Beale, T. The Natural History of the Sperm Whale (John Van Voorst, 1839).
Gordon, J. C. D. Sperm whale groups and social behaviour observed off Sri Lanka. Rep. – Int. Whal. Comm. 37, 205–217 (1987).
Whitehead, H. Babysitting, dive synchrony, and indications of alloparental care in sperm whales. Behav. Ecol. Sociobiol. 38, 237–244 (1996).
Whitehead, H. Sperm Whales: Social Evolution in the Ocean (University of Chicago Press, 2003).
Gero, S., Engelhaupt, D., Rendell, L. & Whitehead, H. Who cares? Between-group variation in alloparental caregiving in sperm whales. Behav. Ecol. 20, 838–843 (2009).
Gero, S., Gordon, J. & Whitehead, H. Calves as social hubs: dynamics of the social network within sperm whale units. Proc. Royal Soc. B: Biol. Sci. 280, 20131113 (2013).
Best, P. B. Social organization in sperm whales, Physeter macrocephalus. Behav. Mar. Anim. 227–289. https://doi.org/10.1007/978-1-4684-2985-5_7 (1979).
Weilgart, L. S. & Whitehead, H. Observations of a sperm whale (Physeter catodon) birth. J. Mammal. 67, 399–401 (1986).
Gambell, R., Lockyer, C. & Ross, G. J. B. Observations on the birth of a sperm whale calf. S Afr. J. Sci. 69, 147–148 (1973).
Correia-Fagundes, C. & Romano, H. Observation of a birth of a sperm whale Physeter macrocephalus at Madeira (NE Atlantic). Bocagiana (Funchal) 1–3 (2013).
Pervushin, A. S. Nabliudeniia Za Rodami u kasholotov. (Observations of delivery of sperm-whales). Zool. Zhurnal. 45, 1892–1893 (1966).
Gambell, R. Aerial observations of sperm whale behaviour based on observations, notes and comments by K.J. Pinkerton. Norsk Hvalfangst-Tidende (The Nor. Whal. Gazette). 57, 126–138 (1968).
Slijper, E. J. Some remarks on gestation and birth in cetacea and other aquatic mammals. Hvalrådets Skrifter: Sci. Results Mar. Biol. Research (1956).
Committee on Taxonomy. List of marine mammal species and subspecies, society for marine mammalogy. https://marinemammalscience.org/science-and-publications/list-marine-mammal-species-subspecies/ (2024).
Best, P., Canham, P. & Macleod, N. Patterns of reproduction in sperm whales, Physeter macrocephalus. Rep. Int. Whal. Comm. 6, 51–79 (1984).
Tønnesen, P., Gero, S., Ladegaard, M., Johnson, M. & Madsen, P. T. First-year sperm whale calves echolocate and perform long, deep dives. Behav Ecol. Sociobiol 72, (2018).
Ransome, N. et al. Observations of parturition in humpback whales (Megaptera novaeangliae) and occurrence of escorting and competitive behavior around birthing females. Mar. Mamm. Sci. 38, 408–432 (2022).
Silvers, L. E., Atkinson, S., Iwasa, M., Combelles, C. & Salden, D. R. A large placenta encountered in the Hawaiian winter grounds of the humpback whale, Megaptera novaeangliae. Mar. Mamm. Sci. 13, 711–716 (1997).
Dunston, D. J. Caudal presentation at birth of a humpback whale, Megaptera nodosa (Bonnaterre). Norsk Hvalfangst-Tidende (The Nor. Whal. Gazette) 553–555 (1957).
Ferreira, M. C. E., Maia-Nogueira, R. & De Jesus, A. Surface observation of a birth of a humpback whale (Megaptera novaeangliae) on the Northeast Coast of Brazil. Latin Am. J. Aquat. Mamm. 9, 160–163 (2011).
Faria, M. A., DeWeerdt, J., Pace, F. & Mayer, F. X. Observation of a humpback whale (Megaptera novaeangliae) birth in the coastal waters of Sainte Marie Island, Madagascar. Aquat. Mamm. 39, 296–305 (2013).
