Male Elephant Research Articles

Elephant Scar Prevalence in the Kasigau Wildlife Corridor, Kenya: Echoes of Human-Elephant Conflict.

Human-elephant conflict (HEC) compromises crop security and threatens elephant conservation. Most commonly, HEC manifests as crop-foraging as elephants modify natural foraging strategies to incorporate crops. Farmers may retaliate by frightening or harming elephants, leaving scars from inflicted wounds. We assessed the prevalence and distribution of scars on the bodies of African savanna elephants (Loxodonta africana) observed in the Kasigau Wildlife Corridor (KWC), part of the Greater Tsavo Ecosystem of Kenya, where conflict is prevalent. We surmised that scars on the body are largely a result of HEC as opposed to scars on the rump or head, which we attributed primarily to elephant-elephant conflict. We hypothesized that: (1) male elephants would have more scars than females; (2) older males would be more likely to have scars than younger males; and (3) most scars would be located on the bodies of elephants. We assessed scars from a photographic catalogue of elephants from the KWC. In line with our hypotheses, male elephants were more likely to have scars than females (32% of males compared to 6% of females); older males had significantly more scars than younger males (61% compared to 24%); and the majority of scars (89%) were located on the body. Scar presence may be useful as an animal-centered indicator to estimate the prevalence and demographic patterns of HEC.

Hormonal variation and temporal dynamics of musth in Asian elephants (Elephas maximus) are associated with age, body condition and the social environment.

The sustainability of endangered Asian elephants in human care is threatened in part by low breeding success and concerns over individual animal wellbeing. Male elephants have received less research attention compared to females, yet males deserve special consideration due to their unique reproductive biology (particularly the sexual state of "musth") and the complex interaction of physiological, environmental, and social pressures they face. We measured fecal androgen metabolites (FAMs), fecal glucocorticoid metabolites (FGMs), and fecal triiodothyronine metabolites (FT3s) collected weekly over approximately 12months from 26 male Asian elephants housed in zoos across the US, hypothesizing that FAM, FGM, and FT3 concentrations would be associated with temporal correlates of musth and would vary further with intrinsic (musth status, age, body condition) and extrinsic (social environment) factors. The duration of each musth episode was positively associated with exposure to male conspecifics and negatively associated with body condition. Further, elevated FAM concentrations were associated with social exposure, age, and body condition, and FGM concentrations also varied with age and body condition. FT3 concentrations were not associated with any factor we measured. We also identified periods of lower FAM concentration than confirmed musth episodes (but still higher than baseline FAM concentrations) that we termed "elevated FAM episodes." The durations of these episodes were negatively correlated with exposure to other male elephants. Together, these results provide evidence that hormone profiles (including those that are predicted to change around musth) vary significantly between male Asian elephants in a way that may be attributed to intrinsic and extrinsic factors. Studies like these serve to enhance the sustainability of ex-situ populations by providing wildlife managers with information to enhance the health, welfare, and reproduction of threatened species like Asian elephants.

Extreme dietary specialisation in adult male southern elephant seals: determining variation between individual trophic diets

Although dietary studies have provided important insights into the causes and ramifications of diet variation for the southern elephant seal (SES) (Mirounga leonina), adult males are comparatively underrepresented within that literature. Individual males can vary morphologically as well as behaviourally, leading to differences in their life history trajectories and outcomes. Therefore, to improve our understanding of the male diet, we sought to determine the degree of dietary variation between as well as within individuals from the West Antarctic Peninsula. Secondly, we investigated whether individual morphological traits, seasonality, and year influenced their dietary variation. Whiskers were sampled from 31 adult male seals and used to measure the bulk stable isotope nitrogen (δ15N). We sequentially segmented each whisker to create a time series of datapoints for each individual, allowing us to compare δ15N variation within each seal as well as assess variation between the seals. We then investigated the relationships between male dietary variation and body length, girth, season, and year. We found that adult male SESs maintained an extremely specialised diet. Variation between individuals was strongly correlated with their body size, with larger seals feeding higher up the trophic web. Interestingly, seasonality and year both influenced variation within the seals’ diets, but only year was seen to influence the variability between seals. We discuss the possible causes and ramifications of dietary specialisation for the SES and highlight the need for combined tracking and stable isotope investigations to improve our understanding of the ontogeny of the seals’ dietary specialisation.

