Cetacea#Classification

{{More citations needed section|date=November 2019}}

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|image1=Eubalaena glacialis (North Atlantic right whale) 1 (30986325841).jpg

|caption1=Skull of the North Atlantic right whale (Mysticeti)

|image2=Orca Schaedel Senckenberg.jpg

|caption2=Skull of the orca (Odontoceti)

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The two parvorders, baleen whales (Mysticeti) and toothed whales (Odontoceti), are thought to have diverged around thirty-four million years ago.{{Cite journal|last1=Cerchio|first1=Salvatore|last2=Tucker|first2=Priscilla|date=1998-06-01|title=Influence of Alignment on the mtDNA Phylogeny of Cetacea: Questionable Support for a Mysticeti/Physeteroidea Clade|journal=Systematic Biology|volume=47|issue=2|pages=336–344|doi=10.1080/106351598260941|pmid=12064231|s2cid=16270218 |issn=1076-836X|doi-access=free}}

Baleen whales have bristles made of keratin instead of teeth. The bristles filter krill and other small invertebrates from seawater. Grey whales feed on bottom-dwelling mollusks. Rorqual family (balaenopterids) use throat pleats to expand their mouths to take in food and sieve out the water.{{Citation |last=Marshall |first=Christopher D. |title=Feeding Morphology |date=2018-01-01 |encyclopedia=Encyclopedia of Marine Mammals (Third Edition) |pages=349–354 |editor-last=Würsig |editor-first=Bernd |url=https://linkinghub.elsevier.com/retrieve/pii/B9780128043271001254 |access-date=2025-02-09 |publisher=Academic Press |doi=10.1016/b978-0-12-804327-1.00125-4 |isbn=978-0-12-804327-1 |editor2-last=Thewissen |editor2-first=J. G. M. |editor3-last=Kovacs |editor3-first=Kit M.}} Balaenids (right whales and bowhead whales) have massive heads that can make up 40% of their body mass. Most mysticetes prefer the food-rich colder waters of the Northern and Southern Hemispheres, migrating to the Equator to give birth. During this process, they are capable of fasting for several months, relying on their fat reserves.

The parvorder of Odontocetes – the toothed whales – include sperm whales, beaked whales, orcas, dolphins and porpoises. Generally their teeth have evolved to catch fish, squid or other marine invertebrates, not for chewing them, so prey is swallowed whole. Teeth are shaped like cones (dolphins and sperm whales), spades (porpoises), pegs (belugas), tusks (narwhals) or variable (beaked whale males). Female beaked whales' teeth are hidden in the gums and are not visible, and most male beaked whales have only two short tusks. Narwhals have vestigial teeth other than their tusk, which is present on males and 15% of females and has millions of nerves to sense water temperature, pressure and salinity. A few toothed whales, such as some orcas, feed on mammals such as pinnipeds and other whales.{{Cite news |last=Roth |first=Annie |date=2022-01-29 |title=Orcas Are Able to Kill and Eat Blue Whales, Scientists Confirm |url=https://www.nytimes.com/2022/01/29/science/orcas-blue-whales.html |access-date=2025-02-09 |work=The New York Times |language=en-US |issn=0362-4331}}{{Cite web |title=BBC Two - Expedition Killer Whale - Capturing the secret lives of pack ice killer whales |url=https://www.bbc.co.uk/programmes/articles/52HKYPGNzVgK62gbtQXz3mD/capturing-the-secret-lives-of-pack-ice-killer-whales |access-date=2025-02-09 |website=BBC |language=en-GB}}

Toothed whales have well-developed senses – their eyesight and hearing are adapted for both air and water, and they have advanced sonar capabilities using their melon. Their hearing is so well-adapted for both air and water that some blind specimens can survive. Some species, such as sperm whales, are well adapted for diving to great depths. Several species of toothed whales show sexual dimorphism, in which the males differ from the females, usually for purposes of sexual display or aggression.{{Citation |last1=Ralls |first1=Katherine |title=Sexual Dimorphism |date=2009-01-01 |encyclopedia=Encyclopedia of Marine Mammals (Second Edition) |pages=1005–1011 |editor-last=Perrin |editor-first=William F. |url=https://linkinghub.elsevier.com/retrieve/pii/B9780123735539002339 |access-date=2025-02-09 |place=London |publisher=Academic Press |doi=10.1016/b978-0-12-373553-9.00233-9 |isbn=978-0-12-373553-9 |last2=Mesnick |first2=Sarah |editor2-last=Würsig |editor2-first=Bernd |editor3-last=Thewissen |editor3-first=J. G. M.}}

File:Dolphin Anatomy.svg

Cetacean bodies are generally similar to those of fish, which can be attributed to their lifestyle and the habitat conditions. Their body is well-adapted to their habitat, although they share essential characteristics with other higher mammals (Eutheria).{{cite news |last1=Groves |last2=Colin |first3=Peter |last3=Grubb |title=Ungulate taxonomy |newspaper=JHU Press |year=2011}} {{page needed|date=April 2018}}

They have a streamlined shape, and their forelimbs are flippers. Almost all have a dorsal fin on their backs, but this can take on many forms, depending on the species. A few species, such as the beluga whale, lack them. Both the flipper and the fin are for stabilization and steering in the water.{{citation needed|date=October 2024}}

The male genitals and the mammary glands of females are sunken into the body.{{cite book |author=Thewissen, J.G.M. |title=The Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea |url=https://books.google.com/books?id=8QPyBwAAQBAJ&q=(penis+OR+genital)&pg=PA383 |date=11 November 2013 |publisher=Springer |isbn=978-1-4899-0159-0 |pages=383–}}{{cite book |author=Miller, Debra Lee |title=Reproductive Biology and Phylogeny of Cetacea: Whales, Porpoises and Dolphins |url=https://books.google.com/books?id=sD3NBQAAQBAJ |date=2007 |publisher=CRC Press |isbn=978-1-4398-4257-7}} The male genitals are attached to a vestigial pelvis.{{Cite book |last=Tinker |first=Spencer Wilkie |url=https://books.google.com/books?id=ASIVAAAAIAAJ&pg=PA93 |title=Whales of the World |date=1988-01-01 |publisher=Brill Archive |isbn=978-0-935848-47-2 |language=en}}

The body is wrapped in a thick layer of fat, known as blubber. This provides thermal insulation and gives cetaceans their smooth, streamlined body shape. In larger species, it can reach a thickness up to {{convert|1/2|m|ft|abbr=off|sp=us|spell=in}}.{{citation needed|date=October 2024}}

Sexual dimorphism evolved in many toothed whales. Sperm whales, narwhals, many members of the beaked whale family, several species of the porpoise family, orcas, pilot whales, eastern spinner dolphins and northern right whale dolphins show this characteristic.{{cite journal |last1=Dines |first1=James |last2=Mesnick |first2=Sarah |last3=Ralls |first3=Katherine |last4=May-Collado |first4=Laura |last5=Agnarsson |first5=Ingi |last6=Dean |first6=Matthew |title=A trade-off between precopulatory and postcopulatory trait investment in male cetaceans |journal=Evolution |date=2015 |volume=69 |issue=6 |pages=1560–1572 |doi=10.1111/evo.12676 |pmid=25929734|s2cid=18292677 }} Males in these species developed external features absent in females that are advantageous in combat or display. For example, male sperm whales are up to 63% percent larger than females, and many beaked whales possess tusks used in competition among males.{{cite journal |last1=Dalebout |first1=Merel |last2=Steel |first2=Debbie |last3=Baker |first3=Scott |title=Phylogeny of the Beaked Whale Genus Mesoplodon (Ziphiidae: Cetacea) Revealed by Nuclear Introns: Implications for the Evolution of Male Tusks|journal=Systematic Biology |date=2008 |volume=57 |issue=6 |pages=857–875 |doi=10.1080/10635150802559257 |pmid=19085329|s2cid=205729032 |doi-access=free }}

Hind legs are not present in cetaceans, nor are any other external body attachments such as a pinna and hair.{{Cite web|url=http://www.eurekalert.org/pub_releases/2006-05/uof-haw052206.php|title=How ancient whales lost their legs, got sleek and conquered the oceans|date=2006-05-22|website=EurekAlert|publisher=University of Florida|access-date=2016-03-20|archive-date=2020-12-02|archive-url=https://web.archive.org/web/20201202154340/https://www.eurekalert.org/pub_releases/2006-05/uof-haw052206.php|url-status=dead}}

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|image1=Beluga premier.gov.ru-1.jpeg

|caption1=Head profile of a beluga whale, featuring the large "melon" region

|image2=SBNMS - feeding humpback whale (31319806616).jpg

|caption2=Close up of a feeding humpback whale, showing baleen

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Whales have an elongated head, especially baleen whales, due to the wide overhanging jaw. Bowhead whale plates can be {{convert|9|m|ft}} long. Their nostril(s) make up the blowhole, with one in toothed whales and two in baleen whales.{{Cite journal|last1=Buono|first1=Mónica R.|last2=Fernández|first2=Marta S.|last3=Fordyce|first3=R. Ewan|last4=Reidenberg|first4=Joy S.|date=2015|title=Anatomy of nasal complex in the southern right whale, Eubalaena australis (Cetacea, Mysticeti)|journal=Journal of Anatomy|language=en|volume=226|issue=1|pages=81–92|doi=10.1111/joa.12250|issn=1469-7580|pmc=4313901|pmid=25440939}}

The nostrils are located on top of the head above the eyes so that the rest of the body can remain submerged while surfacing for air. The back of the skull is significantly shortened and deformed. By shifting the nostrils to the top of the head, the nasal passages extend perpendicularly through the skull.{{cite book|author=Milan Klima |title=Development of the Cetacean Nasal Skull|url=https://books.google.com/books?id=kOHAz9SCL54C |date=29 January 1999 |publisher=Springer |isbn=978-3-540-64996-0 }} {{page needed|date=April 2018}} The teeth or baleen in the upper jaw sit exclusively on the maxilla. The braincase is concentrated through the nasal passage to the front and is correspondingly higher, with individual cranial bones that overlap.{{citation needed|date=October 2024}}

In toothed whales, connective tissue exists in the melon as a head buckle. This is filled with air sacs and fat that aid in buoyancy and biosonar. The sperm whale has a particularly pronounced melon; this is called the spermaceti organ and contains the eponymous spermaceti, hence the name "sperm whale". Even the long tusk of the narwhal is a vice-formed tooth. In many toothed whales, the depression in their skull is due to the formation of a large melon and multiple, asymmetric air bags.{{Cite journal |last1=Cranford |first1=Ted W. |last2=Amundin |first2=Mats |last3=Norris |first3=Kenneth S. |date=1996 |title=Functional morphology and homology in the odontocete nasal complex: Implications for sound generation |url=https://onlinelibrary.wiley.com/doi/10.1002/(SICI)1097-4687(199606)228:3%3C223::AID-JMOR1%3E3.0.CO;2-3 |journal=Journal of Morphology |language=en |volume=228 |issue=3 |pages=223–285 |doi=10.1002/(SICI)1097-4687(199606)228:3<223::AID-JMOR1>3.0.CO;2-3 |pmid=8622183 |issn=1097-4687}}

River dolphins, unlike most other cetaceans, can turn their head 90°. Most other cetaceans have fused neck vertebrae and are unable to turn their head at all.{{citation needed|date=October 2024}}

The baleen of baleen whales consists of long, fibrous strands of keratin. Located in place of the teeth, it has the appearance of a huge fringe and is used to sieve the water for plankton and krill.{{Cite web |title=What is baleen? |url=https://us.whales.org/whales-dolphins/what-is-baleen/ |access-date=2025-01-13 |website=Whale & Dolphin Conservation USA |language=en-US}}

