Anatomical, physiological, and behavioral mechanisms of thermoregulation in elephants


  • Adriana Domínguez-Oliva Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Marcelo Daniel Ghezzi Animal Welfare Area, Faculty of Veterinary Sciences (FCV). Universidad Nacional del Centro de la Provincia de Buenos Aires (UNCPBA), Argentina.
  • Patricia Mora-Medina Facultad de Estudios Superiores Cuautitlán. Universidad Nacional Autónoma de México (UNAM), Mexico.
  • Ismael Hernández-Ávalos Facultad de Estudios Superiores Cuautitlán. Universidad Nacional Autónoma de México (UNAM), Mexico.
  • Joseline Jacome Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Andrea Castellón Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Isabel Falcón Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Fátima Reséndiz Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Nicole Romero Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Raúl Ponce Neurophysiology, Behavior and Animal Welfare Assessment, Xochimilco Campus, Universidad Autónoma Metropolitana, Ciudad de México 04960, Mexico.
  • Daniel Mota-Rojas Universidad Autonoma Metropolitana



Loxodonta africana, Elephas maximus, elephant anatomy, thermal state


Elephants use different thermoregulatory mechanisms that depend on the anatomical and morphological characteristics of the species. The crevices and wrinkles of the skin enhance the water-retention capacity of the epidermis. The highly vascularized ear is another region of particular interest, as its movement and vasomotor changes promote heat dissipation. Generally, these mechanisms are modulated by the hypothalamic thermoregulatory center and by the peripheral response of animals. Nonetheless, elephants are currently exposed to alterations in their habitats, such as global warming and climatic changes, which challenge their homeothermy. This article aims to discuss the thermoregulation mechanisms of African (Loxodonta africana) and Asian (Elephas maximus) elephants from an anatomical, physiological, and behavioral basis. The practical implications of these elements will be analyzed to implement tools, such as infrared thermography, or environmental enrichment, as strategies to promote the thermal balance of elephants.


Anastassakis K (2022) The mission of hair follicles and hair. In: Anastassakis K (ed) Androgenetic alopecia from A to Z. Springer Nature, Switzerland, pp.

Athurupana R, Schmitt D, Santiapillai C (2015) Pinnae movement of captive asian elephants weakly affected by environmental factors. Gajah 43:4–9.

Avni-Magen, Zaken S, Kaufman E, Kelmer G (2017) Use of infrared thermography in early diagnosis of pathologies in asian elephants (Elephas maximus). Israel Journal of Veterinary Medicine 72:22–27.

Ayer A, Mariappa D (1950) External characters of three foetuses of the indian elephant. In: Proceedings of the Indian Academy of Sciences. 31:193–209.

Bansiddhi P, Brown JL, Thitaram C, Punyapornwithaya V, Nganvongpanit K (2020) Elephant tourism in thailand: a review of animal welfare practices and needs. Journal of Applied Animal Welfare Science 23:164–177.

Barandongo ZR, Mfune JKE, Turner WC (2018) Dust-bathing behaviors of african herbivores and the potential risk of inhalational anthrax. Journal of Wildlife Diseases 54:34.

Barlett S (2006) Cardiovascular system. In: Fowler ME, Mikota SK (eds) Biology, medicine and surgery of elephants. Blackwell, Australia, pp 317–324.

Baskaran N (1998) Ranging and resource utilization by Asian elephant (Elephas maximus Linnaeus) in Nilgiri biosphere reserve, South India. Dissertation, Bharathidasan University.

Birkett PJ, Vanak AT, Muggeo VMR, Ferreira SM, Slotow R (2012) Animal perception of seasonal thresholds: changes in elephant movement in relation to rainfall patterns. PLoS One 7:e38363.

Carrington R (1959) Elephants: a short account of their natural history evolution and influence on mankind. Basic Books, USA, pp 272.

Chamaille-Jammes S, Mtare G, Makuwe E, Fritz H (2013) African elephants adjust speed in response to surface water constraint on foraging during the dry-season. PLoS One 8:1–6.