Age-sex structure of drought-driven African elephant (Loxodonta africana) mortality in Hwange National Park, Zimbabwe

Age and sex determination of 101 lower jaws from drought-driven savanna elephant mortalities was done in Hwange National Park (HNP) using the Age Reference Line (ARL) method. We recorded an overall higher number of female elephant death compared to males. The age at death was skewed towards the young and very old individuals, with an overall gradual decrease in frequency into prime cohorts. Female elephant mortalities were more prevalent in young animals, nursing, and very old individuals. A similar mortality trend was observed for male elephants below the age of 22 years. We did not record any elephant between the age of 45–52.5 years for both sexes. We support the notion that the overall distribution of the female's mortality prevalence compared to males is aligned to and influenced by the social and developmental behaviour of savanna elephants. Our findings suggest a natural selection process at play, facilitated by resource limitation during the period of drought. This study provides evidence for the need to get back to the basic ecological principles and understanding of natural ecological processes in the conservation of key species. Although elephant mortalities due to drought represent a small proportion of the entire population in the study area, more work needs to be done to investigate the influence of drought, novel infections and anthropogenic drivers as they have potential to affect savanna elephant populations in the long term.

Standard Identification Certificate for Legal Legislation of a Unique Gene Pool of Thai Domestic Elephants Originating from a Male Elephant Contribution to Breeding

Illegal wildlife trade is a major threat to global biodiversity. Asian elephants (Elephas maximus) are highly valued by various cultures as religious symbols and tourist attractions, which has led to a high demand for captive elephants. Owing to the unviability of captive breeding programs, several captive elephant populations are maintained by illegally obtaining wild Asian elephants. Morbidity and mortality rates among captive populations are high, whereas reproduction is low. In this study, we examined the genetic diversity among elephants using microsatellite genotyping and mitochondrial D-loop sequences of three captive elephant populations. The study results showed very low nucleotide diversity D-loop sequences and high variations in microsatellite genotyping, with an extensive variation of the gene pool estimates from different populations. This suggests that the optimal male selection during breeding could aid in maintaining the genetic diversity among captive populations. Forward genetic simulation revealed a decreasing genetic diversity in the fixed state within 50 generations. However, largely different gene pools can be effectively used to infer original elephant sources; this would facilitate the development of an identification certificate integration with machine learning and image processing to prevent illegal legislation owing to registration fraud between wild and domestic elephants. Implementing the proposed approaches and recommendations would aid in the mitigation of the illegal capture and domestic trade of wild elephants in Thailand and contribute to the success of future conservation plans in the blueprint of sustainable development goals.

Density-dependent habitat selection varies between male and female African elephants

Habitat selection models are the basis of an increasing number of conservation and management programs. Decision-makers rely on accurate models to assess animal distribution over space and time, and to recommend suitable actions that can alleviate human-wildlife conflicts. Despite a rapidly growing number of field studies on habitat selection, there remains a paucity of empirical evidence that selection is a density-dependent process that can impact males and females differently. Based on 11 years of monitoring, we demonstrate that the response of African elephants (Loxodonta africana) to land-cover types varied with population size, and that density-dependent adjustments differed between sexes. Specifically, our longitudinal follow-up of GPS-collared elephants revealed that elephants gradually decrease their selection for open woodlands and forests, as the population increased and the availability of palatable browse species decreased. Both sexes – though males more strongly – increased their travel rate together with their relative probability of selection of roads for travel. Also, elephants displayed a density-dependent increase in their selection of infrastructures, a response that was stronger for males than females. The risk of human-elephant conflicts thus increased with population size, with males being particularly prone to be involved in such conflicts. Overall, we provide rare empirical evidence that density-dependence in fine-grain habitat selection can differ between sexes. This information can be critical to accurately forecast potential human-wildlife conflicts, and for taking targeted and effective conservation and management actions.