File:Preserved sperm whale brain.jpg, considered the largest brain in the world]]

Sperm whales have the largest brain mass of any animal on Earth, averaging {{convert|8,000|cm3|in3|abbr=on}} and {{convert|7.8|kg|lb|abbr=on}} in mature males.{{cite web | url=http://www.nmfs.noaa.gov/pr/species/mammals/cetaceans/spermwhale.htm |title=Sperm Whales brain size| publisher=NOAA Fisheries – Office of Protected Resources |access-date=9 August 2015}} The brain to body mass ratio in some odontocetes, such as belugas and narwhals, is second only to humans.{{cite magazine |url=http://blogs.scientificamerican.com/news-blog/are-whales-smarter-than-we-are/ |author=Fields, R. Douglas |title=Are whales smarter than we are? | magazine=Scientific American |access-date=9 August 2015}} In some whales, however, it is less than half that of humans: 0.9% versus 2.1%.{{citation needed|date=October 2024}}

In cetaceans, evolution in the water has caused changes to the head that have modified brain shape such that the brain folds around the insula and expands more laterally than in terrestrial mammals. As a result, the cetacean prefrontal cortex (compared to that in humans) rather than frontal is laterally positioned.{{cite journal |last1=Gerussi |first1=Tommaso |last2=Graïc |first2=Jean-Marie |last3=Peruffo |first3=Antonella |last4=Behroozi |first4=Mehdi |last5=Schlaffke |first5=Lara |last6=Huggenberger |first6=Stefan |last7=Güntürkün |first7=Onur |last8=Cozzi |first8=Bruno |date=2023 |title=The prefrontal cortex of the bottlenose dolphin (Tursiops truncatus Montagu, 1821): a tractography study and comparison with the human |journal=Brain Structure and Function |volume=228 |issue=8 |pages=1963–1976 |doi=10.1007/s00429-023-02699-8 |issn=1863-2661 |pmc=10517040 |pmid=37660322}}

Brain size was previously considered a major indicator of intelligence. Since most of the brain is used for maintaining bodily functions, greater ratios of brain to body mass may increase the amount of brain mass available for cognitive tasks. Allometric analysis of the relationship between mammalian brain mass (weight) and body mass for different species of mammals shows that larger species generally have larger brains. However, this increase is not fully proportional. Typically the brain mass only increases in proportion to somewhere between the two-thirds power (or the square of the cube root) and the three-quarters power (or the cube of the fourth root) of the body mass.

m_brain ∝ (m_body)^k

where k is between two-thirds and three-quarters. Thus if Species B is twice the size of Species A, its brain size will typically be somewhere between 60% and 70% higher.{{cite web |url=http://pages.ucsd.edu/~jmoore/courses/allometry/allometry.html |title=Allometry |first=Jim |last=Moore| publisher=University of California San Diego |access-date=9 August 2015}} Comparison of a particular animal's brain size with the expected brain size based on such an analysis provides an encephalization quotient that can be used as an indication of animal intelligence.{{cite book|last1=Pontarotti|first1=Pierre|title=Evolutionary Biology: Convergent Evolution, Evolution of Complex Traits|date=2016|publisher=Springer|isbn=978-3-319-41324-2|pages=74}}

The neocortex of many cetaceans is home to elongated spindle neurons that, prior to 2019, were known only in hominids.{{cite journal |author=Watson, K.K. |title=Dendritic architecture of the Von Economo neurons |journal=Neuroscience |volume=141 |issue=3 |pages=1107–1112 |year=2006 |doi=10.1016/j.neuroscience.2006.04.084 |last2=Jones |first2=T. K. |last3=Allman |first3=J. M. |pmid=16797136|s2cid=7745280 }} In humans, these cells are thought to be involved in social conduct, emotions, judgment and theory of mind.{{cite journal |author=Allman, John M. |title=Intuition and autism: a possible role for Von Economo neurons |journal=Trends Cogn Sci |volume=9 |issue=8 |pages=367–373 |year=2005 |doi=10.1016/j.tics.2005.06.008 |last2=Watson |first2=Karli K. |last3=Tetreault |first3=Nicole A. |last4=Hakeem |first4=Atiya Y. |pmid=16002323|s2cid=14850316 }} Cetacean spindle neurons are found in areas of the brain homologous to where they are found in humans, suggesting they perform a similar function.{{cite journal |author=Hof, Patrick R. |title=Structure of the cerebral cortex of the humpback whale, Megaptera novaeangliae (Cetacea, Mysticeti, Balaenopteridae) |journal=The Anatomical Record |volume=290 |issue=1 |pages=1–31 |year=2007 |doi=10.1002/ar.20407 |last2=Van Der Gucht |first2=Estel |pmid=17441195|s2cid=15460266 |doi-access=free }}

{{unreferenced section|date=April 2018}}

File:Whales skeletal system comparison.png

The cetacean skeleton is largely made up of cortical bone, which stabilizes the animal in the water. For this reason, the usual terrestrial compact bones, which are finely woven cancellous bone, are replaced with lighter and more elastic material. In many places, bone elements are replaced by cartilage and even fat, thereby improving their hydrostatic qualities. The ear and the muzzle contain a bone shape that is exclusive to cetaceans with a high density, resembling porcelain. This conducts sound better than other bones, thus aiding biosonar.

The number of vertebrae that make up the spine varies by species, ranging from forty to ninety-three. The cervical spine, found in all mammals, consists of seven vertebrae which, however, are reduced or fused. This fusion provides stability during swimming at the expense of mobility. The fins are carried by the thoracic vertebrae, ranging from nine to seventeen individual vertebrae. The sternum is cartilaginous. The last two to three pairs of ribs are not connected and hang freely in the body wall. The stable lumbar and tail include the other vertebrae. Below the caudal vertebrae is the chevron bone.

The front limbs are paddle-shaped with shortened arms and elongated finger bones, to support movement. They are connected by cartilage. The second and third fingers display a proliferation of the finger members, a so-called hyperphalangy. The shoulder joint is the only functional joint in all cetaceans except for the Amazon river dolphin. The collarbone is completely absent.

File:Humpback whale fluke (2).jpg

Cetaceans have a cartilaginous fluke at the end of their tails that is used for propulsion. The fluke is set horizontally on the body and used with vertical movements, unlike fish and ichthyosaurs, which have vertical tails which move horizontally.{{Cite web|title=Why do whale and dolphin tails go up and down?|url=https://us.whales.org/whales-dolphins/why-do-whale-and-dolphin-tails-go-up-and-down/|access-date=2021-12-23|website=Whale & Dolphin Conservation USA|language=en-US}}

Cetaceans have powerful hearts. Blood oxygen is distributed effectively throughout the body. They are warm-blooded, i.e., they hold a nearly constant body temperature.{{cite web |title=Whales, dolphins and porpoises |url=https://www.dcceew.gov.au/environment/marine/marine-species/cetaceans |website=Department of Climate Change, Energy, the Environment and Water |publisher=Australian Government Department of Climate Change, Energy, the Environment and Water |access-date=19 October 2024}}

{{unreferenced section|date=April 2018}}

Cetaceans have lungs, meaning they breathe air. An individual can last without a breath from a few minutes to over two hours depending on the species. Cetacea are deliberate breathers who must be awake to inhale and exhale. When stale air, warmed from the lungs, is exhaled, it condenses as it meets colder external air. As with a terrestrial mammal breathing out on a cold day, a small cloud of 'steam' appears. This is called the 'spout' and varies across species in shape, angle and height. Species can be identified at a distance using this characteristic.

The structure of the respiratory and circulatory systems is of particular importance for the life of marine mammals. The oxygen balance is effective. Each breath can replace up to 90% of the total lung volume. For land mammals, in comparison, this value is usually about 15%. During inhalation, about twice as much oxygen is absorbed by the lung tissue as in a land mammal. As with all mammals, the oxygen is stored in the blood and the lungs, but in cetaceans, it is also stored in various tissues, mainly in the muscles. The muscle pigment, myoglobin, provides an effective bond. This additional oxygen storage is vital for deep diving, since beyond a depth around {{convert|100|m|ft|abbr=on}}, the lung tissue is almost completely compressed by the water pressure.

The stomach consists of three chambers. The first region is formed by a loose gland and a muscular forestomach (missing in beaked whales); this is followed by the main stomach and the pylorus. Both are equipped with glands to help digestion. A bowel adjoins the stomachs, whose individual sections can only be distinguished histologically. The liver is large and separate from the gall bladder.{{cite book|author1=C. Edward Stevens|author2=Ian D. Hume|title=Comparative Physiology of the Vertebrate Digestive System|publisher=University of Cambridge|year=1995|access-date=5 September 2015|page=51|isbn=978-0-521-44418-7|url= https://books.google.com/books?id=DZuAsci2apAC&q=cetacean+digestive+system&pg=PR15}}

The kidneys are long and flattened. The salt concentration in cetacean blood is lower than that in seawater, requiring kidneys to excrete salt. This allows the animals to drink seawater.{{cite journal|author=Clifford A. Hui|title=Seawater Consumption and Water Flux in the Common Dolphin Delphinus delphis|journal=Chicago Journals|volume=54|issue=4|pages=430–440|year=1981|jstor= 30155836}} The urinary bladder is proportionally smaller in cetaceans than in land mammals.{{cite book|url=https://books.google.com/books?id=TNAIBwAAQBAJ&pg=PR35|title=The Works of John Hunter, F.R.S.|date=26 March 2015|publisher=Cambridge University|isbn=978-1-108-07960-0|page=35|author=John Hunter}} The testes are located internally, without an external scrotum.{{cite book|author1=Bernd Würsig|author2=J.G.M. Thewissen|author3=Kit M. Kovacs|title=Encyclopedia of Marine Mammals|url=https://books.google.com/books?id=T3zGDgAAQBAJ&q=testes+OR+testis+OR+testicles|date=27 November 2017|publisher=Elsevier Science|isbn=978-0-12-804381-3|access-date=19 October 2020|archive-date=29 June 2023|archive-url=https://web.archive.org/web/20230629161516/https://books.google.com/books?id=T3zGDgAAQBAJ&q=testes+OR+testis+OR+testicles|url-status=live}}{{cite book | last1 = Rommel | first1 = S.A. | first2 = D.A. | last2 = Pabst | first3 = W.A. | last3 = McLellan | chapter = Functional anatomy of the cetacean reproductive system, with comparisons to the domestic dog | title = Reproductive Biology and Phylogeny of Cetacea: Whales, Porpoises and Dolphins | date = 2007 | pages = 127–145 | editor-last = Miller | editor-first = D.L. | chapter-url = https://www.researchgate.net/publication/286334625 | doi = 10.1201/b11001| isbn = 9780429063626 }}{{cite magazine | last1 = Rommel | first1 = S.A. | first2 = D.A. | last2 = Pabst | first3 = W.A. | last3 = McLellan | title = Reproductive Thermoregulation in Marine Mammals | journal = American Scientist | volume = 86 | number = 5 | date = 1998 | pages = 440–448 | jstor = 27857097 | url = https://uncw.edu/mmsp/documents/rommeletal1998reproductivethermoregulation.pdf | url-status = live | archive-date = 22 November 2021 | archive-url = https://web.archive.org/web/20211122202231/https://uncw.edu/mmsp/documents/rommeletal1998reproductivethermoregulation.pdf}}{{cite book | last1 = Pabst | first1 = D.A. | last2 = Sentiel | first2 = A.R | first3 = W.A. | last3 = McLellan | chapter = Evolution of thermoregulatory function in cetacean reproductive systems | title = The Emergence of Whales | publisher = Springer US | date = 1998 | pages = 379–397 | series = Advances in Vertebrate Paleobiology | doi = 10.1007/978-1-4899-0159-0_13 | isbn = 978-1-4899-0161-3 | editor-last = Thewissen | editor-first = J.G.M.}} The uterus is bicornuate.