Choudhury A (2004) Human–elephant conflicts in northeast india. Human Dimensions of Wildlife 9:261–270.

Cilulko J, Janiszewski P, Bogdaszewski M, Szczygielska E (2013) Infrared thermal imaging in studies of wild animals. European Journal of Wildlife Research 59:17–23.

Codron J, Lee-Thorp JA, Sponheimer M, Codron D, Grant RC, de Ruiter DJ (2006) Elephant (Loxodonta africana) diets in Kruger National Park, South Africa: spatial and landscape differences. Journal of Mammalogy 87:27–34.

Cole M (1986) The savannas: biogeography and geobotany. Academic Press, London, pp 438.

Diaz SG, DeAngelis DL, Gaines MS, Purdon A, Mole MA, van Aarde RJ (2021) Development and validation of a spatially-explicit agent-based model for space utilization by African savanna elephants (Loxodonta africana) based on determinants of movement. Ecological Modelling 447:1–14.

Du Plessis K, Ganswindt SB, Bertschinger H, Crossey B, Henley MD, Ramahlo M, Ganswindt A, Baotic A, Meehan C (2021) Social and seasonal factors contribute to shifts in male african elephant (Loxodonta africana) foraging and activity patterns in Kruger National Park, South Africa. Animals 11:3070.

Dunkin RC, Wilson D, Way N, Johnson K, Williams TM (2013) Climate influences thermal balance and water use in African and Asian elephants: physiology can predict drivers of elephant distribution. Journal of Experimental Biology 216:2939–2952.

Escobar-Ibarra I, Mota-Rojas D, Gual-Sil F, Sanchez CR, Baschetto F, Alonso-Spilsbury M (2021) Conservation, animal behaviour, and human-animal relantionship in zoos. Why is animal welfare so important? Journal of Animal Behaviour and Biometeorology 9:2111.

Feldhamer GA, Merritt JF, Krajewski C, Rachow JL, Stewart KM (2020) Mammology. Adaptation, diversity, ecology, 5th edn. Johns Hopkins University Press, United States.

Fowler ME (2006) Multisystem disorders. In: Fowler ME, Mikota SK (eds) Biology, medicine and surgery of elephants. Blackwell Publishing, Australia, pp 243–252.

Glaeser SS, Shepherdson D, Lewis K, Prado N, Brown JL, Lee B, Wielebnowski N (2021) Supporting zoo asian elephant (Elephas maximus) welfare and herd dynamics with a more complex and expanded habitat. Animals 11:2566.

Grubb P, Groves CP, Dudley JP, Shoshani J (2000). Living African elephants belong to two species: Loxodonta africana (Blumenbach, 1797) and Loxodonta cyclotis (Matschie, 1900). Elephant 2:1–4.

Hidden P (2009) Thermoregulation in African elephants (Loxodonta africana). Dissertation, University of the Witwatersrand.

Hiley PG (1975) How the elephant keeps its cool. Natural History 84:34–41.

Ikoma Y, Kusumoto-Yoshida I, Yamanaka A, Ootsuka Y, Kuwaki T (2018) Inactivation of serotonergic neurons in the rostral medullary raphé attenuates stress-induced tachypnea and tachycardia in mice. Frontiers in Physiology 9:832.

Isaza R, Hunter R (2004) Drug delivery to captive asian elephants - treating goliath. Current Drug Delivery 1:291–298.

Ishiwata T, Saito T, Hasegawa H, Yazawa T, Kotani Y, Otokawa M, Aihara Y (2005) Changes of body temperature and thermoregulatory responses of freely moving rats during GABAergic pharmacological stimulation to the preoptic area and anterior hypothalamus in several ambient temperatures. Brain Research 1048:32–40.

Jachowski DS, Slotow R, Millspaugh JJ (2012) Physiological stress and refuge behavior by African elephants. PLoS One 7:31818.