The ontogeny of at-sea behaviour in male southern elephant seals (Mirounga leonina) at Marion Island

Megafauna, such as southern elephant seals (SESs) (Mirounga leonina, Linn.), forage in diverse, seemingly limitless habitats. In pelagic settings, their behaviour is more likely to be limited by physiological ability and prey distribution, than physical barriers. For elephant seals, their rapid growth in body size corresponds to changing physiological abilities. These changes are most pronounced for male elephant seals. While most studies have compared male and female SESs in terms of changing body size and physiological ability, few studies have compared the influence of changing body size on behaviour of male SESs from the same population using long term satellite tracking data. We describe age-related differences in movement and diving behaviour for male SESs from Marion Island. We analysed satellite tracking data collected from 23 male SESs seals fitted with Sea Mammal Research Unit Satellite Relay Data Loggers at Marion Island between 2005 and 2011. Each dive was assigned a behavioural mode, either ‘searching’ or ‘transit’, using state-space modelling. We used mixed-effects models to quantify the influence of age and behavioural mode on dive duration, surface duration, dive depth and number of daily dives. Younger seals travelled significantly further from Marion Island and spent most of their dives in transit mode, whereas older seals stayed closer to the island but were in searching mode for most of their tracks. When searching, older seals dived more frequently, displayed longer dive and surface durations, and reached greater depths than younger seals. These differences in diving behaviour seem to reflect changing physiological ability. For male SESs, changes in physiological ability necessitate behavioural plasticity, which may be the key to survival and future breeding success. Robust males are more likely to breed and must therefore, maintain their body size by adapting to local oceanic conditions.

Predictors of testosterone in zoo-managed male African elephants (Loxodonta Africana).

Reproductive complications for both male and female zoo-managed African elephants (Loxodonta africana) contribute to the rapidly declining population. In zoo-managed bull elephants, few studies have explored the potential physiological, physical, social, and environmental factors that influence bull fertility, particularly, androgen production. Testosterone is the essential steroid hormone for male sexual maturation and inadequate concentrations can be detrimental for spermatogenesis. Testosterone, fecal glucocorticoid metabolites, leptin, glucose, insulin, and triglycerides were analyzed from weekly fecal and blood serum samples taken over 6 months from six zoo-managed African elephant bulls (10-19 years of age). Testosterone levels were compared to endocrine factors, weekly social and environmental variables, daily musth signs, and body condition scores (BCS). The glucose-to-insulin ratio (G:I) was the only physiological biomarker found to be positively associated with testosterone. Predictive physical variables included Musth Score (+) and Moderate Exercise (+). Bulls with BCSsignifying overweight (BCS 4) had lower testosterone (36.6 ± 1.6 ng/g fecal extraction [FE]) than bulls with healthyBCS 3; 51.2 ± 4.9 ng/g FE). Numerous social variables influenced testosterone concentrations, including Total Contact Day (+), Female Interaction Day (+), Indirect Contact Day (+), Indirect Contact Night (+) and Total No Contact (-). Both percentage of Time Outdoor and Time Mixed positively influenced testosterone, whereas testosterone decreased for percentage of Time Indoors. Each additional daily browse opportunity increased testosterone by approximately 7 ng/g FE. In managed care, the emphasis should be placed on optimizing these predictors of testosterone production to promote bull reproductive health.

“The male elephant in the room”: a qualitative evidence synthesis exploring male experiences of eating disorders

BackgroundMen are believed to be underrepresented in eating disorder services; there are many reasons presented to account for this such as a lack of recognition and detection. Due to the physical and psychological complexity of an eating disorder it is important to understand this underrepresentation. This qualitative evidence synthesis aimed to explore the literature relating to men’s experiences of an eating disorder, in order to synthesise the findings and offer a more coherent understanding.MethodA systematic search of the literature was undertaken. Inclusion and exclusion criteria were applied to the findings resulting in 14 papers deemed suitable for inclusion. A Meta-Ethnographic approach to synthesising the data of the 89 participants was undertaken.ResultsFour themes were identified: ‘Societal Construction of the Perfect Male’; ‘Striving to Maintain a Masculine Identity’; ‘The Interconnectedness of Control and Self-Worth’, and ‘The Hidden Man’. There appeared to be an underlying concept relating to the conflict of being a man, with what is perceived to be a ‘woman’s illness’ and how this challenged the men’s experiences of masculinity.ConclusionsBeing a man with an eating disorder conflicted with societal norms, exacerbating their experience of having an eating disorder.. To resolve this, gendered norms need to shift, at societal level as well as considering how best to improve understanding and recognition of men with an eating disorder at the first point of help seeking.

