Reproductive thermoregulation in marine mammals: mechanisms, adaptations and evolutionary implications

Keywords: Marine thermoregulation, Reproductive adaptation, Heat exchange, Internal testes, Aquatic ecology

Abstract

Marine mammals inhabit thermally challenging environments in which maintaining internal body temperature is essential for survival, metabolic function, and reproductive success. Unlike terrestrial mammals, many marine male species lack external reproductive structures (e.g. scrotum), raising questions about how reproductive temperatures are regulated in the absence of conventional cooling mechanisms. This review synthesizes current knowledge of reproductive thermoregulation in marine mammals, with a focus on cetaceans and pinnipeds, and highlights key anatomical and physiological adaptations. These include countercurrent heat exchange systems, specialized vascular networks surrounding reproductive organs, and behavioral strategies that facilitate localized cooling while conserving overall body heat. The review also addresses thermal protection of the developing fetus, illustrating how vascularization and insulation help maintain optimal intrauterine conditions. In addition, it examines evolutionary trade-offs associated with internal testes and streamlined body forms, ecological pressures imposed by aquatic environments, and the integration of thermoregulation with diving physiology. Finally, implications for conservation are discussed, particularly in the context of climate change, rising ocean temperatures and human-induced habitat disturbances. By integrating anatomical, physiological, ecological and evolutionary perspectives, this article underscores how marine mammals have evolved highly efficient thermoregulatory strategies, offering insights into reproductive biology and species resilience in dynamic aquatic ecosystems.

Downloads

Download data is not yet available.

References


1.
Morrison SF, Nakamura K: Central mechanisms for thermoregulation. Ann Rev Physiol 2019;81:285–308. doi: 10.1146/annurev-physiol-020518-114546


2.
Mota-Rojas D, Titto CG, Orihuela A, et al: Physiological and behavioral mechanisms of thermoregulation in mammals. Animals (Basel) 2021;11:1733. doi: 10.3390/ani11061733


3.
Knapp BD, Huang KC: The effects of temperature on cellular physiology. Ann Rev Biophys 2022;51:499–526. doi: 10.1146/annurev-biophys-112221-074832


4.
Maroto M, Torvisco SN, García-Merino C, et al: Mechanisms of hormonal, genetic, and temperature regulation of germ cell proliferation, differentiation, and death during spermatogenesis. Biomolecules 2025;15:500. doi: 10.3390/biom15040500


5.
Setchell BP: The Parkes Lecture. Heat and the testis. J Reprod Fertil 1998;114:179–194. doi: 10.1530/jrf.0.1140179


6.
Setchell BP, Breed WG: Anatomy, vasculature and innervation of the male reproductive tract. In: Neill JD: editor. Knobil and Neill’s Physiology of Reproduction. San Diego, CA; Elsevier: 2006. p. 771–825.


7.
Samuels L, Nakstad B, Roos N, et al: Physiological mechanisms of the impact of heat during pregnancy and the clinical implications: review of the evidence from an expert group meeting. Int J Biometeorol 2022;66:1505–1513. doi: 10.1007/s00484-022-02301-6


8.
Konkel L: Taking the heat: potential fetal health effects of hot temperatures. Environ Health Perspect 2019;127:102002. doi: 10.1289/EHP6221


9.
Rommel SA, Pabst DA, McLellan WA, et al: Anatomical evidence for a countercurrent heat exchanger associated with dolphin testes. Anat Rec 1992;232:150–156. doi: 10.1002/ar.1092320117


10.
Pabst DA, Rommel SA, McLellan W: Reproductive thermoregulation in marine mammals. Am Sci 1998;86:440–448. doi: 10.1511/1998.37.901


11.
Rommel SA, Pabst DA, McLellan WA, et al: Temperature regulation of the testes of the bottlenose dolphin (Tursiops truncatus): evidence from colonic temperatures. J Comp Physiol B 1994;164:130–134. doi: 10.1007/BF00301654


12.
Pabst DA, McLellan WA, Rommel SA: Thermoregulation of reproductive organs. In: Reynolds JE III, Rommel SA: editors. Biology of Marine Mammals. Washington, DC; Smithsonian Institution Press: 1999. p. 379–403.