{{see also|Monochromacy}}

Cetacean eyes are set on the sides rather than the front of the head. This means only species with pointed 'beaks' (such as dolphins) have good binocular vision forward and downward. Tear glands secrete greasy tears, which protect the eyes from the salt in the water. The lens is almost spherical, which is most efficient at focusing the minimal light that reaches deep water.{{citation needed|date=February 2025}}{{clarify|the shape is necessary for focusing in water and has little to do with light levels. monochromacy is because the light is monochrome blue at depth|date=February 2025}}

Odontocetes have little to no ability to taste or smell, while mysticetes are believed to have some ability to smell because of their reduced, but functional olfactory system.{{Cite journal|last1=Godfrey|first1=Stephen J.|last2=Geisler|first2=Jonathan|last3=Fitzgerald|first3=Erich M. G.|date=2013|title=On the Olfactory Anatomy in an Archaic Whale (Protocetidae, Cetacea) and the Minke Whale Balaenoptera acutorostrata (Balaenopteridae, Cetacea)|journal=The Anatomical Record|language=en|volume=296|issue=2|pages=257–272|doi=10.1002/ar.22637|pmid=23233318|s2cid=25260840|issn=1932-8494|doi-access=free}}

At least one species, the tucuxi or Guiana dolphin, is able to use electroreception to sense prey.{{cite web |last=Morell |first=Virginia |title=Guiana Dolphins Can Use Electric Signals to Locate Prey |work=Science |publisher=American Association for the Advancement of Science (AAAS) |date=July 2011 |url=http://news.sciencemag.org/sciencenow/2011/07/guiana-dolphins-can-use-electric.html?ref=hp |url-status=dead |archive-url=https://web.archive.org/web/20130530040210/http://news.sciencemag.org/sciencenow/2011/07/guiana-dolphins-can-use-electric.html?ref=hp |archive-date=2013-05-30 }}

Cetaceans are known to possess excellent hearing.{{cite web |last1=Mead |first1=James |title=Cetacea |url=https://school.eb.com/levels/high/article/cetacean/105982 |website=Britannica School High |publisher=Encyclopædia Britannica, Inc. |access-date=3 June 2019}}

File:Delfinekko.gif ]]

The external ear has lost the pinna (visible ear), but still retains a narrow ear canal. The three small bones or ossicles that transmit sound within each ear are dense and compact, and differently shaped from those of land mammals. The semicircular canals are much smaller relative to body size than in other mammals.{{cite book | last=Thewissen | first=J. G. M. | chapter=Hearing | pages=[https://archive.org/details/encyclopediaofma2002unse/page/570 570–572] | title=Encyclopedia of Marine Mammals | editor1-last=Perrin | editor1-first=William R. | editor2-last=Wiirsig | editor2-first=Bernd | editor3-last=Thewissen | editor3-first=J. G. M. | year=2002 | publisher=Academic Press | isbn=978-0-12-551340-1 | chapter-url=https://archive.org/details/encyclopediaofma2002unse/page/570 }}

A bony structure of the middle and inner ear, the auditory bulla, is composed of two compact and dense bones (the periotic and tympanic). It is housed in a cavity in the middle ear; in the Odontoceti (apart from in the physeterids, this cavity is filled with dense foam{{clarify|what is the foam made of?|date=February 2025}} and completely surrounds the bulla, which is connected to the skull only by ligaments. This may isolate the ear from sounds transmitted through the bones of the skull, something that also happens in bats.{{cite book | last=Ketten | first=Darlene R. | chapter=The Marine Mammal Ear: Specializations for Aquatic Audition and Echolocation | pages=717–750 | title=The Evolutionary Biology of Hearing | editor1-last=Webster | editor1-first=Douglas B. | editor2-last=Fay | editor2-first=Richard R. | editor3-last=Popper | editor3-first=Arthur N. | year=1992 | publisher=Springer | chapter-url=https://www.researchgate.net/publication/230691464}} Pages 725–727 used here.

Cetaceans use sound to communicate, using groans, moans, whistles, clicks or the 'singing' of the humpback whale.

Odontoceti are generally capable of echolocation.{{cite book |editor1-last=Perrin |editor1-first=William F. |editor2-last=Wursig |editor2-first=Bernd |editor3-last=Thewissen|editor3-first=J. G. M. |title=Encyclopedia of Marine Mammals |last=Hooker |first=Sascha K. |edition=2 |publisher=Academic Press |page=1176 |year=2009 |isbn=978-0-12-373553-9 |url=https://books.google.com/books?id=2rkHQpToi9sC&q=Encyclopedia%20of%20Marine%20Mammals&pg=PP1}} They can discern the size, shape, surface characteristics, distance and movement of an object. They can search for, chase and catch fast-swimming prey in total darkness. Most Odontoceti can distinguish between prey and nonprey (such as humans or boats); captive Odontoceti can be trained to distinguish between, for example, balls of different sizes or shapes. Echolocation clicks also contain characteristic details unique to each animal, which may suggest that toothed whales can discern between their own click and that of others.{{cite journal |author = de Obaldia, C., Simkus, G. & and Zölzer, U. | title = Estimating the number of sperm whale (Physeter macrocephalus) individuals based on grouping of corresponding clicks

| journal = 41. Jahrestagung für Akustik (DAGA 2015), Nürnberg | year = 2015 | doi = 10.13140/RG.2.1.3764.9765 | url= https://www.researchgate.net/publication/277009691 }}

While differences in ear structure associated with echolocating abilities are found amongst Cetacea, cranial asymmetry has also been found to be a factor in the ability to produce sounds used in echolocation. Mysticeti, who don't have the ability to echolocate, possess general symmetry of the skull and facial region, while Odontoceti display a nasofacial asymmetry that is linked to their echolocating abilities.{{Cite journal |last1=Coombs |first1=Ellen J. |last2=Clavel |first2=Julien |last3=Park |first3=Travis |last4=Churchill |first4=Morgan |last5=Goswami |first5=Anjali |date=2020-07-10 |title=Wonky whales: the evolution of cranial asymmetry in cetaceans |journal=BMC Biology |volume=18 |issue=1 |pages=86 |doi=10.1186/s12915-020-00805-4 |doi-access=free |issn=1741-7007 |pmc=7350770 |pmid=32646447}} Differences in the level of asymmetry also seem to correlate with differences in the types of sounds produced.

Mysticeti have exceptionally thin, wide basilar membranes in their cochleae without stiffening agents, making their ears adapted for processing low to infrasonic frequencies.{{cite journal | last = Ketten | first = Darlene R. | title = Structure and function in whale ears | year = 1997 | journal = The International Journal of Animal Sound and Its Recording | volume = 8 | issue = 1–2 | pages = 103–135 | url = http://csi.whoi.edu/sites/default/files/literature/Full%20Text_29.pdf | doi = 10.1080/09524622.1997.9753356 | bibcode = 1997Bioac...8..103K | access-date = 2013-12-21 | archive-date = 2014-08-01 | archive-url = https://web.archive.org/web/20140801161859/http://csi.whoi.edu/sites/default/files/literature/Full%20Text_29.pdf | url-status = dead }}

The initial karyotype includes a set of chromosomes from 2n = 44. They have four pairs of telocentric chromosomes (whose centromeres sit at one of the telomeres), two to four pairs of subtelocentric and one or two large pairs of submetacentric chromosomes. The remaining chromosomes are metacentric—the centromere is approximately in the middle—and are rather small. All cetaceans have chromosomes 2n = 44, except the sperm whales and pygmy sperm whales, which have 2n = 42.{{cite journal|author=Ulfur Anarson|title=Comparative chromosome studies in Cetacea|journal=Institute of Genetics|year=1974|volume=77|issue=1|doi=10.1111/j.1601-5223.1974.tb01351.x|pages=1–36|pmid=4137586|doi-access=free}}

{{see also|Cetaceans of the Caribbean}}

Cetaceans are found in many aquatic habitats. While many marine species, such as the blue whale, the humpback whale and the orca, have a distribution area that includes nearly the entire ocean, some species occur only locally or in broken populations. These include the vaquita, which inhabits a small part of the Gulf of California and Hector's dolphin, which lives in some coastal waters in New Zealand. Most river dolphin species live exclusively in fresh water.{{cite journal|last1=Cassens|first1=I.|last2=Vicario|first2=S.|last3=Waddell|first3=V.G.|last4=Balchowsky|first4=H.|last5=Van Belle|first5=D.|last6=Ding|first6=W.|last7=Fan|first7=C.|last8=Mohan|first8=R.S.|last9=Simões-Lopes|first9=P.C.|last10=Bastida|first10=R.|last11=Meyer|first11=A.|last12=Stanhope|first12=M.J.|last13=Milinkovitch|first13=M.C.|title=Independent adaptation to riverine habitats allowed survival of ancient cetacean lineages|journal=Proceedings of the National Academy of Sciences of the United States of America|date=2000|volume=97|issue=21|pages=11343–11347|doi=10.1073/pnas.97.21.11343|pmid=11027333|pmc=17202|bibcode=2000PNAS...9711343C|doi-access=free}}

Many species inhabit specific latitudes, often in tropical or subtropical waters, such as Bryde's whale or Risso's dolphin. Others are found only in a specific body of water. The southern right whale dolphin and the hourglass dolphin live only in the Southern Ocean. The narwhal and the beluga live only in the Arctic Ocean. Sowerby's beaked whale and the Clymene dolphin exist only in the Atlantic and the Pacific white-sided dolphin and the northern straight dolphin live only in the North Pacific.{{citation needed|date=October 2024}}

Cosmopolitan species may be found in the Pacific, Atlantic and Indian Oceans. However, northern and southern populations become genetically separated over time. In some species, this separation leads eventually to a divergence of the species, such as produced the southern right whale, North Pacific right whale and North Atlantic right whale.{{cite journal|author=AR Hoelzel|title=Genetic structure of cetacean populations in sympatry, parapatry, and mixed assemblages: implications for conservation policy|journal=Journal of Heredity|year=1998|doi=10.1093/jhered/89.5.451|volume=89|issue=5|pages=451–458|doi-access=free}} Migratory species' reproductive sites often lie in the tropics and their feeding grounds in polar regions.