Behaviour and Welfare Impacts of Releasing Elephants from Overnight Tethers: A Zimbabwean Case Study.

Simple SummaryIn the southern African elephant tourism industry, many elephants are routinely chained or tethered for prolonged periods of time, particularly overnight. There are many negative implications of such restrictive management on elephant welfare. In this pilot study, a group of four male semi-captive African elephants at a tourist facility in Zimbabwe were being taken off overnight tethers and put into small pens, as a strategy to improve elephant welfare. Behavioural data were collected when elephants were on tethers, and approximately four weeks and eight weeks after removal of tethers. Behavioural changes were noted, after removal of tethers, which were indicative of improved welfare (e.g., increased lying rest, increased positive social behaviour and reduced abnormal repetitive behaviours). Importantly, there were no significant increases in aggression (either to their human handlers or to other elephants) following this change in management. To improve elephant welfare in southern African tourism facilities, we strongly advocate management practices which enable greater choice and freedom of movement overnight, which includes ceasing the use of overnight tethers and provision of opportunities for physical interaction with other elephants overnight. Within the southern African elephant tourism industry, chaining or tethering elephants is still a relatively routine practice, despite the known negative impacts. Cited reasons for chaining include fear of aggressive interactions between elephants when handlers are absent, or a general increase in expression of aggressive behaviours (both to other elephants and to their human handlers). In Zimbabwe, concerns expressed include the danger of elephants escaping and entering human-inhabited areas. Four male semi-captive elephants at a Zimbabwe tourist facility were taken off overnight (~12 h) tethers and were placed in small pens (‘bomas’), approximate sizes from 110 m2 to 310 m2), as part of a strategy to improve elephant welfare. Behavioural data were collected from overnight videos from December 2019 to March 2020, between 18:00 to 06:00, using focal, instantaneous sampling (5-min interval). Data were collected for three nights at three time periods: (i) Tethered; (ii) approximately four weeks post-release; (iii) approximately eight weeks post-release. Behavioural change over these time points was analysed using general linear models with quasibinomial error structures. Behavioural changes indicative of improved welfare were observed following these management changes, and no significant increases in aggression were observed either between elephants, or towards their human handlers. Proportion of time engaging in lying rest was higher in the first month after release from tethering (mean ± SD, 50 ± 14%) than when elephants were tethered (20 ± 18%) (p < 0.05). Additionally, although not statistically significant, stereotypies were reduced when elephants were no longer tethered (4 ± 6% observations tethered compared to 2 ± 2% off tethers), and positive social behaviour also increased (1 ± 1% on tethers, 2 ± 2% off tethers), with the greatest improvements seen in the pair-housed elephants. To improve elephant welfare in southern African tourism facilities we strongly advocate that less restrictive management practices which enable greater choice and freedom of movement overnight are implemented.

Cephalopod diet of juvenile male southern elephant seals Mirounga leonina at Marion Island, South Indian Ocean

Cephalopod diet of juvenile male southern elephant seals Mirounga leonina at Marion Island, South Indian Ocean

A novel technique for aging male African elephants (Loxodonta africana) using craniofacial photogrammetry and geometric morphometrics

A novel technique for aging male African elephants (Loxodonta africana) using craniofacial photogrammetry and geometric morphometrics

Characterization of Longitudinal Testosterone, Cortisol, and Musth in Male Asian Elephants (Elephas maximus), Effects of Aging, and Adrenal Responses to Social Changes and Health Events.