13.
Irving L, Hart JS: The metabolism and insulation of seals as bare-skinned mammals in cold water. Can J Zool 2011;35:497–511. doi: 10.1139/z57-041


14.
Pabst DA, McLellan WA, Rommel SA: How do marine mammals thermoregulate? In: Reynolds JE III, Rommel SA: editors. Biology of Marine Mammals. Washington, DC; Smithsonian Institution Press: 1999. p. 379–403.


15.
Hokkanen JE: Temperature regulation of marine mammals. J Theor Biol 1990;145:465–485. doi: 10.1016/s0022-5193(05)80482-5


16.
Setchell BP: The effects of heat on the testes of mammals. In: Proceedings of the International Symposium on Animal Biology of Reproduction; 2006 November 15-18; Belo Horizonte, Brazil Anim Reprod 2006;3:81–91.


17.
Hansen P: Effects of heat stress on mammalian reproduction. Philos Trans R Soc Lond B Biol Sci 2009;364:3341–3350. doi: 10.1098/rstb.2009.0131


18.
Paul C, Murray AA, Spears N, et al: A single, mild, transient scrotal heat stress causes DNA damage, subfertility and impairs formation of blastocysts in mice. Reproduction 2008;136:73–84. doi: 10.1530/REP-08-0036


19.
Rockett JC, Mapp FL, Garges JB, et al: Effects of hyperthermia on spermatogenesis, apoptosis, gene expression, and fertility in adult male mice. Biol Reprod 2001;65:229–239. doi: 10.1095/biolreprod65.1.229


20.
Kastelic JP, Rizzoto G, Thundathil J: Testicular vascular cone development and its association with scrotal thermoregulation, semen quality and sperm production in bulls. Animal 2018;12(Suppl 1):s133–s141. doi: 10.1017/S1751731118001167


21.
Waites GM: Thermoregulation of the scrotum and testis: studies in animals and significance for man. Adv Exp Med Biol 1991;286:9–17. doi: 10.1007/978-1-4684-5913-5_2


22.
Ford SP: Control of blood flow to the gravid uterus of domestic livestock species. J Anim Sci 1995;73:1852–1860. doi: 10.2527/1995.7361852x


23.
Baumgard LH, Rhoads RP, Jr: Effects of heat stress on postabsorptive metabolism and energetics. Annu Rev Anim Biosci 2013;1:311–337. doi: 10.1146/annurev-animal-031412-103644


24.
Kooyman GL: Diverse Divers: Physiology and Behavior. Berlin; Springer-Verlag: 1989.


25.
Rommel SA, Pabst DA, McLellan WA: Functional morphology of the vascular plexuses associated with the cetacean uterus. Anat Rec 1993;237:414–426. doi: 10.1002/ar.1092370414


26.
Favilla AB, Horning M, Costa DP: Advances in thermal physiology of diving marine mammals: the dual role of peripheral perfusion. Temperature (Austin) 2021;9:46–66. doi: 10.1080/23328940.2021.1988817


27.
Hindle AG, Horning M, Mellish J-AE: Estimating total body heat dissipation in air and water from skin surface heat flux telemetry in Weddell seals. Anim Biotelem 2015;3:1–11. doi: 10.1186/s40317-015-0081-4


28.
Kvadsheim PH, Folkow LP: Blubber and flipper heat transfer in harp seals. Acta Physiol Scand 1997;161:385–395. doi: 10.1046/j.1365-201X.1997.00235.x