Thirty-two species are found in European waters, including twenty-five toothed and seven baleen species.{{citation needed|date=October 2024}}

Many species of whales migrate on a latitudinal basis to move between seasonal habitats. For example, the gray whale migrates {{convert|10,000|mi}} round trip. The journey begins at winter birthing grounds in warm lagoons along Baja California, and traverses {{convert|5,000|-|7,000|mi}} of coastline to summer feeding grounds in the Bering, Chuckchi and Beaufort seas off the coast of Alaska.{{cite web |url=https://journeynorth.org/tm/gwhale/annual/map.html |title=Gray Whale Migration |website=journeynorth.org |access-date=3 July 2021 |url-status=live|archive-url=https://web.archive.org/web/20190609231527/https://journeynorth.org/tm/gwhale/annual/map.html |archive-date=2019-06-09 }}

Conscious breathing cetaceans sleep but cannot afford to be unconscious for long, because they may drown. While knowledge of sleep in wild cetaceans is limited, toothed cetaceans in captivity have been recorded to exhibit unihemispheric slow-wave sleep (USWS), which means they sleep with one side of their brain at a time, so that they may swim, breathe consciously and avoid both predators and social contact during their period of rest.{{cite journal | vauthors = Sekiguchi Y, Arai K, Kohshima S | title = Sleep behaviour: sleep in continuously active dolphins | journal = Nature | volume = 441 | issue = 7096 | pages = E9-10; discussion E11 | date = June 2006 | pmid = 16791150 | doi = 10.1038/nature04898 | s2cid = 4406032 | bibcode = 2006Natur.441E...9S }}

A 2008 study found that sperm whales sleep in vertical postures just under the surface in passive shallow 'drift-dives', generally during the day, during which whales do not respond to passing vessels unless they are in contact, leading to the suggestion that whales possibly sleep during such dives.{{cite journal | vauthors = Miller PJ, Aoki K, Rendell LE, Amano M | title = Stereotypical resting behavior of the sperm whale | journal = Current Biology | volume = 18 | issue = 1 | pages = R21-3 | date = January 2008 | pmid = 18177706 | doi = 10.1016/j.cub.2007.11.003 | s2cid = 10587736 | doi-access = free | bibcode = 2008CBio...18..R21M }}

{{see also|Physiology of underwater diving#Mammals}}

While diving, the animals reduce their oxygen consumption by lowering the heart activity and blood circulation; individual organs receive no oxygen during this time. Some rorquals can dive for up to 40 minutes, sperm whales between 60 and 90 minutes and bottlenose whales for two hours. Diving depths average about {{convert|100|m|ft|abbr=on}}. Species such as sperm whales can dive to {{convert|3,000|m|ft|abbr=on}}, although more commonly {{convert|1,200|m|ft}}.{{cite journal|last=Scholander|first=Per Fredrik|title=Experimental investigations on the respiratory function in diving mammals and birds|year=1940|journal=Hvalraadets Skrifter|volume=22}}{{cite journal|author1=Bruno Cozzi|author2=Paola Bagnoli|author3=Fabio Acocella|author4=Maria Laura Costantino|title=Structure and biomechanical properties of the trachea of the striped dolphin Stenella coeruleoalba: Evidence for evolutionary adaptations to diving|journal=The Anatomical Record|volume=284|issue=1|pages=500–510|year=2005|doi=10.1002/ar.a.20182|pmid=15791584}}

Most cetaceans are social animals, although a few species live in pairs or are solitary. A group, known as a pod, usually consists of ten to fifty animals, but on occasion, such as mass availability of food or during mating season, groups may encompass more than one thousand individuals. Inter-species socialization can occur.{{cite book|editor1=Janet Mann|editor2=Richard C. Connor|editor3=Peter L. Tyack|editor4=Hal Whitehead| display-editors = 3|title=Cetacean Societies: Field Study of Dolphins and Whales|publisher=University of Chicago}}

Pods have a fixed hierarchy, with the priority positions determined by biting, pushing or ramming. The behavior in the group is aggressive only in situations of stress such as lack of food, but usually it is peaceful. Contact swimming, mutual fondling and nudging are common. The playful behavior of the animals, which is manifested in air jumps, somersaults, surfing, or fin hitting, occurs more often than not in smaller cetaceans, such as dolphins and porpoises.

Males in some baleen species communicate via whale song, sequences of high pitched sounds. These "songs" can be heard for hundreds of kilometers. Each population generally shares a distinct song, which evolves over time. Sometimes, an individual can be identified by its distinctive vocals, such as the 52-hertz whale that sings at a higher frequency than other whales. Some individuals are capable of generating over 600 distinct sounds. In baleen species such as humpbacks, blues and fins, male-specific song is believed to be used to attract and display fitness to females.{{cite journal |last1=Janik |first1=Vincent |title=Cetacean vocal learning and communication |journal=Current Opinion in Neurobiology |date=2014 |volume=28 |pages=60–65 |doi= 10.1016/j.conb.2014.06.010|pmid=25057816 |s2cid=40334723 }}

Pod groups also hunt, often with other species. Many species of dolphins accompany large tunas on hunting expeditions, following large schools of fish. The orca hunts in pods and targets belugas and even larger whales. Humpback whales, among others, form in collaboration bubble carpets to herd krill or plankton into bait balls before lunging at them.

{{Main|Cetacean intelligence}}File:Whales Bubble Net Feeding-edit1.jpg

Cetacea are known to teach, learn, cooperate, scheme and grieve.{{cite news |last=Siebert |first=Charles |title=Watching Whales Watching Us |work=The New York Times Magazine |date=8 July 2009 |access-date=29 August 2015 |url=https://www.nytimes.com/2009/07/12/magazine/12whales-t.html?pagewanted=all}}

Smaller cetaceans, such as dolphins and porpoises, engage in complex play behavior, including such things as producing stable underwater toroidal air-core vortex rings or "bubble rings". The two main methods of bubble ring production are rapid puffing of air into the water and allowing it to rise to the surface, forming a ring, or swimming repeatedly in a circle and then stopping to inject air into the helical vortex currents thus formed. They also appear to enjoy biting the vortex rings, so that they burst into many separate bubbles and then rise quickly to the surface. Whales produce bubble nets to aid in herding prey.{{cite journal |author1=Wiley, David|s2cid=55168063 |title=Underwater components of humpback whale bubble-net feeding behaviour |journal=Behaviour |volume=148 |issue=5 |pages=575–602 |year=2011 |doi=10.1163/000579511X570893 |display-authors=etal}}

File:Orca porpoising.jpg]]

Larger whales are also thought to engage in play. The southern right whale elevates its tail fluke above the water, remaining in the same position for a considerable time. This is known as "sailing". It appears to be a form of play and is most commonly seen off the coast of Argentina and South Africa.{{cite book |last1=Carwardine |first1=M. H. |last2=Hoyt |first2=E. |year=1998 |title=Whales, Dolphins and Porpoises |journal=NSW: Reader's Digest |isbn=978-0-86449-096-4}} Humpback whales also display this behaviour.{{Cite web |author=A. O. L. Staff |date=2016-07-15 |title=Humpback whale 'tail-sails' as she watches her calf off the Maui coast |url=https://www.aol.com/article/2016/03/11/humpback-whale-tail-sails-as-she-watches-her-calf-off-maui-coast/21326396/?guccounter=1 |access-date=2025-02-08 |website=www.aol.com |language=en-US}}

Self-awareness appears to be a sign of abstract thinking. Self-awareness, although not well-defined, is believed to be a precursor to more advanced processes such as metacognitive reasoning (thinking about thinking) that humans exploit. Dolphins appear to possess self-awareness.{{cite web | url=http://www.livescience.com/4272-elephant-awareness-mirrors-humans.html | title=Elephant Self-Awareness Mirrors Humans |publisher=Live Science |date=30 October 2006 |access-date=29 August 2015}} The most widely used test for self-awareness in animals is the mirror test, in which a temporary dye is placed on an animal's body and the animal is then presented with a mirror. Researchers then explore whether the animal shows signs of self-recognition.{{cite news |url=https://www.nytimes.com/2001/05/01/science/brainy-dolphins-pass-the-human-mirror-test.html |title=Mirror test |author=Derr, Mark |newspaper=New York Times |access-date=3 August 2015|date=May 2001 }}

Critics claim that the results of these tests are susceptible to the Clever Hans effect. This test is much less definitive than when used for primates. Primates can touch the mark or the mirror, while dolphins cannot, making their alleged self-recognition behavior less certain. Skeptics argue that behaviors said to identify self-awareness resemble existing social behaviors, so researchers could be misinterpreting self-awareness for social responses. Advocates counter that the behaviors are different from normal responses to another individual. Dolphins show less definitive behavior of self-awareness, because they have no pointing ability.

In 1995, Marten and Psarakos used video to test dolphin self-awareness.{{cite journal|last1=Marten|first1=Ken|last2=Psarakos|first2=Suchi|title=Using Self-View Television to Distinguish between Self-Examination and Social Behavior in the Bottlenose Dolphin (Tursiops truncatus)|journal=Consciousness and Cognition|volume=4|number=2|date=June 1995|doi=10.1006/ccog.1995.1026|pages=205–224|pmid=8521259|s2cid=44372881}} They showed dolphins real-time footage of themselves, recorded footage and another dolphin. They concluded that their evidence suggested self-awareness rather than social behavior. While this particular study has not been replicated, dolphins later "passed" the mirror test.

Collective decisions are an important part of life as a cetacean for the many species that spend time in groups (whether these be temporary such as the fission-fusion dynamics of many smaller dolphin species or long-term stable associations as are seen in killer whale and sperm whale matrilines).{{cite journal | vauthors = Zwamborn E, Boon N, Whitehead H | title = Collective Decision-making in Aquatic Mammals | journal = Mammal Review| volume = 53 | issue = 4 | pages = 238–253| date = October 2023 | doi = 10.1111/mam.12321 | s2cid = 261141293 | doi-access = free | bibcode = 2023MamRv..53..238Z }} Little is known about how these decisions work, though studies have found evidence messy consensus decisions in groups of sperm whales and leadership in other species like bottlenose dolphins and killer whales.{{citation needed|date=October 2024}}

Most cetaceans sexually mature at seven to 10 years. An exception to this is the La Plata dolphin, which is sexually mature at two years, but lives only to about 20. The sperm whale reaches sexual maturity within about 20 years and has a lifespan between 50 and 100 years.

For most species, reproduction is seasonal. Ovulation coincides with male fertility. This cycle is usually coupled with seasonal movements that can be observed in many species. Most toothed whales have no fixed bonds. In many species, females choose several partners during a season. Baleen whales are largely monogamous within each reproductive period.{{citation needed|date=October 2024}}

Gestation ranges from 9 to 16 months. Duration is not necessarily a function of size. Porpoises and blue whales gestate for about 11 months. As with all mammals other than marsupials and monotremes, the embryo is fed by the placenta, an organ that draws nutrients from the mother's bloodstream. Mammals without placentas either lay minuscule eggs (monotremes) or bear minuscule offspring (marsupials).{{citation needed|date=October 2024}}

Cetaceans usually bear one calf. In the case of twins, one usually dies, because the mother cannot produce sufficient milk for both. In modern cetaceans, the fetus is usually positioned for a tail-first delivery. Contrary to popular belief, this is not to minimize the risk of drowning during delivery. More likely it has to do with the mechanics of birthing and the shape of the fetus.{{Cite journal |last1=Miedema |first1=Feiko |last2=Klein |first2=Nicole |last3=Blackburn |first3=Daniel G. |last4=Sander |first4=P. Martin |last5=Maxwell |first5=Erin E. |last6=Griebeler |first6=Eva M. |last7=Scheyer |first7=Torsten M. |date=2023-04-18 |title=Heads or tails first? Evolution of fetal orientation in ichthyosaurs, with a scrutiny of the prevailing hypothesis |journal=BMC Ecology and Evolution |volume=23 |issue=1 |pages=12 |doi=10.1186/s12862-023-02110-4 |doi-access=free |issn=2730-7182 |pmc=10114408 |pmid=37072698}} After birth, the mother carries the infant to the surface for its first breath. At birth, they are about one-third of their adult length and tend to be independently active, comparable to terrestrial mammals.{{citation needed|date=October 2024}}

Like other placental mammals, cetaceans give birth to well-developed calves and nurse them with milk from their mammary glands. When suckling, the mother actively splashes milk into the mouth of the calf, using the muscles of her mammary glands, as the calf has no lips. This milk usually has a high-fat content, ranging from 16 to 46%, causing the calf to increase rapidly in size and weight.