Simple SummaryThe conservation of endangered species and sustainability of managed populations requires good health and welfare of individuals. Male elephants experience a biological phenomenon called “musth”, characterized by a suite of physiological and behavioral changes which serve to facilitate copulation between the sexes, but can also pose unique challenges under human care. This study characterized longitudinal patterns of circulating testosterone and cortisol in relation to musth in four adult Asian elephant bulls and assessed age-related effects on musth activity and adrenal responses to social changes and clinical health events. All bulls exhibited regular annual musth cycles, and there was no clear evidence of chronically elevated cortisol secretion outside of musth. Circulating cortisol covaried positively with testosterone and musth, highlighting intrinsic patterns that should be considered when evaluating the impact of changes on adrenal glucocorticoid activity. Testosterone decreased with age in sexually mature bulls, whereas age-related changes in cortisol varied across individuals, with the three older bulls showing the greatest rate of change during musth. Changes in cortisol were associated with treatment for Mycobacterium tuberculosis in two bulls, but not with activation of disease. In contrast to physiological factors, there was no evidence that social changes involving male herdmates impacted adrenal glucocorticoid activity in the short term. This study highlights the importance of longitudinal hormone monitoring to track changes in physiological function and identify factors that may influence welfare, which is important for making more informed decisions on how to manage male elephants under varying degrees of human care.The conservation of endangered species and sustainability of managed populations requires considerations to ensure the health and welfare of individuals. Male elephants experience a biological phenomenon called “musth”, which is characterized by increased testosterone production, temporal gland secretion and urine dribbling, heightened aggression and sexual behavior, and therefore can pose unique challenges for human safety and animal welfare. This study characterized longitudinal (9 to 22 years) patterns of circulating testosterone and cortisol in relation to musth in four adult Asian elephant bulls spanning ages from 12 to 54 years. Age-related effects on musth activity and adrenal responses to social changes and clinical health events were also examined. All bulls exhibited regular annual musth cycles. Circulating cortisol covaried positively with testosterone and musth, highlighting intrinsic patterns that should be considered when evaluating the impact of social, health, and environmental changes on adrenal glucocorticoid activity. Except for an end-of-life cortisol increase in one bull, there was no clear evidence of chronically elevated cortisol secretion outside of musth in any individual. Testosterone decreased with age in sexually mature bulls, whereas age-related changes in cortisol varied across individuals, with the three older bulls showing the greatest rate of change during musth versus inter-musth periods. In contrast to physiological factors, there was no evidence of social factors, such as addition of a new male and death of male herdmates, impacting adrenal glucocorticoid activity in these bulls in the short term. Changes in cortisol were associated with treatment for Mycobacterium tuberculosis (M. tb) in two bulls, increasing after start of treatment and decreasing with cessation of treatment, but were not clearly associated with activation of disease. This study highlights the importance of longitudinal hormone monitoring to track changes in physiological function and responses to social, health, and environmental change in elephant bulls, which is important for making more informed decisions on how to manage male elephants under varying degrees of human care to ensure welfare and safety.

Social Behavior and Group Formation in Male Asian Elephants (Elephas maximus): The Effects of Age and Musth in Wild and Zoo-Housed Animals.

Simple SummaryThe conservation of wild and zoo-housed Asian elephants partly depends on developing our understanding of male elephant social behavior. Once thought to be solitary, we now know that male elephants can display rich social exchanges with both males and females. However, these interactions are expected to change with age and around the sexually active state of “musth.” We used behavioral data from wild and zoo-housed elephant populations to investigate how age, musth, and environmental factors influence how male elephants socialize and associate with group members. In Sri Lanka, only male elephants of older age (>20 years) exhibited signs of musth, but even some of the youngest zoo-housed elephants underwent musth (as young as 11 years). Additionally, we found that age and musth status predicted whether a wild male elephant associated with females, other males or was alone, as well as the number of conspecifics (males and females) in the same group with which a male was observed. Finally, rates of aggression, prosocial behavior (affiliative behaviors that promote positive social bonds between elephants), and submissive behavior exhibited by wild and zoo-housed male elephants were associated with age, musth status, the number of elephants presented, and group type (all-male or mixed sex). These results provide motivation for future studies of social behavior in male Asian elephants, as they will contribute to the reproduction and conservation of this endangered species.Asian elephants are endangered, and the long-term viability of the species depends on integrative approaches to address the sustainability of in-situ and ex-situ populations. Growing evidence shows that male elephants exhibit extensive and flexible social behavior that rivals the complexity of that of females. Male elephant sociality is expected to change dramatically around the unique sexual state of musth. However, data related to male Asian elephant sociality is lacking. Here, we conducted complementary observations in Wasgamuwa National Park, Sri Lanka, and North American zoos of male Asian elephant social behavior. Age and musth status, along with other factors, were associated with variation in social behavior and group formation of males. In wild male elephants, both musth status and age impacted elephant associations within all-male and mixed-sex groups: non-musth elephants were generally sighted less often in mixed-sex groups as they aged, while the inverse occurred with musth elephants. Musth status interacted with age to predict the number of conspecifics with which a wild male elephant associated: younger males were observed with more females during non-musth (but the opposite was true during musth), and male elephants between 20 and 30 years were observed with the highest number of male conspecifics except during musth. Finally, we found variation in aggression, prosocial behavior, and submissive behavior was influenced by intrinsic (age and musth status) and extrinsic factors (group size and type) in similar ways in both populations; prosocial behavior was most common and was influenced by the number of conspecifics present (both populations), and age, group type, and musth status (zoo population), while aggression was rare, especially among older elephants. We suggest that longitudinal studies of this threatened species will be particularly helpful to promote the reproduction and conservation of Asian elephants in in-situ and ex-situ environments.