29.
Fougeres E, McLellan WA, Westgate AJ, et al: The relationship between heat flow and vasculature in the dorsal fin of wild bottlenose dolphins (Tursiops truncatus). J Exp Biol 2002;205:3475–3486. doi: 10.1242/jeb.205.22.3475


30.
Blix AS, Fay FH, Ronald K: On testicular cooling in phocid seals. Polar Res 2010;1:231–233. doi: 10.3402/polar.v1i3.6989


31.
Scholander PF, Schevill WE: Counter-current vascular heat exchange in the fins of whales. J Appl Physiol 1955;8:279–282. doi: 10.1152/jappl.1955.8.3.279


32.
Ekdale EG, Kienle SS: Passive restriction of blood flow and counter-current heat exchange via lingual retia in the tongue of a neonatal gray whale (Eschrichtius robustus). Anat Rec (Hoboken) 2015;298:675–679. doi: 10.1002/ar.23111


33.
Castellini M, Thermoregulation: Encyclopedia of Marine Mammals. In: Perrin WF, Würsig B, Thewissen JGM: editors. 2nd edition, San Diego, CA, USA, Academic Pressx: 2009. pp. 1166–1171.


34.
Favilla AB, Costa DP: Thermoregulatory strategies of diving air-breathing marine vertebrates: a review. Front Ecol Evol 2020;8:555509. doi: 10.3389/fevo.2020.555509


35.
Iverson SJ, Blubber: Encyclopedia of Marine Mammals. In: Perrin WF, Würsig B, Thewissen JGM: editors. 2nd edition, San Diego, CA, USA, Academic Press: 2009. pp. 115–120.


36.
Rommel S, Pabst D, McLellan W: Functional anatomy of the cetacean reproductive system, with comparisons to the domestic dog. In: Miller DL: editor. Reproductive Biology and Phylogeny of Cetacea: Whales, Porpoises and Dolphins. 1st edition, Boca Raton, FL; CRC Press: 2007. p. 127–145.


37.
Whittow GC: Thermoregulatory adaptations in marine mammals: interacting effects of exercise and body mass: a review. Mar Mamm Sci 2006;3:220–241. doi: 10.1111/j.1748-7692.1987.tb00165.x


38.
Panneton WM: The mammalian diving response: an enigmatic reflex to preserve life? Physiology (Bethesda) 2013;28:284–297. doi: 10.1152/physiol.00020.2013


39.
Werdelin L, Nilsonne A: The evolution of the scrotum and testicular descent in mammals: a phylogenetic view. J Theor Biol 1999;196:61–72. doi: 10.1006/jtbi.1998.0821


40.
Kleisner K, Ivell R, Flegr J: The evolutionary history of testicular externalization and the origin of the scrotum. J Biosci 2010;35:27–37. doi: 10.1007/s12038-010-0005-7


41.
Liu J, Lu Z, Wang X, et al: From land to sea: unique evolution in reproductive strategies of marine mammals. Inno Life 2024;2:100090. doi: 10.59717/j.xinn-life.2024.100090


42.
Liwanag HEM, Berta A, Costa DP, et al: Morphological and thermal properties of mammalian insulation: the evolutionary transition to blubber in pinnipeds. Biol J Linn Soc 2012;107:774–787. doi: 10.1111/j.1095-8312.2012.01992.x


43.
Chetan-Welsh H, Hendry L: Convergent Evolution Explained with 13 Examples. Natural History Museum. Available from: https://www.nhm.ac.uk/discover/convergent-evolution.html [cited 28 March 2026].