Kinahan AA, Inge-moller R, Bateman PW, Kotze A, Scantlebury M (2007a) Body temperature daily rhythm adaptations in African savanna elephants (Loxodonta africana). Physiology and Behavior 92:560–565.

Kinahan AA, Pimm SL, Van Aarde RJ (2007b) Ambient temperature as a determinant of landscape use in the savanna elephant, Loxodonta africana. Journal of Thermal Biology 32:47–58.

Kingma B (2012) The thermoneutral zone: implications for metabolic studies. Frontiers in Bioscience 4:1975.

Kosaruk W, Brown JL, Plangsangmas T, Towiboon P, Punyapornwithaya V, Silva-Fletcher A, Thitaram C, Khonmee J, Edwards KL, Somgird C (2020) Effect of tourist activities on fecal and salivary glucocorticoids and immunoglobulin A in female captive Asian elephants in thailand. Animals. 10:1928.

Kruk B, Davydov AF (1977) Effect of ambient temperature on thermal sensitivity of POA area in the rabbit. Journal of Thermal Biology 2:75–78.

Lamps LW, Smoller BR, Rasmussen LEL, Slade BE, Fritsch G, Goodwin TE (2001) Characterization of interdigital glands in the Asian elephant (Elephas maximus). Research in Veterinary Science 71:197–200.

Langman VA, Rowe M, Forthman D, Langman N, Black J, Walker T (2003) Quantifying shade using a standard environment. Zoo Biology 22:253–260.

Lee PC, Moss CJ (1995) Statural growth in known‐age African elephants (Loxodonta africana). Journal of Zoology 236:29–41.

Lezama-García K, Mota-Rojas D, Pereira AMF, Martínez-Burnes J, Ghezzi M, Domínguez A, Gómez J, de Mira Geraldo A, Lendez P, Hernández-Ávalos I, Falcón I, Olmos-Hernández A, Wang D (2022) Transient Receptor Potential (TRP) and thermoregulation in animals: structural biology and neurophysiological aspects. Animals 12:106.

Lillywhite HB, Stein BR (1987) Surface sculpturing and water retention of elephant skin. Journal of Zoology 211:727–734.

Low ZWK, Li Z, Owh C, Chee PL, Ye E, Dan K, Chan SY, Young DJ, Loh XJ (2020) Recent innovations in artificial skin. Biomaterials Science 8:776–797.

Marchant GH, Shoshani J (2007) Head muscles of Loxodonta africana and Elephas maximus with comments on Mammuthus primigenius muscles. Quaternary International 169–170:186–191.

Martin P, Bateson P (1993) Measuring behaviour. an introductory guide, 2nd edn. Cambridge University Press, United Kingdom.

Martins AF, Bennett NC, Clavel S, Groenewald H, Hensman S, Hoby S, Joris A, Manger PR, Milinkovitch MC (2018) Locally-curved geometry generates bending cracks in the African elephant skin. Nature Communications 9:3865.

Mason GJ (2010) Species differences in responses to captivity: Stress, welfare and the comparative method. Trends in Ecology and Evolution 25:713–721.

McCafferty DJ (2007) The value of infrared thermography for research on mammals: Previous applications and future directions. Mammal Review 37:207–223.

Meehan CL, Hogan JN, Bonaparte-Saller MK, Mench JA (2016) Housing and social environments of African (Loxodonta africana) and Asian (Elephas maximus) Elephants in North American Zoos PLOS One 11:e0146703.

Mikota SK (2006) Integument system. In: Fowler ME, Mikota SK (eds) Biology, medicine and surgery of elephants. Blackwell Publishing, Australia, pp 253–262.

Mitchell D, Maloney S, Jessen C, Laburn H, Kamerman P, Mitchell G, Fuller A (2002) Adaptive heterothermy and selective brain cooling in arid-zone mammals. Comparative biochemistry and Physiology Part B 131:571–585.