Male African Elephant (Loxodonta africana) Behavioral Responses to Estrous Call Playbacks May Inform Conservation Management Tools.

Simple SummaryDuring annual periods of heightened sexual activity (musth), male African elephants expend a significant amount of energy communicating their reproductive status, as well as searching and competing for potential mates. To locate females, musth males may risk venturing outside protected areas and into landscapes shared with humans where conflict between wildlife and people can occur. Adverse interactions between elephants and people can be detrimental to human livelihoods, resulting in negative attitudes towards elephants and, in some cases, retaliatory killings. Interactions with aggressive musth males can also be life-threatening to community members who attempt to confront them. Mitigation strategies that effectively target the reproductive motivations of musth males may offer solutions as the human–elephant interface continues to expand. In this study, we build on earlier research showing that playbacks of female elephant reproductive calls, (i.e., estrous rumbles) can change the movement trajectory and behavior of male elephants in Etosha National Park, Namibia. Individuals belonging to three male groups were opportunistically subjected to playback experiments and evaluated based on their reaction intensity. Our results demonstrate that mature musth adults are more likely to change directions and approach the source of a female’s estrous call than mature, sexually-inactive adult elephants. We also show that post-dispersal young males that were not in musth also respond strongly to the stimulus. These findings support further exploration of mitigation solutions that incorporate elephant behavior, reproductive status, and context-specific vocalizations.Driven by reproductive motives, male African elephants (Loxodonta africana) in musth often expand their home ranges to locate estrous females. This extended range, coupled with heightened aggression often observed in musth males, can be particularly problematic in regions where human-modified landscapes and elephant territories increasingly overlap. Several mitigation tools have been tested to resolve a wide range of human–elephant conflicts with varying degrees of success due to geographical disparities and habituation. We present findings on the potential application of estrous call playbacks in manipulating the behavior and movement of male elephants non-invasively, particularly mature musth adults and younger post-dispersal males, in Etosha National Park. Estrous vocalizations were presented across 26 experimental trials to mature musth adults (n = 5), mature non-musth adults (n = 6), and non-musth males belonging to younger, post-dispersal age classes (n = 8), with behavioral responses scored on a gradient scale from 0–1. Both mature musth adults and younger non-musth elephants were significantly more likely to respond with the highest intensity by approaching the acoustic source compared to mature non-musth adults that avoided the call. However, younger males tested in the presence of an older, higher-ranking male tended to react with a lower intensity than those tested alone. This result likely demonstrates the influence of social hierarchy and associations on male elephant behavior. We also observed a significant increase in physiological response, measured by defecation rate, across all male groups in response to the estrous call playbacks. Our findings suggest that using estrous calls as acoustic deterrents may effectively and non-invasively aid in reducing tension at the human–elephant interface, depending on the age, social context, and reproductive status of the male elephant.