In many small cetaceans, suckling lasts for about four months. In large species, it lasts for over a year and involves a strong bond between mother and offspring.{{citation needed|date=October 2024}}

In several species of both whales and dolphins, alloparenting has been observed.{{Cite journal |last1=Konrad |first1=Christine M |last2=Frasier |first2=Timothy R |last3=Whitehead |first3=Hal |last4=Gero |first4=Shane |date=2019-03-04 |title=Kin selection and allocare in sperm whales |url=https://academic.oup.com/beheco/article-abstract/30/1/194/5144394?redirectedFrom=fulltext |journal=Behavioral Ecology |volume=30 |issue=1 |pages=194–201 |doi=10.1093/beheco/ary143 |issn=1045-2249}}{{Cite news |last=Weston |first=Phoebe |date=2023-03-10 |title='Extraordinary' sighting of orca with baby pilot whale astounds scientists |url=https://www.theguardian.com/environment/2023/mar/10/killer-whale-orca-adopts-abducts-pilot-whale-calf-aoe |access-date=2025-02-09 |work=The Guardian |language=en-GB |issn=0261-3077}}

This reproductive strategy provides a few offspring that have a high survival rate.{{citation needed|date=October 2024}}

Among cetaceans, whales are distinguished by an unusual longevity compared to other higher mammals. Some species, such as the bowhead whale (Balaena mysticetus), can reach over 200 years. Based on the annual rings of the bony otic capsule, the age of the oldest known specimen is a male determined to be 211 years at the time of death.{{cite journal|author1=John C George|author2=Jeffrey Bada|author3=Judith Zeh|author3-link= Judy Zeh |author4=Laura Scott|author5=Stephen E Brown|author6=Todd O'Hara|author7=Robert Suydam|title=Age and growth estimates of bowhead whales (Balaena mysticetus) via aspartic acid racemization|journal=Canadian Journal of Zoology|year=1999|volume=77|issue=4|doi=10.1139/z99-015|pages=571–580|bibcode=1999CaJZ...77..571G }}

{{Main|Whale fall}}

Upon death, whale carcasses fall to the deep ocean and provide a substantial habitat for marine life. Evidence of whale falls in present-day and fossil records shows that deep-sea whale falls support a rich assemblage of creatures, with a global diversity of 407 species, comparable to other neritic biodiversity hotspots, such as cold seeps and hydrothermal vents.{{cite book |url=http://www.soest.hawaii.edu/oceanography/faculty/csmith/Files/Smith%20and%20Baco%202003.pdf |archive-url=https://web.archive.org/web/20060817174541/http://www.soest.hawaii.edu/oceanography/faculty/csmith/Files/Smith%20and%20Baco%202003.pdf |archive-date=2006-08-17 |url-status=live |title=Ecology of Whale Falls at the Deep-Sea Floor |author1=Smith, Craig R. |author2=Baco, Amy R. |volume=41 |date=2003 |pages=311–354 |access-date=23 August 2014 |doi=10.1201/9780203180594.ch6|series=Oceanography and Marine Biology - an Annual Review |doi-broken-date=2024-11-11 |isbn=978-0-415-25462-5 }}

Deterioration of whale carcasses happens through three stages. Initially, organisms such as sharks and hagfish scavenge the soft tissues at a rapid rate over a period of months and as long as two years. This is followed by the colonization of bones and surrounding sediments (which contain organic matter) by enrichment opportunists, such as crustaceans and polychaetes, throughout a period of years. Finally, sulfophilic bacteria reduce the bones releasing hydrogen sulfide enabling the growth of chemoautotrophic organisms, which in turn, support organisms such as mussels, clams, limpets and sea snails. This stage may last for decades and supports a rich assemblage of species, averaging 185 per site.{{cite journal |author1=Fujiwara, Yoshihiro|title=Three-year investigations into sperm whale-fall ecosystems in Japan |journal=Marine Ecology |volume=28 |issue=1 |pages=219–230|date=16 February 2007|display-authors=etal |doi=10.1111/j.1439-0485.2007.00150.x|bibcode=2007MarEc..28..219F|doi-access=free}}

Brucellosis affects almost all mammals. It is distributed worldwide, while fishing and pollution have caused porpoise population density pockets, which risks further infection and disease spreading. Brucella ceti, most prevalent in dolphins, has been shown to cause chronic disease, increasing the chance of failed birth and miscarriages, male infertility, neurobrucellosis, cardiopathies, bone and skin lesions, strandings and death. Until 2008, no case had ever been reported in porpoises, but isolated populations have an increased risk and consequentially a high mortality rate.{{cite journal|first1=Guzmán-Verri|last1=Caterina| first2=Rocío|last2=González-Barrientos|first3=Gabriela|last3=Hernández-Mora|first4=Juan-Alberto|last4=Morales|first5=Elías|last5=Baquero-Calvo|first6=Esteban|last6=Chaves-Olarte|first7=Edgardo|last7=Moreno|title=Brucella ceti and Brucellosis in Cetaceans|journal=Frontiers in Cellular and Infection Microbiology|volume=2|pages=3|year=2012|doi=10.3389/fcimb.2012.00003|pmid=22919595|pmc=3417395|doi-access=free}}

{{main|Evolution of cetaceans}}

The direct ancestors of today's cetaceans are probably found within the Dorudontidae whose most famous member, Dorudon, lived at the same time as Basilosaurus. Both groups had already developed some of the typical anatomical features of today's whales, such as the fixed bulla, which replaces the mammalian eardrum, as well as sound-conducting elements for submerged directional hearing. Their wrists were stiffened and probably contributed to the typical build of flippers. The hind legs existed, however, but were significantly reduced in size and with a vestigial pelvis connection.

File:Adult female and fetal Maiacetus.jpg

The fossil record traces the gradual transition from terrestrial to aquatic life. The regression of the hind limbs allowed greater flexibility of the spine. This made it possible for whales to move around with the vertical tail hitting the water. The front legs transformed into flippers, costing them their mobility on land.{{cite web |url=https://evolution.berkeley.edu/what-are-evograms/the-evolution-of-whales/ |title=The Evolution of Whales |publisher=Understanding Evolution, University of California Museum of Paleontology |accessdate=21 January 2025}}

One of the oldest members of ancient cetaceans (Archaeoceti) is Pakicetus from the Middle Eocene of Pakistan. This is an animal the size of a wolf, whose skeleton is known only partially. It had functioning legs and lived near the shore. This suggests the animal could still move on land. The long snout had carnivorous dentition.

The transition from land to sea dates to about 49 million years ago, with the Ambulocetus ("running whale"), also discovered in Pakistan. It was up to {{convert|3|m|ft|abbr=on}} long. The limbs of this archaeocete were leg-like, but it was already fully aquatic, indicating that a switch to a lifestyle independent from land happened extraordinarily quickly.{{Cite journal |last1=Ando |first1=Konami |last2=Fujiwara |first2=Shin-ichi |date=2016-07-10 |title=Farewell to life on land - thoracic strength as a new indicator to determine paleoecology in secondary aquatic mammals |journal=Journal of Anatomy |volume=229 |issue=6 |pages=768–777 |doi=10.1111/joa.12518 |issn=0021-8782 |pmc=5108153 |pmid=27396988}} The snout was elongated with overhead nostrils and eyes. The tail was strong and supported movement through water. Ambulocetus probably lived in mangroves in brackish water and fed in the riparian zone as a predator of fish and other vertebrates.{{cite journal |last=Thewissen |first=Hans |author-link=Hans Thewissen |year=1994 |title=Phylogenetic aspects of Cetacean origins: A morphological perspective |journal=Journal of Mammalian Evolution |pages=157–184 |doi=10.1007/bf01473527 |volume=2 |issue=3 |s2cid=27675176}}

Dating from about 45 million years ago are species such as Indocetus, Kutchicetus, Rodhocetus and Andrewsiphius, all of which were adapted to life in water. The hind limbs of these species were regressed and their body shapes resemble modern whales. Protocetidae family member Rodhocetus is considered the first to be fully aquatic. The body was streamlined and delicate with extended hand and foot bones. The merged pelvic lumbar spine was present, making it possible to support the floating movement of the tail. It was likely a good swimmer, but could probably move only clumsily on land, much like a modern seal.

Since the late Eocene, about 40 million years ago, cetaceans populated the subtropical oceans and no longer emerged on land. An example is the 18 metre long Basilosaurus, sometimes called Zeuglodon. The transition from land to water was completed in about 10 million years. The Wadi Al-Hitan ("Whale Valley") in Egypt contains numerous skeletons of Basilosaurus, as well as other marine vertebrates.{{cite web |title=First intact fossil of prehistoric whale discovered in Wadi Al-Hitan |url=https://www.iucn.org/content/first-intact-fossil-prehistoric-whale-discovered-wadi-al-hitan |publisher=International Union for Conservation of Nature |date=9 June 2015}}

{{anchor|Classification}}

Molecular biology, immunology, and fossils show that cetaceans are phylogenetically closely related with the even-toed ungulates (Artiodactyla). Whales' direct lineage began in the early Eocene, around 55.8 million years ago, with early artiodactyls. Most molecular biological evidence suggests that hippos are the closest living relatives. Common anatomical features include similarities in the morphology of the posterior molars, and the bony ring on the temporal bone (bulla) and the involucre, a skull feature that was previously associated only with cetaceans.{{cite journal |last1=Thewissen|first1=J. G. M. |last2=Cooper |first2=Lisa Noelle|last3=Clementz |first3=Mark T. |last4=Bajpai |first4=Sunil |last5=Tiwari |first5=B. N. |title=Whales originated from aquatic artiodactyls in the Eocene epoch of India |year=2007 |journal=Nature |volume=450 |issue=7173 |pages=1190–4 |url=http://repository.ias.ac.in/4642/1/316.pdf |archive-url=https://web.archive.org/web/20131224104529/http://repository.ias.ac.in/4642/1/316.pdf |archive-date=2013-12-24 |url-status=live |doi=10.1038/nature06343 |pmid=18097400 |bibcode=2007Natur.450.1190T |s2cid=4416444 |author1-link=Hans Thewissen }} Since the fossil record suggests that the morphologically distinct hippo lineage dates back only about 15 million years, Cetacea and hippos apparently diverged from a common ancestor that was morphologically distinct from either.{{cite journal|title = More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen|author = Gatesy, J.|journal = Molecular Biology and Evolution|date=1 May 1997| volume = 14|pages = 537–543|pmid = 9159931|issue = 5|doi = 10.1093/oxfordjournals.molbev.a025790|doi-access = free}}{{cite journal |author1=Boisserie, Jean-Renaud |author2=Lihoreau, Fabrice |author3=Brunet, Michel |year= 2005|title= The position of Hippopotamidae within Cetartiodactyla|journal= Proceedings of the National Academy of Sciences |volume= 102 |issue= 5|pages= 1537–1541|doi= 10.1073/pnas.0409518102|pmid= 15677331 |pmc=547867|bibcode= 2005PNAS..102.1537B|doi-access=free }}{{cite web|title = Scientists find missing link between the dolphin, whale and its closest relative, the hippo|date = 2005-01-25|access-date = 2011-01-08|url = http://www.innovations-report.com/html/reports/life_sciences/report-39309.html|work = Science News Daily}} The most striking common feature is the talus, a bone in the upper ankle. Early cetaceans, archaeocetes, show double castors, which occur only in even-toed ungulates. Corresponding findings are from Tethys Sea deposits in northern India and Pakistan. The Tethys Sea was a shallow sea between the Asian continent and northward-bound Indian plate.{{citation needed|date=October 2023}}

File:Ichthyosaur vs dolphin.svg as streamlined swimmers with fish and aquatic reptiles. ]]

Molecular and morphological evidence suggests that artiodactyls as traditionally defined are paraphyletic with respect to cetaceans. Cetaceans are deeply nested within the artiodactyls; the two groups together form a clade, a natural group with a common ancestor, for which the name Cetartiodactyla is sometimes used. Modern nomenclature divides Artiodactyla (or Cetartiodactyla) into four subordinate taxa: camelids (Tylopoda), pigs and peccaries (Suina), ruminants (Ruminantia), and hippos plus whales (Whippomorpha). The Cetacea's presumed location within Artiodactyla can be represented in the following cladogram:

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|last19= Spaulding |first19= M. |last20= Velazco |first20= P.M.