Mole MA, Rodrigues D’Araujo S, van Aarde RJ, Mitchell D, Fuller A (2016) Coping with heat: behavioural and physiological responses of savanna elephants in their natural habitat. Conservation Physiology 4:44.

Mole MA, Rodrigues DÁraujo S, van Aarde RJ, Mitchell D, Fuller A (2018) Savanna elephants maintain homeothermy under African heat. Journal of Comparative Physiology B 188:889–897.

Morf J, Schibler U (2013) Body temperature cycles: Gatekeepers of circadian clocks. Cell Cycle 12:539–540.

Morgan BJ, Lee PC (2003) Forest elephant (Loxodonta africana cyclotis) stature in the Réserve de Faune du Petit Loango, Gabon. Journal of Zoology 259:337–344.

Morrison SF (2016) Central control of body temperature. F1000 Research 5:880.

Morrison SF, Nakamura K (2019) Central mechanisms for thermoregulation. Annual Review of Physiology 81:285–308.

Mota-Rojas D, Mariti C, Zdeinert A, Riggio G, Mora-Medina P, del Mar Reyes A, Gazzano A, Domínguez-Oliva A, Lezama-García K, José-Pérez N, Hernández-Ávalos I (2021a) Anthropomorphism and its adverse effects on the distress and welfare of companion animals. Animals 11:3263.

Mota-Rojas D, Titto CG, Orihuela A, Martínez-Burnes J, Gómez-Prado J, Torres-Bernal F, Flores-Padilla K, Carvajal-de la Fuente V, Wang D (2021b) Physiological and behavioral mechanisms of thermoregulation in mammals. Animals 11:1733.

Mota-Rojas D, Titto CG, de Mira Geraldo A, Martínez-Burnes J, Gómez J, Hernández-Ávalos I, Casas A, Domínguez A, José N, Bertoni A, Reyes B, Pereira AMF (2021c) Efficacy and function of feathers, hair, and glabrous skin in the thermoregulation strategies of domestic animals. Animals 11:3472.

Mota-Rojas D, Wang D, Titto CG, Gómez-Prado J, Carvajal-De la Fuente V, Ghezzi M, Boscato-Funes L, Barrios-García H, Torres-Bernal F, Casas-Alvarado A, Martínez-Burnes J (2021d) Pathophysiology of fever and application of infrared thermography (IRT) in the detection of sick domestic animals: Recent advances. Animals 11:2316.

Mota-Rojas D, Miranda-Cortés A, Casas-Alvarado A, Mora-Medina P, Boscato-Funes L, Hernández-Ávalos I (2021e) Neurobiología y modulación de la hipertermia inducida por estrés agudo y fiebre en los animales. Abanico Veterinario 11: 1–17.

Mota-Rojas D, Pereira AMF, Wang D, Martínez-Burnes J, Ghezzi M, Hernández-Avalos I, Lendez P, Mora-Medina P, Casas A, Olmos-Hernández A, Domínguez A, Bertoni A, de Mira Geraldo A (2021f) Clinical applications and factors involved in validating thermal windows used in infrared thermography in cattle and river Buffalo to assess health and productivity. Animals 11:2247.

Mota-Rojas D, Napolitano F, Braghieri A, et al (2021g) Thermal Biology in River Buffalo in the Humid Tropics: Neurophysiological and Behavioral Responses Assessed by Infrared Thermography. Journal of Animal Behaviour and Biometeorology 9:2103.

Mota-Rojas D, Wang D, Titto CG, et al (2022) Neonatal infrared thermography images in the hypothermic ruminant model: Anatomical-morphological-physiological aspects and mechanisms for thermoregulation. Front Vet Sci 9:963205.

Mpakairi KS, Ndaimani H, Tagwireyi P, Zvidzai M, Madiri TH (2020) Futuristic climate change scenario predicts a shrinking habitat for the African elephant (Loxodonta africana): evidence from Hwange National Park, Zimbabwe. European Journal of Wildlife Research 66:1.