Behavioral characterization of musth in Asian elephants (Elephas maximus): Defining progressive stages of male sexual behavior in in-situ and ex-situ populations

Complementary studies of wild and zoo-housed animals offer insight into behavioral variation across a range of conditions including the context under which various behaviors evolved in natural settings. This information can be used to improve the sustainability of in-situ and ex-situ populations and enhance the well-being of individuals. Managed ex-situ populations are critical to the long-term existence of Asian elephants, yet relatively little is known about male reproductive behavior compared to females. Male elephants undergo a unique sexual state called “musth” that further complicates in-situ and ex-situ management strategies. The ability to manage musth males to enhance breeding success and overall wellness of elephants is dependent upon better understanding how intrinsic and extrinsic factors influence male behavioral variation around musth. Here, we observed 62 free-ranging male Asian elephants in Sri Lanka and compared their behavior to observations from 26 elephants managed in facilities around the US. We hypothesized that musth is associated with significant behavioral changes that can be used to define distinct stages in the progression of musth. During observations, we quantified environmental variables and recorded musth status of each focal elephant using visual indicators (temporal gland secretions and urine dribbling). We showed that musth’s behavioral correlates (including changes in locomotion, foraging, alertness, and chemosensory behavior) were remarkably similar in wild and zoo-housed elephants. We also found that behavioral variation around musth was also associated with intrinsic (e.g., musth stage, age) and extrinsic factors (e.g., space availability, temperature) in zoo-housed, but not wild, elephants, indicating that musth is potentially plastic in changing environments. As musth progressed, we noted distinct behavioral signatures that define four stages of sexual activity in male elephants: non-musth, early musth, full musth, and post-musth. Finally, although we did not observe significant changes in overall social behavior (including aggression) during musth, we found that elephants increased the frequency with which they displayed certain behaviors associated with communication (e.g., alertness, chemosensory behavior, ear-flapping) in both populations. Together, these results indicate the significant behavioral changes that occur during musth in wild and zoo-housed elephants, and that musth progresses in distinct behavioral stages that can be easily distinguished by visual indicators. Studies like these serve to provide wildlife managers with information about a species’ unique, evolved behavioral strategies and how these seemingly fixed behaviors may be influenced by intrinsic and extrinsic factors in predictable ways.

Challenges to Elephant Connectivity From Border Fences in the World's Largest Transfrontier Conservation Area

To more effectively protect biodiversity and promote sustainable development, transfrontier conservation areas (TFCAs) aim to enhance wildlife flows across national borders. This is true of the world's largest terrestrial TFCA, the Kavango-Zambezi (KAZA), home to half of Africa's savannah elephants that move across five countries in a mixed-use landscape. We used GPS tracking data from &amp;gt;100 collared elephants to evaluate how fences between Namibia and Botswana impact transboundary connectivity in KAZA. For female elephants these fences formed an impenetrable boundary, with no exchange between animals collared in Botswana and those collared in Namibia. Male elephants did cross border fences, although they remained a partial boundary, with 7 of 21 males accounting for most crossings. Our results suggest a review of fence alignment and de-commissioning of some fencing separating Namibia and Botswana, combined with increased support for fence-free interventions that reduce wildlife-livestock interactions, should be considered to meet the objectives of KAZA.

Physical and behavioral indicators associated with hormonal changes during musth in zoo-housed and free-ranging Asian elephants (Elephas maximus)

In-situ and ex-situ Asian elephant populations are threatened with extinction, and male elephants pose unique challenges to long-term sustainability. The heightened sexual state of “musth” is accompanied by a suite of physical, behavioral and physiological changes. Furthermore, musth is unique to male elephants and requires special consideration when developing short- and long-term management strategies for elephants in the wild and in human care. The purpose of this study was to identify associations between fecal hormone metabolites [fecal androgen metabolites, FAM; fecal glucocorticoid metabolites, FGM; and fecal triiodothyronine (T3) metabolites, FT3] and visible musth indicators [temporal gland secretions (TGS) and urine dribbling (UD)], and behavioral changes around musth. From fecal samples collected non-invasively from wild elephants in Wasgamuwa National Park, Sri Lanka, and zoo-housed elephants in the United States, we hypothesized that (1) TGS and/or UD would be associated with changes in FAM, FGM, and/or FT3 concentrations; (2) variation in fecal hormone metabolites would be associated with increased locomotion and chemosensory behavior, and decreased foraging; and (3) relationships we identified would be similar between wild and zoo-housed elephants. We found that FAM concentrations changed significantly with TGS and UD activity in both wild and zoo elephants. Further while FGM concentrations were higher with increased TGS and UD in zoo elephants, the opposite pattern occurred in wild elephants. We did not identify substantial change in FT3 concentrations with TGS/UD activity. Behavioral changes in zoo elephants were significantly associated with FAM concentration as predicted, but these relationships were more difficult to identify in wild elephants due to lower sample availability. Further, FGM concentration was directly related to time spent locomoting in zoo elephants, but no other apparent association existed between FGM concentration with other behaviors in zoo elephants, or in any behaviors in wild elephants. Likewise, we did not report associations between FT3 and any behaviors we measured. This study contributes to our understanding of the complex response patterns that male Asian elephants exhibit around musth, and it provides another example of complementary in-situ–ex-situ research that can be directly applied to improve the well-being of elephants and other wildlife.