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|last3= Hasegawa |first3= M. |last4= Asher |first4= R.J.

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{{cite journal

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}} (see e.g. Fig S10)

{{Clade

|style=font-size:100%;line-height:100%;width:820px;

|label1=Artiodactyla

|1={{Clade

|1=Tylopoda (camels)50 px

|label2=Artiofabula

|2={{Clade

|1=Suina (pigs)50 px

|label2=Cetruminantia

|2={{Clade

|label1=Ruminantia (ruminants)

|1={{Clade

|1=Tragulidae (mouse deer)50 px

|2=Pecora (horn bearers)50 px

}}

|label2=Cetancodonta/Whippomorpha

|sublabel2=54 mya

|2={{Clade

|1=Hippopotamidae (hippopotamuses)50 px

|2=Cetacea (whales)50 px

}}

}}

}}

}}

}}

{{Further|List of cetaceans|List of extinct cetaceans}}

Within Cetacea, the two parvorders are baleen whales (Mysticeti) which owe their name to their baleen, and toothed whales (Odontoceti), which have teeth shaped like cones, spades, pegs, or tusks, and can perceive their environment through biosonar.{{citation needed|date=October 2024}}

{{Cladogram

|title=Relationship of extinct and extant cetaceans

:

|caption={{center|† Extinct taxa}}

|align=left

|cladogram=

{{Clade

|style=font-size:85%;line-height:100%;width:580px;

|label1=

|1={{Clade

|1=Raoellidae †

|label2=Cetacea

|2={{Clade

|1=Pakicetidae †

|2={{Clade

|1=Ambulocetidae † 50 px

|2={{Clade

|1=Remingtonocetidae † 50 px

|2={{Clade

|1=Rodhocetus †

|2={{Clade

|1=Dorudon †

|2={{Clade

|1=Basilosaurus † 50 px

|2={{Clade

|label1=Mysticeti

|sublabel1=(baleen whales, 25 mya)

|1={{Clade

|1=Janjucetus †

|2={{Clade

|label1=Balaenidae

|1=Right whales 50 px

|2={{Clade

|label1=Neobalaenidae

|1=Pygmy right whale 50 px

|2={{Clade

|label1=Balaenopteridae

|1=Rorquals 50 px

|label2=Eschrichtiidae

|2=Gray whales 50 px

}}

}}

}}

}}

|label2=Odontoceti

|sublabel2=(toothed whales)

|2={{Clade

|1={{Clade

|label1=Kogiidae

|1=Dwarf sperm whales 50 px

|label2=Physeteridae

|2=Sperm whales 50 px

}}

|2={{Clade

|label1=Platanistidae

|1=South Asian river dolphins 50 px

|2={{Clade

|label1=Ziphiidae

|1=Beaked whales 50 px

|2={{Clade

|label1=Inioidea

|sublabel1=(South American
river dolphins)

|1={{Clade

|label1=Lipotidae

|1=Yangtze river dolphin 50 px

|2={{Clade

|1=Iniidae 50 px

|2=Pontoporiidae 50 px

}}

}}

|label2=Delphinoidea

|2={{Clade

|label1=Delphinidae

|sublabel1=("true" dolphins)

|1=Oceanic dolphins 50 px

|2={{Clade

|label1=Monodontidae

|1=Belugas & Narwhals 50 px

|label2=Phocoenidae

|2=Porpoises 50 px

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

{{clear}}

The terms whale and dolphin are informal:

• Mysticeti:

:*Whales, with four families: Balaenidae (right and bowhead whales), Cetotheriidae (pygmy right whales), Balaenopteridae (rorquals), Eschrichtiidae (grey whales)

• Odontoceti:

:*Whales: with four families: Monodontidae (belugas and narwhals), Physeteridae (sperm whales), Kogiidae (dwarf and pygmy sperm whales), and Ziphiidae (beaked whales)

:*Dolphins, with five families: Delphinidae (oceanic dolphins), Platanistidae (South Asian river dolphins), Lipotidae (old world river dolphins) Iniidae (new world river dolphins), and Pontoporiidae (La Plata dolphins)

:*Porpoises, with one family: Phocoenidae

The term 'great whales' covers those currently regulated by the International Whaling Commission:{{cite web |title=Small cetaceans |website=iwc.int |publisher=International Whaling Commission |url=https://iwc.int/smallcetacean |access-date=2018-04-08}} the Odontoceti families Physeteridae (sperm whales), Ziphiidae (beaked whales), and Kogiidae (pygmy and dwarf sperm whales); and Mysticeti families Balaenidae (right and bowhead whales), Cetotheriidae (pygmy right whales), Eschrichtiidae (grey whales), as well as part of the family Balaenopteridae (minke, Bryde's, sei, blue and fin; not Eden's and Omura's whales).{{Cite web |title=Lives of Whales |website=iwc.int |publisher=International Whaling Commission |url=https://iwc.int/lives |access-date=2018-04-08}}

The primary threats to cetaceans come from people, both directly from whaling or drive hunting and indirect threats from fishing and pollution.{{cite book|author=Cara E. Miller|title=Current State of Knowledge of Cetacean Threats, Diversity, and Habitats in the Pacific Islands Region|publisher=Whale and Dolphin Conservation Society|year=2007|access-date=5 September 2015|isbn=978-0-646-47224-9|url=http://uk.whales.org/sites/default/files/whales-and-dolphins-in-the-pacific-islands.pdf|archive-url=https://web.archive.org/web/20150908003502/http://uk.whales.org/sites/default/files/whales-and-dolphins-in-the-pacific-islands.pdf|archive-date=8 September 2015|url-status=dead}}

{{Main|Whaling|History of Whaling|Dolphin drive hunting}}

{{multiple image

| direction = vertical

| header = Methods of whaling

| header_align = center

| width = 220

| caption_align = center

| image1 = Japan Factory Ship Nisshin Maru Whaling Mother and Calf.jpg

| alt1 = Mother and calf minke whales are pulled through the rear of a Japanese research ship.

| caption1 = Japanese research ship whaling mother and calf minke whales.

| image2 = Hvalba 26-08-06 (3).jpg

| alt2 = A dolphin caught in a drive hunt is transported away by a forklift.

| caption2 = An Atlantic white-sided dolphin caught in a drive hunt in Hvalba on the Faroe Islands being taken away with a forklift.

| align =

| total_width =

}}

Whaling is the practice of hunting whales, mainly baleen and sperm whales. This activity has gone on since the Stone Age.{{Cite web |date=2024-10-11 |title=Whaling - Commercial, Industrial, Japan {{!}} Britannica |url=https://www.britannica.com/topic/whaling |access-date=2024-10-28 |website=Britannica Online Encyclopedia |language=en}}

In the Middle Ages, reasons for whaling included their meat, oil usable as fuel and the jawbone, which was used in house construction. At the end of the Middle Ages, early whaling fleets aimed at baleen whales, such as bowheads. In the 16th and 17th centuries, the Dutch fleet had about 300 whaling ships with 18,000 crewmen.{{citation needed|date=October 2024}}

In the 18th and 19th centuries, baleen whales especially were hunted for their baleen, which was used as a replacement for wood, or in products requiring strength and flexibility such as corsets and crinoline skirts. In addition, the spermaceti found in the sperm whale was used as a machine lubricant and the ambergris as a material for pharmaceutical and perfume industries. In the second half of the 19th century, the explosive harpoon was invented, leading to a massive increase in the catch size.{{citation needed|date=October 2024}}

Large ships were used as "mother" ships for the whale handlers. In the first half of the 20th century, whales were of great importance as a supplier of raw materials. Whales were intensively hunted during this time; in the 1930s, 30,000 whales were killed. This increased to over 40,000 animals per year up to the 1960s, when stocks of large baleen whales collapsed.{{citation needed|date=October 2024}}

Most hunted whales are now threatened, with some great whale populations exploited to the brink of extinction. Atlantic and Korean gray whale populations were completely eradicated and the North Atlantic right whale population fell to some 300–600. The blue whale population is estimated to be around 14,000.{{citation needed|date=October 2024}}

The first efforts to protect whales came in 1931. Some particularly endangered species, such as the humpback whale (which then numbered about 100 animals),{{Cite web |title=A History of the International Whaling Commission (IWC) |url=https://wwf.panda.org/discover/knowledge_hub/endangered_species/cetaceans/cetaceans/iwc/history |access-date=2024-08-08 |website=wwf.panda.org |language=en}} were placed under international protection and the first protected areas were established. In 1946, the International Whaling Commission (IWC) was established, to monitor and secure whale stocks. Whaling of 14 large species for commercial purposes was prohibited worldwide by this organization from 1985 to 2005, though some countries do not honor the prohibition.{{citation needed|date=October 2024}}

File:Whales caught recently.png

The stocks of species such as humpback and blue whales have recovered, though they are still threatened. The United States Congress passed the Marine Mammal Protection Act of 1972 sustain the marine mammal population. It prohibits the taking of marine mammals except for several hundred per year taken in Alaska. Japanese whaling ships are allowed to hunt whales of different species for ostensibly scientific purposes.{{citation needed|date=October 2024}}

Aboriginal whaling is still permitted. About 1,200 pilot whales were taken in the Faroe Islands in 2017,{{Cite web |url=http://heimabeiti.fo/default.asp?menu=400 |title=Hagar & seyðamark |website=heimabeiti.fo |access-date=2018-04-07 |archive-date=2014-09-24 |archive-url=https://web.archive.org/web/20140924041544/http://www.heimabeiti.fo/default.asp?menu=400 |url-status=dead }} and about 900 narwhals and 800 belugas per year are taken in Alaska, Canada, Greenland, and Siberia. About 150 minke are taken in Greenland per year, 120 gray whales in Siberia and 50 bowheads in Alaska, as aboriginal whaling, besides the 600 minke taken commercially by Norway, 300 minke and 100 sei taken by Japan and up to 100 fin whales taken by Iceland.{{Cite web |url=https://iwc.int/total-catches |title=Total Catches |website=iwc.int |language=en |access-date=2018-04-07}} Iceland and Norway do not recognize the ban and operate commercial whaling. Norway and Japan are committed to ending the ban.{{citation needed|date=October 2024}}

Dolphins and other smaller cetaceans are sometimes hunted in an activity known as dolphin drive hunting. This is accomplished by driving a pod together with boats, usually into a bay or onto a beach. Their escape is prevented by closing off the route to the ocean with other boats or nets. Dolphins are hunted this way in several places around the world, including the Solomon Islands, the Faroe Islands, Peru and Japan (the most well-known practitioner). Dolphins are mostly hunted for their meat, though some end up in dolphinaria. Despite the controversy thousands of dolphins are caught in drive hunts each year.{{citation needed|date=October 2024}}

{{See also|Cetacean bycatch}}

File:Domino whale-bone hg.jpg made of baleen]]

Dolphin pods often reside near large tuna shoals. This is known to fishermen, who look for dolphins to catch tuna. Dolphins are much easier to spot from a distance than tuna, since they regularly breathe. The fishermen pull their nets hundreds of meters wide in a circle around the dolphin groups, in the expectation that they will net a tuna shoal. When the nets are pulled together, the dolphins become entangled under water and drown. Line fisheries in larger rivers are threats to river dolphins.{{citation needed|date=October 2024}}

A greater threat than by-catch for small cetaceans is targeted hunting. In Southeast Asia, they are sold as fish-replacement to locals, since the region's edible fish promise higher revenues from exports. In the Mediterranean, small cetaceans are targeted to ease pressure on edible fish.