Mumby HS, Mar KU, Thitaram C, Courtiol A, Towiboon P, Min-Oo Z, Htut-Aung Y, Brown JL, Lummaa V (2015) Stress and body condition are associated with climate and demography in Asian elephants. Conservation Physiology 3:cov030.

Napolitano F, Mota-Rojas D, Guerrero-Legarreta I, Orihuela A (2020) El búfalo de agua en Latinoamérica, hallazgos recientes. 3ra ed. BM Editores, México.

Narasimhan A (2008) Why do elephants have big ear flaps? Resonance 13:638–647.

Pastorini J, Nishantha HG, Janaka HK, Isler K, Fernando P (2010) Water body use by Asian elephants in Southern Sri Lanka. Tropical Conservation Science 3:412–422.

Peeks M, Badarnah L (2021) Textured building façades: utilizing morphological adaptations found in nature for evaporative cooling. Biomimetics 6:24.

Phillips PK, Heath JE (1995) Dependency of surface temperature regulation on body size in terrestrial mammals. Journal of Thermal Biology 20:281–289

Phillips PK, Heath JE (1992) Heat exchange by the pinna of the african elephant (Loxodonta africana). Comparative Biochemistry and Physiology Part A: Physiology 101:693–699.

Pontzer H, Rimbach R, Paltan J, Ivory EL, Kendall CJ (2020) Air temperature and diet influence body composition and water turnover in zoo-living African elephants (Loxodonta africana). Royal Society Open Science 7:201155.

Purdon A, van Aarde RJ (2017) Water provisioning in Kruger National Park alters elephant spatial utilisation patterns. Journal of Arid Environments 141:45–51.

Rees P (2002a) Seasonal and roofing material influence on the thermoregulation by captive asian elephants and its implications for captive elephant welfare. Journal of Thermal Biology 27:353–358

Rees PA (2002b) Asian elephants (Elephas maximus) dust bathe in response to an increase in environmental temperature. Journal of Thermal Biology 27:353–358

Rowe MF, Bakken GS, Ratliff JJ, Langman VA (2013) Heat storage in Asian elephants during submaximal exercise: behavioral regulation of thermoregulatory constraints on activity in endothermic gigantotherms. Journal of Experimental Biology 216:1774–1785.

Rubio-Martínez LM, Hendrickson DA, Stetter M, Zuba JR, Marais HJ (2014) Laparoscopic vasectomy in African elephants (Loxodonta africana). Veterinary Surgery 43:507–514.

Shelton DS, Alberts JR (2018) Development of behavioral responses to thermal challenges. Developmental Psychobiology 60:5–14.

Shoshani J, Eisenberg JF (1982) Mammalian species. Elephas maximus. American Society of Mammalogist 182:1–8.

Shoshani J, Foley C (2000) Frequently asked questions about elephants. Elephant 2:78–87.

Spearman RIC (1970) The epidermis and its keratinisation in the African elephant (Loxodonta africana). Zoologica Africana 5:327–338.

Stoinski TS, Daniel E, Maple TL (2000) A preliminary study of the behavioral effects of feeding enrichment on African elephants. Zoo Biology 19:485–493.

Strauss WM, Hetem RS, Mitchell D, Maloney SK, O’Brien HD, Meyer LCR, Fuller A (2017) Body water conservation through selective brain cooling by the carotid rete: a physiological feature for surviving climate change? Conservation Physiology 5:1–15.

Šumbera R, Zelová J, Kunc P, Knížková I, Burda H (2007) Patterns of surface temperatures in two mole-rats (Bathyergidae) with different social systems as revealed by IR-thermography. Physiology and Behavior 92:526–532.

Terrien J (2011) Behavioral thermoregulation in mammals: a review. Frontiers in Bioscience 16:1428.

Thaker M, Gupte PR, Prins HHT, Slotow R, Vanak AT (2019) Fine-scale tracking of ambient temperature and movement reveals shuttling behavior of elephants to water. Frontiers in Ecology and Evolution 7:4.

