the complex context-dependent effects call for cautious interpretation of relationships between stress physiology and phenotypic quality.