{{Main|Cetacean stranding}}

A stranding is when a cetacean leaves the water to lie on a beach. In some cases, groups of whales strand together. The best known are mass strandings of pilot whales and sperm whales. Stranded cetaceans usually die, because their as much as {{convert|90|MT|ST}} body weight compresses their lungs or breaks their ribs. Smaller whales can die of heatstroke because of their thermal insulation.{{citation needed|date=October 2024}}

File:Ecomare - gestrande bultrug op Razende Bol (bultrug2012-razende-bol-412-sw).jpg

The causes are not clear. Possible reasons for mass beachings are:

• toxic contaminants
• debilitating parasites (in the respiratory tract, brain or middle ear)
• infections (bacterial or viral)
• flight from predators (including humans)
• social bonds within a group, so that the pod follows a stranded animal
• disturbance of their magnetic senses by natural anomalies in the Earth's magnetic field
• injuries
• noise pollution by shipping traffic, seismic surveys and military sonar experiments

Since 2000, whale strandings frequently occurred following military sonar testing. In December 2001, the US Navy admitted partial responsibility for the beaching and the deaths of several marine mammals in March 2000. The coauthor of the interim report stated that animals killed by active sonar of some Navy ships were injured. Generally, underwater noise, which is still on the increase, is increasingly tied to strandings; because it impairs communication and sense of direction.{{cite journal |author=Schrope, Mark. |title=Whale deaths caused by US Navy's sonar |journal=Nature |volume=415 |issue=106 |pages=106 |year=2003 |doi=10.1038/415106a|bibcode = 2002Natur.415..106S |pmid=11805797|s2cid=52827761 |doi-access=free }}

Climate change influences the major wind systems and ocean currents, which also lead to cetacean strandings. Researchers studying strandings on the Tasmanian coast from 1920 to 2002 found that greater strandings occurred at certain time intervals. Years with increased strandings were associated with severe storms, which initiated cold water flows close to the coast. In nutrient-rich, cold water, cetaceans expect large prey animals, so they follow the cold water currents into shallower waters, where the risk is higher for strandings. Whales and dolphins who live in pods may accompany sick or debilitated pod members into shallow water, stranding them at low tide.{{Cite journal |last1=Evans |first1=K |last2=Thresher |first2=R |last3=Warneke |first3=R.M |last4=Bradshaw |first4=C.J.A |last5=Pook |first5=M |last6=Thiele |first6=D |last7=Hindell |first7=M.A |date=2005-06-22 |title=Periodic variability in cetacean strandings: links to large-scale climate events |journal=Biology Letters |volume=1 |issue=2 |pages=147–150 |doi=10.1098/rsbl.2005.0313 |issn=1744-9561 |pmc=1626231 |pmid=17148151}}

{{See also|Marine mammals and sonar}}

{{Quote box|quote=Worldwide, use of active sonar has been linked to about 50 marine mammal strandings between 1996 and 2006. In all of these occurrences, there were other contributing factors, such as unusual (steep and complex) underwater geography, limited egress routes, and a specific species of marine mammal—beaked whales—that are suspected to be more sensitive to sound than other marine mammals.

|source=—Rear Admiral Lawrence Rice|align=left|width=30%}}

Heavy metals, residues of many plant and insect venoms and plastic waste flotsam are not biodegradable. Sometimes, cetaceans consume these hazardous materials, mistaking them for food items. As a result, the animals are more susceptible to disease and have fewer offspring.

Damage to the ozone layer reduces plankton reproduction because of its resulting radiation. This shrinks the food supply for many marine animals, but the filter-feeding baleen whales are most impacted. Even the Nekton is, in addition to intensive exploitation, damaged by the radiation.

Food supplies are also reduced long-term by ocean acidification due to increased absorption of increased atmospheric carbon dioxide. The CO₂ reacts with water to form carbonic acid, which reduces the construction of the calcium carbonate skeletons of food supplies for zooplankton that baleen whales depend on.

The military and resource extraction industries operate strong sonar and blasting operations. Marine seismic surveys use loud, low-frequency sound that show what is lying underneath the Earth's surface.{{cite journal|last1=Nowacek|first1=Douglas|last2=Donovan|first2=Greg|last3=Gailey|first3=Glenn|last4=Racca|first4=Roberto|last5=Reeves|first5=Randall|last6=Vedenev|first6=Alexander|last7=Weller|first7=David|last8=Southall|first8=Brandon|title=Responsible Practices for Minimizing and Monitoring Environmental Impacts of Marine Seismic Surveys with an Emphasis on Marine Mammal|journal=Aquatic Mammals|date=2013|volume=39|issue=4 |pages=356–377|doi=10.1578/am.39.4.2013.356|bibcode=2013AqMam..39..356N }} Vessel traffic also increases noise in the oceans. Such noise can disrupt cetacean behavior such as their use of biosonar for orientation and communication. Severe instances can panic them, driving them to the surface. This leads to bubbles in blood gases and can cause decompression sickness.{{Cite book|author1=M. Andre|title=Europe Oceans 2005|author2=T. Johansson|author3=E. Delory|author4=M. van der Schaar|publisher=Oceans 2005–Europe|year=2005|volume=2|pages=1028–1032 Vol. 2|doi=10.1109/OCEANSE.2005.1513199|chapter=Cetacean biosonar and noise pollution|isbn=978-0-7803-9103-1|s2cid=31676969}} Naval exercises with sonar regularly results in fallen cetaceans that wash up with fatal decompression. Sounds can be disruptive at distances of more than {{convert|100|km|mi}}. Damage varies across frequency and species.{{citation needed|date=October 2024}}

{{See also|Cryptid whale|Whale#In myth, literature and art}}

File:Trolual (Gessner).jpg

In Aristotle's time, the fourth century BCE, whales were regarded as fish due to their superficial similarity. Aristotle, however, observed many physiological and anatomical similarities with the terrestrial vertebrates, such as blood (circulation), lungs, uterus and fin anatomy.{{Cite book |last=Aristotle |url=https://penelope.uchicago.edu/aristotle/histanimals8.html |title=The History of Animals, Book VIII |language=en |translator-last=Thompson |translator-first=D'Arcy Wentworth |chapter=Chapter 2 |access-date=April 16, 2022 |archive-url=https://archive.today/20220416102355/https://penelope.uchicago.edu/aristotle/histanimals8.html |archive-date=April 16, 2022}} His detailed descriptions were assimilated by the Romans, but mixed with a more accurate knowledge of the dolphins, as mentioned by Pliny the Elder in his Natural history. In the art of this and subsequent periods, dolphins are portrayed with a high-arched head (typical of porpoises) and a long snout. The harbour porpoise was one of the most accessible species for early cetologists; because it could be seen close to land, inhabiting shallow coastal areas of Europe. Much of the findings that apply to all cetaceans were first discovered in porpoises.{{cite book|author= Conrad Gesner|url=http://etext.lib.virginia.edu/toc/modeng/public/AriHian.html|title=Historiae animalium |date=6 September 2008 |archive-url= https://web.archive.org/web/20080906090248/http://etext.lib.virginia.edu/toc/modeng/public/AriHian.html |archive-date=6 September 2008 |access-date=4 September 2015}} One of the first anatomical descriptions of the airways of a harbor porpoise dates from 1671 by John Ray. It nevertheless referred to the porpoise as a fish.{{cite journal|author=J. Ray|title=An account of the dissection of a porpess|journal=Philosophical Transactions of the Royal Society of London|volume=6|issue=69–80|year=1671|pages=2274–2279|bibcode = 1671RSPT....6.2274R|doi=10.1098/rstl.1671.0048|s2cid=186210473|doi-access=free}}{{cite journal|author=Susanne Prahl|title=Studies for the construction of epicranialen airway when porpoise (Phocoena phocoena Linnaeus, 1758)|journal=Dissertation for the Doctoral Degree of the Department of Biology of the Faculty of Mathematics, Computer Science and Natural Sciences at the University of Hamburg|year=2007|page=6}}

{{Blockquote|The tube in the head, through which this kind fish takes its breath and spitting water, located in front of the brain and ends outwardly in a simple hole, but inside it is divided by a downward bony septum, as if it were two nostrils; but underneath it opens up again in the mouth in a void.|John Ray, 1671, the earliest description of cetacean airways}}

In the 10th edition of Systema Naturae (1758), Swedish biologist and taxonomist Carl Linnaeus asserted that cetaceans were mammals and not fish. His groundbreaking binomial system formed the basis of modern whale classification.{{citation needed|date=October 2024}}

{{More citations needed section|date=August 2024}}

File:Destruction of Leviathan.png, 1865]]

File:Tarentum.jpg riding a dolphin]]

Stone Age petroglyphs, such as those in Roddoy and Reppa (Norway), and the Bangudae Petroglyphs in South Korea, depict them.{{cite web| url = http://www.pcas.org/Vol36N2/11Meighan.pdf| title = PCAS Quarterly - Rock Art on the Channel Islands of California}}{{cite news| url = http://news.bbc.co.uk/1/hi/sci/tech/3638853.stm| title = BBC News - Rock Art Hints at Whaling Origins| date = 20 April 2004}} Whale bones were used for many purposes. In the Neolithic settlement of Skara Brae on Orkney sauce pans were made from whale vertebrae.{{citation needed|date=October 2024}} The whale was first mentioned in ancient Greece by Homer. There, it is called Ketos, from which was derived the Roman word for whale, Cetus. In the Bible especially, the leviathan plays a role as a sea monster. The prophet Jonah, on his flight from the city of Nineveh, is swallowed by a whale.

Dolphins are mentioned far more often than whales. Aristotle discusses the sacred animals of the Greeks in his Historia Animalium. The Greeks admired the dolphin as a "king of the aquatic animals" and referred to them erroneously as fish. Dolphins appear in Greek mythology. Because of their intelligence, they rescued multiple people from drowning. They were said to love music, probably because of their own song, and in the legends they saved famous musicians, such as Arion of Lesbos from Methymna. Dolphins belong to the domain of Poseidon and led him to his wife Amphitrite. Dolphins are associated with other gods, such as Apollo, Dionysus and Aphrodite. The Greeks paid tribute to both whales and dolphins with their own constellation. The constellation of the Whale (Ketos, lat. Cetus) is located south of the Dolphin (Delphi, lat. Delphinus) north of the zodiac. Ancient art often included dolphin representations, including the Cretan Minoans. A particularly popular representation is that of Arion or Taras riding on a dolphin. In early Christian art, the dolphin is a popular motif, at times used as a symbol of Christ.

