Physical, Physiological, and Behavioral Correlates of Musth in Captive African Elephants ( Loxodonta africana )
Abstract
Although musth in male African elephants (Loxodonta africana) is known to be associated with increased aggressiveness, urine dribbling (UD), temporal gland secretion (TGS), and elevated androgens, the temporal relationship between these changes has not been examined. Here, we describe the pattern of musth-related characteristics in 14 captive elephant bulls by combining long-term observations of physical and behavioral changes with physiological data on testicular and adrenal function. The length of musth periods was highly variable but according to our data set not related to age. Our data also confirm that musth is associated with elevated androgens and, in this respect, show that TGS and UD are downstream effects of this elevation, with TGS responding earlier and to lower androgen levels than UD. Because the majority of musth periods were associated with a decrease in glucocorticoid levels, our data also indicate that musth does not represent a physiological stress mediated by the hypothalamic-pituitary-adrenal axis. Furthermore, we demonstrate that the occurrence of musth is associated with increased aggression and that this is presumably androgen mediated because aggressive males had higher androgen levels. Collectively, the information generated contributes to a better understanding of what characterizes and initiates musth in captive African elephants and provides a basis for further studies designed to examine in more detail the factors regulating the intensity and duration of musth.
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This heightened hormonal reactivity may be a physiological mechanism that supports older males' ability to gain or maintain high-status positions within the dynamic social hierarchy of male elephants.
“This heightened hormonal reactivity may be a physiological mechanism that supports their ability to gain or maintain high-status positions within the dynamic social hierarchy of male elephants [60,71].”
From Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR (2025) · cites this paper as [60]
Musth is associated with increased androgen production.
“and is associated with increased androgen production [60,61].”
From Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR (2025) · cites this paper as [60]
Cites
A Simple Sequentially Rejective Multiple Test Procedure
(1979) · Scandinavian Journal of Statistics · doi:10.2307/4615733 · Cited by 21,526 · see citing works
Observational study of behavior: sampling methods
Altmann, J. (1974) · Behaviour · doi:10.1163/156853974x00534 · Cited by 13,891 · see citing works
Rutting Behavior in African Elephants: the Phenomenon of Musth
Poole, J. H. (1987) · Behaviour · doi:10.1163/156853986x00171 · Cited by 258 · see citing works
Mate guarding, reproductive success and female choice in African elephants
Poole, J. H. (1989) · Animal Behaviour · doi:10.1016/0003-3472(89)90068-7 · Cited by 242 · see citing works
Announcing intent: the aggressive state of musth in African elephants
Poole, J. H. (1989) · Animal Behaviour · doi:10.1016/0003-3472(89)90014-6 · Cited by 240 · see citing works
Ganswindt, A., Palme, R., Heistermann, M., Borragan, S., & Hodges, J. K. (2003) · General and Comparative Endocrinology · doi:10.1016/s0016-6480(03)00251-x · Cited by 213 · see citing works
Self-citationMusth in the African elephant, Loxodonta africana
Poole, J. H., & Moss, C. J. (1981) · Nature · doi:10.1038/292830a0 · Cited by 164 · see citing works
Barrett, G. M., Shimizu, K., Bardi, M., Asaba, S., & Mori, A. (2002) · Hormones and Behavior · doi:10.1006/hbeh.2002.1804 · Cited by 155 · see citing works
Ganswindt, A., Heistermann, M., Borragan, S., & Hodges, J. K. (2002) · Zoo Biology · doi:10.1002/zoo.10034 · Cited by 137 · see citing works
Self-citationReproductive Behavior of the Asiatic Elephant
Eisenberg, J. F., McKay, G. M., & Jainudeen, M. R. (1971) · Behaviour · doi:10.1163/156853971x00087 · Cited by 134 · see citing works
Testart, J., Frydman, R. R. F., Feinstein, M.-C., et al. (1981) · Fertility and Sterility · doi:10.1016/s0015-0282(16)45617-7 · Cited by 124 · see citing works
Fieß, M., Heistermann, M., & Hodges, J. K. (1999) · General and Comparative Endocrinology · doi:10.1006/gcen.1999.7287 · Cited by 113 · see citing works
Self-citationViau, V., & Meaney, M. (2004) · Journal of Endocrinology · doi:10.1677/joe.0.1810223 · Cited by 93 · see citing works
Place, N. J., & Kenagy, G. J. (2000) · Journal of Comparative Physiology B · doi:10.1007/s003600050282 · Cited by 91 · see citing works
Testosterone and Dihydrotestosterone Concentrations in Elephant Serum and Temporal Gland Secretions
Rasmussen, L. E. L., Buss, I. O., Hess, D. L., & Schmidt, M. J. (1984) · Biology of Reproduction · doi:10.1095/biolreprod30.2.352 · Cited by 90 · see citing works
Musth and urinary testosterone concentrations in the African elephant ( Loxodonta africana )
