Social behavior and group formation in male Asian Elephants (Elephas maximus): the effects of age and musth in wild and zoo-housed animals
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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.
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In 2017, males made up only 21% of the Asian elephant population in North America but will eventually approach 50%.
“In 2017, males made up only 21% of the Asian elephant population in North America but will eventually approach 50%”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as LaDue et al., 2022b
The Conclusions state that males comprised 27% of the North American Asian elephant population in 2017, not 21%. They also state that the proportion will eventually approach 50%. The supplied passage has no page number. (LLM, conf=0.93)
Experimental — accuracy not yet measuredAffiliation was the most common type of social behavior observed in the wild (72.5%) and in all-male groups of Asian elephants housed in zoos (68.5%).
“Affiliation was also the most common type of social behavior observed in the wild (72.5%) and in all-male groups of Asian elephants housed in zoos (68.5%)”
From Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR (2025) · cites this paper as [56]
The Results (§3.2; page not provided) identify prosocial behavior as the most frequent social behavior and report 72.5% for wild elephants and 68.5% for zoo-housed elephants. The 68.5% figure applies to zoo-housed elephants overall, not specifically to all-male zoo groups; the paper says prosocial behavior was more common in mixed-sex zoo groups. (LLM, conf=0.90)
Experimental — accuracy not yet measuredIncreasing evidence indicates that male elephants display extensive social behaviors that rival the complexity of those of females.
“Increasing evidence indicates that male elephants display extensive social behaviors that rival the complexity of that of females”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as LaDue et al., 2022b
The study animals exhibited behavior consistent with other populations.
“indicating that the study animals exhibited behavior consistent with other populations [56].”
From Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR (2025) · cites this paper as [56]
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BLOOD ANALYTE CHANGES ASSOCIATED WITH MUSTH IN TWO CAPTIVE ASIAN ELEPHANT (ELEPHAS MAXIMUS) BULLS
Minor, R. L., Edes, A. N., Kanarr, M. L., Cannizzo, S. A., Griffioen, J. A., Stacy, N. I., Cray, C., Paris, S., Brown, J. L., & Rainwater, K. L. (2026) · Journal of Zoo and Wildlife Medicine · doi:10.1638/2025-0064
Pérez, A. B. L., Goldenberg, S. Z., Brown, J. L., & Khonmee, J. (2026) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2026.107067
Gartland, K. N., Bovee, E., Reed, A. N., & Fuller, G. A. (2026) · American Journal of Primatology · doi:10.1002/ajp.70183
- Authors
- LaDue, C. A., Vandercone, R. P. G., Kiso, W. K., & Freeman, E. W.
- Affiliations
- George Mason University (US)
- Rajarata University of Sri Lanka (LK)
- White Oak Conservation Foundation, Yulee, FL 32097, USA
- Corresponding author
- LaDue, Chase A.
- Venue
- Animals
- Publisher
- MDPI AG
- Published
- 2022-05-08
- Volume
- 12
- Issue
- 9
- Pages
- 1215
- ISSN
- 2076-2615 (ISSN-L)
- PMC
- PMC9100748
- OpenAlex
- W4229335296
- Cited by
- 26
- References
- 75
- Citations by year
- 2026: 4 · 2025: 10 · 2024: 4 · 2023: 6 · 2022: 2
- FWCI
- 5.20
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Animals (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- Funders
- American Society of Mammalogists
- Animal Behavior Society
- Asian Elephant Support
- Cosmos Club Foundation
- Elephant Managers Association
- The Explorers Club
- Feld Entertainment, Inc.
- Ringling Bros
- Center for the Study and Conservation of the Asian Elephant
- Fulbright US Student Program
- George Mason University (Department of Biology, Office of the Provost, and School of Integrative Studies)
- IDEA WILD
- International Elephant Foundation
- National Geographic Society
- Oklahoma City Zoo Conservation Action Now
- Saint Louis Zoo WildCare Institute
- Virginia Academy of Science
- Topics
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
- Keywords
- Asian elephant, male social behavior, musth, group formation, age effects, aggression, prosocial behavior, submissiveness, in situ and ex situ conservation, group size
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