44.
Chatten A, Grieve I, Meligoniti E, et al: Predicting the effects of climate change on the fertility of aquatic animals using a meta-analytic approach. Ecol Lett 2025;28:e70054. doi: 10.1111/ele.70054


45.
Wild S, Krützen M, Rankin RW, et al: Long-term decline in survival and reproduction of dolphins following a marine heatwave. Curr Biol 2019;29:R239𠄓R240. doi: 10.1016/j.cub.2019.02.047


46.
Davis RW: Metabolism and thermoregulation. In: Marine Mammals. Cham; Springer: 2019. p. 57–87.


47.
Kebke A, Samarra F, Derous D: Climate change and cetacean health: impacts and future directions. Phil Trans R Soc B 2022;377:20210249. doi: 10.1098/rstb.2021.0249


48.
Jepson PD, Deaville R, Barber JL, et al: PCB pollution continues to impact populations of orcas and other dolphins in European waters. Sci Rep 2016;6:18573. doi: 10.1038/srep18573


49.
Meager JJ, Limpus C: Mortality of inshore marine mammals in eastern Australia is predicted by freshwater discharge and air temperature. PLoS One 2014;9:e94849. doi: 10.1371/journal.pone.0094849


50.
National Research Council (US) Committee on Potential Impacts of Ambient Noise in the Ocean on Marine Mammals: Ocean Noise and Marine Mammals. Washington, DC; National Academies Press; 2003. p. 3. Effects of Noise on Marine Mammals. Available from: https://www.ncbi.nlm.nih.gov/books/NBK221255/ [cited 28 March 2026].


51.
Erbe C, Marley SA, Schoeman RP, et al: The effects of ship noise on marine mammals-a review. Front Mar Sci 2019;6:606. doi: 10.3389/fmars.2019.00606


52.
McDonald B: Impacts of Disturbance on Marine Mammals: Physiological and Behavioral Responses to Stressors. University Scholar Series. 2022. p. 48. Available from: https://scholarworks.sjsu.edu/uss/48 [cited 28 March 2026].


53.
Harding S, Cousins N: Review of the Impacts of Anthropogenic Underwater Noise on Marine Biodiversity and Approaches to Manage and Mitigate Them. Technical Series No. 99. Montreal: Secretariat of the Convention on Biological Diversity; 2022. Available from: https://www.cbd.int/doc/publications/cbd-ts-99-en.pdf [cited 28 March 2026].


54.
Gulland FMD, Baker JD, Howe M, et al: A review of climate change effects on marine mammals in United States waters: past predictions, observed impacts, current research and conservation imperatives. Clim Change Ecol 2022;3:100054. doi: 10.1016/j.ecochg.2022.100054


55.
Aulia C, Maulina J, Edelwis TW, et al: Survival strategies of marine mammals in extreme environments: a study of whales and seals. BIO Web Conf 2024;134:06014. doi: 10.1051/bioconf/202413406014


56.
Burgess EA, Lanyon JM: Reproductive science methods for marine mammals. Adv Exp Med Biol 2026;16:553–604. doi: 10.1007/978-3-031-87707-0_17


57.
Lanyon JM, Burgess EA: Reproductive science methods for wild, fully-marine mammals: current approaches and future applications. Adv Exp Med Biol 2019;1200:363–411. doi: 10.1007/978-3-030-23633-5_13


58.
Cottrell B, Kalacska M, Arroyo P, et al: 3D reconstructions of stranded marine mammals via easily accessible remote sensing tools for use in morphometrics and visualizations. Front Mar Sci 2025;12:1485788. doi: 10.3389/fmars.2025.1485788


59.
van Elk CE: Pathology of Stranded Marine Mammals. PhD thesis. Rotterdam; Erasmus University: 2021. Available from: https://repub.eur.nl/pub/135597/thesis.pdf [cited 29 March 2026].


60.
Okamura T, Smith AR, Mikami T, et al: Three-dimensional vascular structure of caudal and dorsal fins of a dwarf sperm whale. Ecol Evol 2024;14:e70727. doi: 10.1002/ece3.70727


61.
Walsh MT, Jones D, Rankin D, et al: Diagnostic visualization techniques in marine mammals: CAT, NMR, ultrasound, endoscopy, thermography, and radiography. In: International Association for Aquatic Animal Medicine Conference Proceedings. 1998: Section VI, Clinical Reports I. Available from: https://www.vin.com/doc/?id=3863761 [cited 29 March 2026].


