Male elephant management in AZA institutions: Current status and priorities for the future.

Asian elephant (Elephas maximus) and African savanna elephant (Loxodonta africana) populations collectively managed by ex-situ facilities accredited by the Association of Zoos and Aquariums (AZA) face sustainability challenges. Among the priorities to strengthen animal wellbeing and population sustainability is male elephant management. We conducted a survey of AZA facilities currently housing male elephants to assess the status, challenges, and priorities in three areas of male elephant management: musth, socialization, and semen collection. Surveys were administered to elephant care teams at AZA-accredited institutions between November 2022 and February 2023, and we received responses from 34 institutions (91.9% of AZA-accredited facilities holding adult male elephants), housing 32 adult male Asians and 26 adult male Africans. Most facilities prioritized breeding and male socialization over musth management and semen collection (although most facilities acknowledged that all these efforts are important), citing leadership support and staffing as most important to achieve male management goals. Behaviors most commonly accompanying musth included reduced appetite, difficulty training or shifting, human-directed aggression, and interest in females. Musth timing was variable between males and facilities. Most males were well-socialized with females and/or other males, though elephant compatibility and facility design were limiting factors in managing socialization. Although 60.6% of facilities collected semen or were training for semen collection, very few male elephants could reliably provide viable semen samples, challenging assisted reproductive efforts that could bolster population sustainability in both species. Together, our results provide a better understanding of the state of male elephant management, offering specific areas deserving of research and development to enhance wellbeing and sustainability.

Asian elephants distinguish sexual status and identity of unfamiliar elephants using urinary odours.

Despite the ubiquity of odours in mammals, few studies have documented the natural olfactory abilities of many 'non-model' species such as the Asian elephant. As Asian elephants are endangered, we may apply odours to more effectively manage threatened populations. We implemented a habituation-discrimination paradigm for the first time in Asian elephants to test the ability of elephants to discriminate between unfamiliar male elephant urine, hypothesizing that elephants would successfully distinguish non-musth from musth urine and also distinguish identity between two closely related individuals. We conducted two bioassay series, exposing three female and three male zoo-housed elephants to the same urine sample (non-musth urine in the first series, and urine from an unfamiliar individual in the second) over 5 days. On the sixth day, we simultaneously presented each elephant with a novel sample (either musth urine or urine from a second unfamiliar individual) alongside the habituated urine sample, comparing rates of chemosensory response to each sample to indicate discrimination. All elephants successfully discriminated non-musth from musth urine, and also urine from two unfamiliar half-brothers. Our results further demonstrate the remarkable olfactory abilities of elephants with promising implications for conservation and management.

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 12 months 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.

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.

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

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.

The Chemical Ecology of Elephants: 21st Century Additions to Our Understanding and Future Outlooks.

Chemical signals are the oldest and most ubiquitous means of mediating intra- and interspecific interactions. The three extant species of elephants, the Asian elephant and the two African species, savanna and forest share sociobiological patterns in which chemical signals play a vital role. Elephants emit secretions and excretions and display behaviors that reveal the importance of odors in their interactions. In this review, we begin with a brief introduction of research in elephant chemical ecology leading up to the 21st century, and then we summarize the body of work that has built upon it and occurred in the last c. 20 years. The 21st century has expanded our understanding on elephant chemical ecology, revealing their use of odors to detect potential threats and make dietary choices. Furthermore, complementary in situ and ex situ studies have allowed the careful observations of captive elephants to be extended to fieldwork involving their wild counterparts. While important advances have been made in the 21st century, further work should investigate the roles of chemical signaling in elephants and how these signals interact with other sensory modalities. All three elephant species are threatened with extinction, and we suggest that chemical ecology can be applied for targeted conservation efforts.

The behavioral effects of exhibit size versus complexity in African elephants: A potential solution for smaller spaces.

Population-level analyses suggest that habitat complexity, but not necessarily space availability, has important welfare outcomes for elephants in human care. At the Dallas Zoo, the opening of a new exhibit complex allowed us to measure the behavior of two female African elephants across three treatments to evaluate the independent effects of complexity and space. Preoccupancy observations were conducted in the elephants' older exhibit, which consisted of a smaller, more simple yard (630 m2 ). Subsequent postoccupancy observations measured behavior in two different spaces in the new exhibit: a larger, complex yard (15,000 m2 ), and a smaller, but complex yard (1,520 m2 ). The elephants' overall activity levels were greater in complex habitats, regardless of their size. Similar effects of habitat complexity oversize were observed with greater rates of foraging and lower rates of being stationary. Furthermore, elephants were out of view of visitors significantly more in the small, simple yard compared to either of the more complex habitats. However, exhibit size affected the incidence of stereotypic behavior (with lower rates of stereotypy in the larger exhibit compared to the smaller yards) and investigatory behavior (elephants investigated their environments more with increasing size and complexity). Behavioral diversity also increased with exhibit size and complexity. These results indicate that space availability alone is not sufficient to enhance the behavioral welfare of zoo elephants. Therefore, facilities with limited space can still encourage species-appropriate behaviors and improved welfare for the elephants in their care by converting a small, simple area into a more complex habitat.