Poole, J. H., Kasman, L. H., Ramsay, E. C., & Lasley, B. L. (1984) · Reproduction · doi:10.1530/jrf.0.0700255 · Cited by 87 · see citing works
Kenagy, G. J., Place, N. J., & Veloso, C. (1999) · General and Comparative Endocrinology · doi:10.1006/gcen.1999.7307 · Cited by 77 · see citing works
Lincoln, G. A., & Ratnasooriya, W. D. (1996) · Reproduction · doi:10.1530/jrf.0.1080107 · Cited by 61 · see citing works
Rasmussen, L. E. L., Hall-Martin, A. J., & Hess, D. L. (1996) · Journal of Mammalogy · doi:10.2307/1382819 · Cited by 52 · see citing works
Serum testosterone and musth in captive male African and Asian elephants
Cooper, K. A., Harder, J. D., Clawson, D. H., Fredrick, D. L., Lodge, G. A., Peachey, H., Spellmire, T. J., & Winstel, D. P. (1990) · Zoo Biology · doi:10.1002/zoo.1430090405 · Cited by 49 · see citing works
The value of ultrasound, gonadotropin, and estradiol measurements for precise ovulation prediction
Bryce, R. L., Shuter, B., Sinosich, M. J., et al. (1982) · Fertility and Sterility · doi:10.1016/s0015-0282(16)45974-1 · Cited by 49 · see citing works
Brannian, J. D., Griffin, F. M., & Terranova, P. F. (1989) · Zoo Biology · doi:10.1002/zoo.1430080207 · Cited by 15 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
Brown, J. L., Somerville, M., Riddle, H. S., Keele, M., Duer, C. K., & Freeman, E. W. (2007) · General and Comparative Endocrinology · doi:10.1016/j.ygcen.2007.01.006 · Cited by 83 · see citing works
RASMUSSEN, H., GANSWINDT, A., DOUGLASHAMILTON, I., & VOLLRATH, F. (2008) · Hormones and Behavior · doi:10.1016/j.yhbeh.2008.05.008 · Cited by 58 · see citing works
Kumar, V., Palugulla Reddy, V., Kokkiligadda, A., Shivaji, S., & Umapathy, G. (2014) · General and Comparative Endocrinology · doi:10.1016/j.ygcen.2014.03.024 · Cited by 55 · see citing works
Bansiddhi, P., Brown, J. L., Khonmee, J., Norkaew, T., Nganvongpanit, K., Punyapornwithaya, V., Angkawanish, T., Somgird, C., & Thitaram, C. (2019) · PLoS ONE · doi:10.1371/journal.pone.0221537 · Cited by 46 · see citing works
Facilitating the social behaviour of bull elephants in zoos
Hartley, M., Wood, A., & Yon, L. (2019) · International Zoo Yearbook · doi:10.1111/izy.12245 · Cited by 24 · see citing works
Physiological plasticity in elephants: highly dynamic glucocorticoids in African and Asian elephants
Pokharel, S. S., & Brown, J. L. (2023) · Conservation Physiology · doi:10.1093/conphys/coad088 · Cited by 23 · see citing works
LaDue, C. A., Hunt, K. E., Samaraweera, M. G. S. M., Vandercone, R. P. G., Kiso, W. K., & Freeman, E. W. (2022) · Theriogenology Wild · doi:10.1016/j.therwi.2022.100011 · Cited by 18 · see citing works
Schreier, A. L., Readyhough, T. S., Moresco, A., Davis, M., & Joseph, S. (2023) · Journal of Applied Animal Welfare Science · doi:10.1080/10888705.2021.1908141 · Cited by 15 · see citing works
Long-Term Olfactory Memory in African Elephants
Hoerner, F., Lawrenz, A., Oerke, A.-K., Müller, D. W. H., Azogu-Sepe, I., Roller, M., Damerau, K., & Preisfeld, A. (2023) · Animals · doi:10.3390/ani13040679 · Cited by 8 · see citing works
Morris‐Drake, A., & Mumby, H. S. (2018) · Mammal Review · doi:10.1111/mam.12106 · Cited by 8 · see citing works
Nighttime behavior and the length of social relationships in male Asian elephants
Thevarajah, S. J., Readyhough, T. S., Davis, M., Moresco, A., Joseph, S., & Schreier, A. L. (2023) · Journal of Applied Animal Welfare Science · doi:10.1080/10888705.2021.1998777 · Cited by 6 · see citing works
Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR
López Pérez, A. B., Brown, J. L., Goldenberg, S. Z., Rey, A., & Khonmee, J. (2025) · PLOS One · doi:10.1371/journal.pone.0332944 · Cited by 2 · see citing works
- Authors
- Ganswindt, A., Heistermann, M., & Hodges, K.
- Affiliations
- German Primate Center (DE)
- Corresponding author
- Ganswindt, André
- Heistermann, Michael
- Hodges, Keith
- ORCID
- Ganswindt, André
- Venue
- Physiological and Biochemical Zoology
- Publisher
- University of Chicago Press
- Published
- 2005-07
- Volume
- 78
- Issue
- 4
- Pages
- 505-514
- ISSN
- 1522-2152 (ISSN-L)
- 1537-5293
- OpenAlex
- W2035312355
- Cited by
- 58
- References
- 22
- Citations by year
- 2026: 4 · 2025: 2 · 2024: 1 · 2023: 3 · 2022: 6 · 2021: 5 · 2020: 4 · 2019: 4 · 2018: 2 · 2017: 2 · 2016: 2 · 2015: 3 · 2014: 2 · 2013: 3 · 2012: 2
- FWCI
- 2.14
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Physiological and Biochemical Zoology (publishedVersion)
- PubMed (publishedVersion)
- Nelson Mandela University Research Repository (submittedVersion)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
- Keywords
- musth, Loxodonta africana, captive elephants, androgens, aggression, glucocorticoids, hypothalamic-pituitary-adrenal axis, testicular function, adrenal function
- MeSH terms
- Adrenal Glands, Aggression, Analysis of Variance, Androgens, Animals, Animals, Zoo, Elephants, Feces, Glucocorticoids, Male, Reproduction, Sexual Behavior, Animal, Testis, Time Factors, Observation
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