62.
Gray R, Canfield P, Rogers T: Histology of selected tissues of the leopard seal and implications for functional adaptations to an aquatic lifestyle. J Anat 2006;209:179–199. doi: 10.1111/j.1469-7580.2006.00591.x


63.
White C, Colefax AP, Parra GJ: Drone infrared thermography for detecting skin thermal anomalies in bottlenose dolphins: preliminary insights. Ecol Evol 2026;16:e72892. doi: 10.1002/ece3.72892


64.
Russell JP, Osborn SD, Herrick KES, et al: Infrared thermography of the blowhole as a potential diagnostic tool for health assessment in killer whales (Orcinus orca). Animals 2024;14:1867. doi: 10.3390/ani14131867


65.
Melero M, Rodríguez-Prieto V, Rubio-García A, et al: Thermal reference points as an index for monitoring body temperature in marine mammals. BMC Res Notes 2015;8:411. doi: 10.1186/s13104-015-1383-6


66.
Kaidarova A, Geraldi NR, Wilson RP, et al: Wearable sensors for monitoring marine environments and their inhabitants. Nat Biotechnol 2023;41:1208–1220. doi: 10.1038/s41587-023-01827-3


67.
Papastavrou V, Ryan C: Ethical standards for research on marine mammals. Res Ethics 2023;19:390–408. doi: 10.1177/17470161231182066


68.
Horton TW, Hauser N, Cassel S, et al: Non-invasive measurement of humpback whale vital signs using unoccupied aerial system infrared thermography. Front Mar Sci 2019;6:466. doi: 10.3389/fmars.2019.00466


69.
The Society for Marine Mammalogy: Guideline for Treatment of Marine Mammals. Available from: http://www.marinemammalscience.org/about-us/ethics/marine-mammaltreatment-guidelines/ [cited 29 March 2026].


70.
Freitas C, Villegas-Ríos D, Moland E, et al: Sea temperature effects on depth use and habitat selection in a marine fish community. J Anim Ecol 2021;90:1001–1013. doi: 10.1111/1365-2656.13497


71.
Albouy C, Delattre V, Donati G, et al: Global vulnerability of marine mammals to global warming. Sci Rep 2020;10:548. doi: 10.1038/s41598-019-57280-3


72.
Fedak MA, Thompson D: Behavioural and physiological options in diving seals. In: Boyd IL: editor. Physiology and bioenergetics. Marine Mammals: Advances in Behavioural and Population Biology: Proceedings of a Symposium held at The Zoological Society of London on 9th and 10th April 1992. Oxford, 1993; online edition, Oxford Academic, 31 October 2023.


73.
Xu X, Rioux TP, Castellani MP: Three dimensional models of human thermoregulation: a review. J Therm Biol 2023;112:103491. doi: 10.1016/j.jtherbio.2023.103491


74.
Fish FE, Kocak D: Biomimetics and marine technology: an introduction. Mar Technol Soc J 2011;45:8–13. doi: 10.4031/MTSJ.45.4.14


75.
Shi W, Atlar M, Norman R: Learning from humpback whales for improving the energy capturing performance of tidal turbine blades. In: Ölçer AI, Kitada M, Dalaklis D, et al: editors. Trends and Challenges in Maritime Energy Management. Charm; Springer International Publishing: 2018. p. 479–497.


76.
Anderson IA, Vincent J, Montgomery J: Biomimetics beneath the waves. In: Ocean Innovation. 1st edition, Boca Raton; CRC Press: 2016. p. 202.
Published
2026-10-01
How to Cite
Kasimanickam , R., & Kasimanickam , V. (2026). Reproductive thermoregulation in marine mammals: mechanisms, adaptations and evolutionary implications. Clinical Theriogenology, 18. https://doi.org/10.58292/CT.v18.14255
Section
Review Reports