Thitaram C, Matchimakul P, Pongkan W, Tangphokhanon W, Maktrirat R, Khonmee J, Sathanawongs A, Kongtueng P, Nganvongpanit K (2018) Histology of 24 organs from Asian elephant calves (Elephas maximus). PeerJ. 6:e4947.

Tupone D, Madden CJ, Morrison SF (2014) Autonomic regulation of brown adipose tissue thermogenesis in health and disease: potential clinical applications for altering BAT thermogenesis. Frontiers in Neuroscience 8:1–14.

Uchida Y, Onishi K, Tokizawa K, Nagashima K (2017) Regional differences of cFos immunoreactive cells in the preoptic areas in hypothalamus associated with heat and cold responses in mice. Neuroscience Letters 665:130–134.

Vanitha V, Baskaran N (2010) Seasonal and roofing material influence on the thermoregulation by captive asian elephants and its implications for captive elephant welfare. Gaja 33:35–40

Wang H, Wang P, Zhao X, Zhang W, Li J, Xu C, Xie P (2021) What triggered the Asian elephant’s northward migration across southwestern Yunnan? The Innovation 2:100142.

Weissenböck NM, Arnold W, Ruf T (2012) Taking the heat: thermoregulation in Asian elephants under different climatic conditions. Journal of Comparative Physiology B 182:311–319.

Weissenböck NM, Weiss CM, Schwammer HM, Kratochvil H (2010) Thermal windows on the body surface of African elephants (Loxodonta africana) studied by infrared thermography. Journal of Thermal Biology 35:182–188.

Wenwen L, Yang Y, Peng L, Ruchun T, Yunchuan D, Li L, Zhang L (2019) Identifying climate refugia and its potential impact on small population of Asian elephant (Elephas maximus) in China. Global Ecology and Conservation 19:e00664.

Williams’ TM (1990) Heat transfer in elephants: thermal partitioning based on skin temperature profiles. Journal of Zoology 22:235–245.

Wright PG, Luck CP (1984) Do elephants need to sweat? South African Journal of Zoology 19:270–274.

Yang X, Ruan H-B (2015) Neuronal control of adaptive thermogenesis. Frontiers in Endocrinology 6:149.

Zakharian E, Cao C, Rohacs T (2010) Gating of Transient Receptor Potential Melastatin 8 (TRPM8) channels activated by cold and chemical agonists in planar lipid bilayers. Journal of Neuroscience 30:12526–12534.

Mechanisms of central and peripheral thermoregulation. The image illustrates the integration of the thermal stimuli detected by the corpuscles of Ruffini or Krause in the face of heat or cold, respectively. Although the central pathways in both cases are very similar and use the DRG, POA, MnPO, or rRPA (black lines), the main difference is observed in the LPBN regions that are activated by stimuli of a different nature. The red lines show the thermoregulatory mechanisms in warm environments, where LPBd neurons produce vasodilation. In contrast, with cold stimuli (blue lines), the LPBel neurons are stimulated to generate sympathetic vasoconstriction and stimulate motor fibers that promote shivering and non-shivering thermogenesis. DRG: dorsal root ganglia; LPBd: dorsal part of the lateral parabrachial nucleus; LPBeL: Lateral parabrachial nucleus in its external region; POA: preoptic area of the hypothalamus; MnPO: median preoptic nucleus; MPO: medial preoptic area; DMH: dorsomedial hypothalamus; rRPA: rostral raphe pallidus.



PUBLISHED: 2022-09-22

How to Cite

Domínguez-Oliva, A., Ghezzi, M. D., Mora-Medina, P., Hernández-Ávalos, I., Jacome, J., Castellón, A., Falcón, I., Reséndiz, F., Romero, N., Ponce, R., & Mota-Rojas, D. (2022). Anatomical, physiological, and behavioral mechanisms of thermoregulation in elephants. Journal of Animal Behaviour and Biometeorology, 10(4), 2233.



Review Article

Most read articles by the same author(s)