Patton, D. & Lawless, S. Surface and underwater observation of a humpback whale (Megaptera novaeangliae) birth in progress off Lahaina, Maui, and subsequent encounter of the female with a healthy calf. Aquat. Mamm. 47, 550–558 (2021).
Jacobsen, J. K. San Francisco,. The birth of a wild killer whale (Orcinus orca). in 4th Biennial Conference on the Biology of Marine Mammals 62 (1981).
Stacey, P. J. & Baird, R. W. Birth of a ‘resident’ killer whale off Victoria, British Columbia, Canada. Mar. Mamm. Sci. 13, 504–508 (1997).
Notarbartolo-Di-Sciara, G., Barbaccia, G. & Azzellino, A. Birth at sea of a false killer whale, Pseudorca crassidens. Mar. Mamm. Sci. 13, 508–511 (1997).
Beland, P., Faucher, A. & Corbeil, P. Observations on the birth of a beluga whale (Delphinapterus leucas) in the St Lawrence Estuary, Quebec, Canada. Can. J. Zool. 68, 1327–1329 (1990).
Lynase, M. Birth of a wild white whale. Whales Etcetera: Publication Connecticut Cetacean Soc. 1, 10–11 (1984).
Zani, M. A., Taylor, J. K. D. & Kraus, S. D. Observation of a right whale (Eubalaena glacialis) birth in the coastal waters of the Southeast united States. Aquat. Mamm. 34, 21–24 (2008).
Foley, H. J. et al. Observations of a Western North Atlantic right whale (Eubalaena glacialis) birth offshore of the protected Southeast U.S. Critical habitat. Mar. Mamm. Sci. 27, E234–E240 (2011).
Shuttleworth, L., Appleby, A., Appleby, R. & Vermeulen, E. First direct observation of a successful Southern right whale (Eubalaena australis) birth in South African coastal waters. Mar. Mamm. Sci. 40, e13091 (2024).
Balcomb, I. I. I. The birth of a gray whale. Pac. Discovery. 27, 28–31 (1974).
Leatherwood, S. & Beach, D. W. A California gray whale calf (Eschrichtius robustus) born outside the calving lagoons. Bull. South. Calif. Acad. Sci. 74, 45–46 (1975).
Mills, J. G. & Mills, J. E. Observations of a gray whale birth. Bull. South. Calif. Acad. Sci. 78, 192–196 (1979).
Perrtree, R. M. et al. First observed wild birth and acoustic record of a possible infanticide attempt on a common bottlenose dolphin (Tursiops truncatus). Mar. Mamm. Sci. 32, 376–385 (2016).
Gero, S. & Whitehead, H. Suckling behavior in sperm whale calves: observations and hypotheses. Mar. Mamm. Sci. 23, 398–413 (2007).
Gero, S., Gordon, J. & Whitehead, H. Calves as social hubs: dynamics of the social network within sperm whale units. Proc. Biol. Sci. 280, 20131113 (2013).
Gordon, J. et al. Distribution, movements and residency of sperm whales off the Commonwealth of Dominica, Eastern Caribbean: implications for the development and regulation of the local whalewatching industry. Rep. Int. Whal. Commn. 48, 551–557 (1998).
Gero, S. et al. Behavior and social structure of the sperm whales of Dominica, West Indies. Mar. Mamm. Sci. 30, 905–922 (2014).
Konrad, C. M., Gero, S., Frasier, T. & Whitehead, H. Kinship influences sperm whale social organization within, but generally not among, social units. R Soc. Open. Sci. 5, 180914 (2018).
Gero, S. & Whitehead, H. Critical decline of the Eastern Caribbean sperm whale population. PLoS One. 11, e0162019 (2016).
Gubnitsky, G., Mevorach, Y., Gero, S., Gruber, D. & Diamant, R. Automatic detection and annotation of Eastern Caribbean sperm whale codas. Sci. Rep. 15, 12790 (2025).
Leitao, A. et al. Evidence of social learning across symbolic cultural barriers in sperm whales. Elife 13, RP96362 (2024).
Begus, G., Sprouse, R., Leban, A., Silva, M. & Gero, S. Vowels and dipthong-like spectral patterns in sperm whale codas. Open Mind accepted (2025).