Determining the Dietary Preferences of Wild Asian Elephants (Elephas maximus) in Taman Negara National Park, Malaysia Based on Sex and Age using trnL DNA Metabarcoding Analysis.

The world's largest terrestrial mammal, Asian elephants, are known to have enormous feeding needs. Several factors such as season, sex, age, and daily activities influence the amount of food required by an individual. Generally, captive elephants have a limited choice of food on a daily basis compared with that of elephants in the wild. Elephants in captivity are fed according to a prepared feeding schedule, whereas wild elephants are free to choose the type of plants that they consume in their natural habitat. In the past, ecological observations have been widely used to determine the diet of wild elephants. However, the molecular approach has never been carried out. In the present study, we aimed to; 1) identify the plant diet of wild Asian elephants in Taman Negara National Park (TNNP) according to their sex and age using high-throughput DNA metabarcoding; and 2) determine the dietary formulation of captive elephants based on the generated plant metabarcoding database. DNA was extracted from 24 individual fecal samples collected using noninvasive sampling techniques from TNNP and the National Elephant Conservation Centre (NECC) Kuala Gandah. Seven pooled samples from male adult, female adult, male subadult, female subadult, male juvenile, female juvenile, and captive elephants were amplified and sequenced targeting the trnL region (50-150 base pairs). The CLC Genomic Workbench and PAST 4.02 software were used for data analysis. In total, 24 orders, 41 families, 233 genera, and 306 species of plants were successfully detected in the diet of the Asian elephants. The most abundant plant genera consumed were Sporobolus (21.88%), Musa (21.48%), and Ficus (10.80%). Plant variation was lower in samples from male elephants than in those from female elephants. The plant species identified were correlated with the nutrient benefits required by elephants. Adults and subadults consumed more plant species than were consumed by juvenile elephants. However, there was no significant difference between ages and sexes. The findings of this study can be used as guidance by the Department of Wildlife and National Parks for the management of captive elephants, especially in NECC Kuala Gandah.

Diffuse large B cell lymphoma and a novel gammaherpesvirus in northern elephant seals Mirounga angustirostris.

Two emaciated male northern elephant seal (NES) Mirounga angustirostris pups were admitted to The Marine Mammal Center (Sausalito, California, USA) and treated for malnutrition. Complete blood counts showed a progressive moderate to marked leukocytosis characterized by a predominance of large monomorphic mononuclear cells of probable lymphoid origin, frequently with flower-shaped nuclei. Both seals were euthanized due to suspected lymphoid neoplasia. At necropsy, most lymph nodes in both pups were markedly enlarged, some with distinct white nodules, the spleens were diffusely enlarged, and the intestinal mucosae were thickened. Histopathologic features consistent with disseminated large cell lymphoma were identified to varying degrees of severity in lymph nodes, bone marrow, liver, tonsils, spleen, liver, intestines, kidneys, lower urinary tract, and several other organs. Immunohistochemical staining of neoplastic cells was most consistent with B lymphocyte origin, with most cells staining positively for Pax 5 and CD20 with admixed small CD3-positive T lymphocytes and CD204-positive macrophages. PCR and sequencing identified a novel gammaherpesvirus, herein called miroungine gammaherpesvirus 3, from affected tissues. This virus is in a clade outside of named genera that utilize hosts in the suborder Caniformia. The present study is the first description of diffuse large B cell lymphoma with leukemic manifestation and concomitant detection of a novel gammaherpesvirus in free-living NESs. Further research regarding the prevalence of this new gammaherpesvirus and its associated pathogenesis in this species is indicated.