Schulz, T. M. The Production and Exchange of Sperm Whale Coda Vocalizations (Dalhousie University, 2007).
Trevathan, W. R. Human birth: an evolutionary perspective. Hum. Birth: Evolutionary Perspective. https://doi.org/10.4324/9780203789599 (2017).
Blanchet, M. A., Nance, T., Ast, C., Wahlberg, M. & Acquarone, M. First case of a monitored pregnancy of a harbour porpoise (Phocoena phocoena) under human care. Aquat. Mamm. 34, 9–20 (2008).
Hague, E. L., McCaffrey, N., Stockin, K. A. & Orbach, D. N. Previously undocumented long-finned pilot whale (Globicephala melas) placental expulsion in coastal waters of Shetland, United Kingdom. Aquat. Mamm. 48, 610–616 (2022).
Darwin, C. On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (John Murray, 1859).
Thewissen, J. G. M. & Bajpai, S. Whale origins as a poster child for macroevolution. Bioscience 51, 1037–1049 (2001).
Uhen, M. D. The origin(s) of whales. Annu. Rev. Earth Planet. Sci. 38, 189–219 (2010).
Gatesy, J. et al. A phylogenetic blueprint for a modern whale. Mol. Phylogenet Evol. 66, 479–506 (2013).
McGowen, M. R., Gatesy, J. & Wildman, D. E. Molecular evolution tracks macroevolutionary transitions in Cetacea. Trends Ecol. Evol. 29, 336–346 (2014).
Gero, S., Whitehead, H. & Rendell, L. Individual, unit and vocal clan level identity cues in sperm whale codas. R Soc. Open. Sci. https://doi.org/10.1098/rsos.150372 (2016).
Hersh, T. A. et al. Evidence from sperm whale clans of symbolic marking in non-human cultures. Proc. Natl. Acad. Sci. U S A. 119, e2201692119 (2022).
Smultea, M. et al. Cetacean mother-calf behavior observed from a small aircraft off Southern California. Anim. Behav. Cogn. 4, 1–23 (2017).
Nielsen, M. L. K., Sprogis, K. R., Bejder, L., Madsen, P. T. & Christiansen, F. Behavioural development in Southern right whale calves. Mar. Ecol. Prog Ser. 629, 219–234 (2019).
McGowen, M. R., Spaulding, M. & Gatesy, J. Divergence date Estimation and a comprehensive molecular tree of extant cetaceans. Mol. Phylogenet Evol. 53, 891–906 (2009).
Miller, P. J. O., Johnson, M. P., Tyack, P. L. & Terray, E. A. Swimming gaits, passive drag and buoyancy of diving sperm whales Physeter macrocephalus. J. Exp. Biol. 207, 1953–1967 (2004).
Slijper, E. J. Some remarks on gestation and birth in cetacea and other aquatic mammals. Hvalrådets Skrifter: Sci. Results Mar. Biol. Res. 41, 1–56 (1956).
Glarou, M. et al. Estimating body mass of sperm whales from aerial photographs. Mar. Mamm. Sci. 39, 251–273 (2023).
Sato, K. et al. Stroke frequency, but not swimming speed, is related to body size in free-ranging seabirds, pinnipeds and cetaceans. Proc. Royal Soc. B: Biol. Sci. 274, 471–477 (2007).
Lambert, O., Bianucci, G. & De Muizon, C. Macroraptorial sperm whales (Cetacea, Odontoceti, Physeteroidea) from the Miocene of Peru. Zool. J. Linn. Soc. 179, 404–474 (2017).
Gubbins, C., Mcowan, B., Lynn, S. K., Hooper, S. & Reiss, D. Mother-infant spatial relations in captive bottlenose dolphins, Tursiops truncatus. Mar. Mamm. Sci. 15, 751–765 (1999).
Noren, S. R., Biedenbach, G. & Edwards, E. F. Ontogeny of swim performance and mechanics in bottlenose dolphins (Tursiops truncatus). J. Exp. Biol. 209, 4724–4731 (2006).
Uhen, M. D. Evolution of marine mammals: back to the sea after 300 million years. Anat. Rec. 290, 514–522 (2007).
Heyning, J. E., Lento, G. M. & Hoelzel, A. R. The Evolution of Marine Mammals. in Marine Mammal Biology: An Evolutionary Approach 38–72 (2002).
Gingerich, P. D. et al. New protocetid whale from the middle Eocene of Pakistan: birth on land, precocial development, and sexual dimorphism. PLoS One. 4, e4366 (2009).
Cooper, L. N., Suydam, R. & Thewissen, J. G. M. Cetacean Evolution: Copulatory and Birthing Consequences of Pelvic and Hindlimb Reduction. in Sex in Cetaceans 65–83Springer International Publishing, https://doi.org/10.1007/978-3-031-35651-3_4 (2023).
Reidenberg, J. S. & Laitman, J. T. Cetacean prenatal development. in Encyclopedia of Marine Mammals 220–230 (Elsevier, doi:https://doi.org/10.1016/B978-0-12-373553-9.00056-0. (2009).
Gol’Din, P. Case of cephalic presentation of foetus in a harbour porpoise Phocoena phocoena (Cetacea, Phocoenidae), with notes on other aquatic mammals. Vestn Zool. 45, 34–38 (2011).
Marsh, H., Heinsohn, G., Marsh, L., Breeding & Cycle Life history and population dynamics of the dugong, Dugon dugon (Sirenia: dugongidae. Aust J. Zool. 32, 767 (1984).
Fernández, M., Morteo, E., Delfín-Alfonso, C. A. & Hernández-Candelario, I. Harassing behavior by short-finned pilot whales (Globicephala macrorhynchus) towards a mother and calf sperm whale (Physeter macrocephalus) pair. Aquat. Mamm. 48, 621–625 (2022).
Díaz-Gamboa, R. E. & Gendron, D. Guerrero-de La Rosa, F. Aggressive behavior of short-finned pilot whales towards sperm whales in the Gulf of California: insight into food competition. Aquat. Mamm. 48, 529–532 (2022).
Weller, D. W. et al. Observations of an interaction between sperm whales and short-finned pilot whales in the Gulf of Mexico. Mar. Mamm. Sci. 12, 588–594 (1996).
Pitman, R. L., Ballance, L. T., Mesnick, S. I. & Chivers, S. J. Killer whale predation on sperm whales: observations and implications. Mar. Mamm. Sci. 17, 494–507 (2001).
Whitehead, H. & Rendell, L. The cultural lives of whales and dolphins. Cult. Lives Whales Dolphins. https://doi.org/10.7208/chicago/9780226187426.001.0001 (2015).
Pitman, R. L. et al. Humpback whales interfering when mammal-eating killer whales attack other species: mobbing behavior and interspecific altruism? Mar. Mamm. Sci. 33, 7–58 (2017).
Whitehead, H. & Weilgart, L. The sperm whale: social females and roving males. in Cetacean Societies: Field Stud. Dolphins Whales 154–172 (2000).
Morange, Z., Similä, T., Rikardsen, A. H. & Blanchet, M. A. Seasonal changes in the feeding aggregation structure of male sperm whales (Physeter macrocephalus) in Northern Norway. Mar. Mamm. Sci. 41, e13219 (2024).
Kobayashi, H., Whitehead, H. & Amano, M. Long-term associations among male sperm whales (Physeter macrocephalus). PLoS One. 15, e0244204 (2020).
Ward-Paige, C. A. et al. Large-scale absence of sharks on reefs in the Greater-Caribbean: a footprint of human pressures. PLoS One. 5, e11968 (2010).
Blount, D. et al. Flukebook: an open-source AI platform for cetacean photo identification. Mammalian Biology. 102, 1005–1023 (2022).
Sharma, P. et al. Contextual and combinatorial structure in sperm whale vocalisations. Nat. Commun. 15, 3617 (2024).
Swofford, D. L. PAUP* Phylogenetic Analysis Using Parsimony * (and Other Methods). Version 4.0. Sinauer AssociatesSunderland, Massachusetts, (2002).