{{See also|History of Whaling|Cryptid whales}}

File:La Baleine.jpg

File:Im Februar 1598 an der holländischen Küste gestrandeter Walfisch.jpg

St. Brendan described in his travel story Navigatio Sancti Brendani an encounter with a whale, between the years 565–573.{{citation needed|date=October 2024}} Most descriptions of large whales from the Middle Ages until the whaling era, beginning in the 17th century, were of beached whales. Raymond Gilmore documented seventeen sperm whales in the estuary of the Elbe from 1723 to 1959 and thirty-one animals on the coast of Great Britain in 1784. In 1827, a blue whale beached itself off the coast of Ostend. Whales were used as attractions in museums and traveling exhibitions.{{citation needed|date=October 2024}}

Whalers from the 17th to 19th centuries depicted whales in drawings and recounted tales of their occupation. Although they knew that whales were harmless giants, they described battles with harpooned animals. These included descriptions of sea monsters, including huge whales, sharks, sea snakes, giant squid and octopuses.{{citation needed|date=October 2024}} Among the first whalers who described their experiences on whaling trips was Captain William Scoresby from Great Britain, who published the book Northern Whale Fishery, describing the hunt for northern baleen whales. This was followed by Thomas Beale, a British surgeon, in his book Some observations on the natural history of the sperm whale in 1835; and Frederick Debell Bennett's The tale of a whale hunt in 1840. Whales were described in narrative literature and paintings, most famously in the novels Moby Dick by Herman Melville and Twenty Thousand Leagues Under the Seas by Jules Verne.{{citation needed|date=October 2024}}

Baleen was used to make vessel components such as the bottom of a bucket in the Scottish National Museum. The Norsemen crafted ornamented plates from baleen.{{citation needed|date=October 2024}} In the Canadian Arctic (east coast) in Punuk and Thule culture (1000–1600 C.E.),{{cite journal|title=The circumpolar zone|last1=Cunliffe|first1=B.|last2=Gosden|first2=C.|last3=Joyce|first3=R.|journal=The Oxford Handbook of Archaeology}} baleen was used to construct houses in place of wood as roof support for winter houses.{{cite journal|author=J. Savelle| title=The Role of Architectural utility in the formation of archaeological Whale Bone Assemblages|journal=Journal of Archaeological Science|volume=24| issue=10|year=1997|pages=869–885|doi=10.1006/jasc.1996.0167| bibcode=1997JArSc..24..869S}}

{{Further|Whale#Interactions with humans}}

File:Sea World1.jpgs and false killer whales]]

In the 20th century, perceptions of cetaceans changed. They transformed from monsters into creatures of wonder, as science revealed them to be intelligent and peaceful animals. Hunting was replaced by whale and dolphin tourism. This change is reflected in films and novels. For example, the protagonist of the series Flipper was a bottle-nose dolphin. The TV series SeaQuest DSV (1993–1996), the movies Free Willy and Star Trek IV: The Voyage Home, and the book series The Hitchhiker's Guide to the Galaxy by Douglas Adams are examples.{{cite web|url=http://www.movieretriever.com/videohound_lists/90895/Whales|title=Movie Retriever: Whales|author=unknown|work=movieretriever.com|url-status=dead|archive-url=https://web.archive.org/web/20151015214719/http://www.movieretriever.com/videohound_lists/90895/Whales|archive-date=2015-10-15}}

The study of whale songs also produced a popular album, Songs of the Humpback Whale.{{Cite web |last=O'Dell |first=Cary |title="Songs of the Humpback Whale" (1970) |url=https://www.loc.gov/static/programs/national-recording-preservation-board/documents/humpback%20whales.pdf |website=Library of Congress}}

Whales and dolphins have been kept in captivity for use in education, research and entertainment since the 19th century.{{Cite web |last=Bosworth |first=Amanda |title=Barnum's Whales: The Showman and the Forging of Modern Animal Captivity |url=https://www.historians.org/perspectives-article/barnums-whales-the-showman-and-the-forging-of-modern-animal-captivity/ |access-date=2025-02-09 |website=Historians |language=en-US}}

{{Main|Beluga whale#Captivity}}

Beluga whales were the first whales to be kept in captivity. Other species were too rare, too shy or too big. The first was shown at Barnum's Museum in New York City in 1861.{{cite web | url=http://chnm.gmu.edu/lostmuseum/lm/190/ | title=The Whales, New York Tribune, August 9, 1861 | access-date=5 December 2011 | date=9 August 1861 | work=New York Tribune}} For most of the 20th century, Canada was the predominant source. They were taken from the St. Lawrence River estuary until the late 1960s, after which they were predominantly taken from the Churchill River estuary until capture was banned in 1992.{{cite web|url=http://webpages.charter.net/hrynyshyn/pdfs/Beluga_Report_web2006.pdf |title=Beluga Whales in Captivity: Hunted, Poisoned, Unprotected |access-date=26 December 2014 |year=2006 |work=Special Report on Captivity 2006 |publisher=Canadian Marine Environment Protection Society |url-status=dead |archive-url=https://web.archive.org/web/20141226215306/http://webpages.charter.net/hrynyshyn/pdfs/Beluga_Report_web2006.pdf |archive-date=26 December 2014 }} Russia then became the largest provider. Belugas are caught in the Amur Darya delta and their eastern coast and are transported domestically to aquaria or dolphinaria in Moscow, St. Petersburg and Sochi, or exported to countries such as Canada. They have not been domesticated.{{cite web | url=http://www.waza.org/en/zoo/visit-the-zoo/aquatic-mammals-1254385523/delphinapterus-leucas | title=Beluga (Delphinapterus leucas) Facts – Distribution – In the Zoo | access-date=5 December 2011 | publisher=World Association of Zoos and Aquariums | archive-url=https://web.archive.org/web/20120210101419/http://www.waza.org/en/zoo/visit-the-zoo/aquatic-mammals-1254385523/delphinapterus-leucas | archive-date=10 February 2012 | url-status=dead }}

As of 2006, 30 belugas lived in Canada and 28 in the United States. 42 deaths in captivity had been reported. A single specimen can reportedly fetch up to US$100,000 (£64,160). The beluga's popularity is due to its unique color and its facial expressions. The latter is possible because while most cetacean "smiles" are fixed, the extra movement afforded by the beluga's unfused cervical vertebrae allows a greater range of apparent expression.{{cite book |author=Bonner, Nigel |title=Whales |publisher=Facts on File |isbn=978-0-7137-0887-5 |pages=[https://archive.org/details/whales0000bonn/page/17 17, 23–24] |year=1980 |url=https://archive.org/details/whales0000bonn/page/17 }}

{{Main|Captive killer whales}}

{{See also|SeaWorld#Criticism and resulting impact|Incidents at SeaWorld parks}}

File:2009-Seaworld-Shamu.jpg

The orca's intelligence, trainability, striking appearance, playfulness in captivity and sheer size have made it a popular exhibit at aquaria and aquatic theme parks. From 1976 to 1997, fifty-five whales were taken from the wild in Iceland, nineteen from Japan and three from Argentina. These figures exclude animals that died during capture. Live captures fell dramatically in the 1990s and by 1999, about 40% of the forty-eight animals on display in the world were captive-born.{{cite web |publisher=National Marine Fisheries Service (NMFS) Northwest Regional Office |author=NMFS |year=2005 |url=http://www.nwr.noaa.gov/Marine-Mammals/Whales-Dolphins-Porpoise/Killer-Whales/Conservation-Planning/upload/SRKW-propConsPlan.pdf |title=Conservation Plan for Southern Resident Killer Whales (Orcinus orca) |location=Seattle, U.S. |access-date=January 2, 2009 |archive-url=https://web.archive.org/web/20080626121719/http://www.nwr.noaa.gov/Marine-Mammals/Whales-Dolphins-Porpoise/Killer-Whales/Conservation-Planning/upload/SRKW-propConsPlan.pdf |archive-date=June 26, 2008 |url-status=dead |pages=43–44}}

Organizations such as World Animal Protection and the Whale and Dolphin Conservation campaign against the practice of keeping them in captivity.{{citation needed|date=October 2024}}

In captivity, they often develop pathologies, such as the dorsal fin collapse seen in 60–90% of captive males. Captives have reduced life expectancy, on average only living into their 20s, although some live longer, including several over 30 years old and two, Corky II and Lolita, in their mid-40s. In the wild, females who survive infancy live 46 years on average and up to 70–80 years. Wild males who survive infancy live 31 years on average and can reach 50–60 years.{{cite web|author=Rose, N. A.|year=2011|url=http://www.hsi.org/assets/pdfs/orca_white_paper.pdf |archive-url=https://web.archive.org/web/20111026123839/http://www.hsi.org/assets/pdfs/orca_white_paper.pdf |archive-date=2011-10-26 |url-status=live|title=Killer Controversy: Why Orcas Should No Longer Be Kept in Captivity|publisher=Humane Society International and the Humane Society of the United States|access-date=December 21, 2014}}

Captivity usually bears little resemblance to wild habitat and captive whales' social groups are foreign to those found in the wild. Critics claim captive life is stressful due to these factors and the requirement to perform circus tricks that are not part of wild orca behavior. Wild orca may travel up to {{convert|160|km|mi|-1}} in a day and critics say the animals are too big and intelligent to be suitable for captivity.{{cite news|url= https://www.cbsnews.com/news/whale-attack-renews-captive-animal-debate/|title=Whale Attack Renews Captive Animal Debate|newspaper=CBS News|date=March 1, 2010|access-date=6 September 2015}} Captives occasionally act aggressively towards themselves, their tankmates, or humans, which critics say is a result of stress.{{cite book|author=Susan Jean Armstrong|title=Animal Ethics Reader| isbn=978-0-415-27589-7|year=2003|publisher=Psychology Press }} Orcas are well known for their performances in shows, but the number of orcas kept in captivity is small, especially when compared to the number of bottlenose dolphins, with only forty-four captive orcas being held in aquaria as of 2012.{{cite web|url=http://www.orcahome.de/orcastat.htm|title=Orcas in Captivity – A look at killer whales in aquariums and parks|date=23 November 2009|access-date=6 September 2015|archive-url=https://web.archive.org/web/20070602050516/http://www.orcahome.de/orcastat.htm|archive-date=2 June 2007|url-status=dead}}

Each country has its own tank requirements; in the US, the minimum enclosure size is set by the Code of Federal Regulations, 9 CFR E § 3.104, under the Specifications for the Humane Handling, Care, Treatment and Transportation of Marine Mammals.{{cite journal|url=http://www.ecfr.gov/cgi-bin/text-idx?SID=c7a201bd3f7d31d5f8218167efcd49ba&node=9:1.0.1.1.3.5.31.5&rgn=div8|title=Chapter I: Space requirements|journal=Electronic Code of Federal Regulation|volume= 1|access-date=6 September 2015}}

File:Dawn Brancheau - Riders on the Storm.jpg]]

Aggression among captive orcas is common. They attack each other and their trainers as well. In 2013, SeaWorld's treatment of orcas in captivity was the basis of the movie Blackfish, which documents the history of Tilikum, an orca at SeaWorld Orlando, who had been involved in the deaths of three people.Whiting, Candace Calloway. [https://www.huffpost.com/entry/blackfish-killer-whales_b_4166923 In the Wake of Blackfish – Is it Time to Retire the Last Killer Whale Whose Capture Was Shown in the Film?"], HuffPost, October 29, 2013. Retrieved October 29, 2013. The film led to proposals by some lawmakers to ban captivity of cetaceans, and led SeaWorld to announce in 2016 that it would phase out its orca program.{{Cite web|url=https://www.forbes.com/sites/dalebuss/2016/03/24/it-was-a-losing-battle-seaworlds-ceo-on-its-abrupt-change-and-what-comes-next/#5754041a223b|title=Shamu Goes Out With the Tide: SeaWorld CEO On Its Abrupt Change – And What Comes Next|last=Buss|first=Dale|date=2016-03-24|website=Forbes|access-date=2016-03-26}}

File:Short-finned Pilot Whale (8793172995).jpg pilot whale with trainers]]

Dolphins and porpoises are kept in captivity. Bottlenose dolphins are the most common, as they are relatively easy to train, have a long lifespan in captivity and have a friendly appearance. Bottlenose dolphins live in captivity across the world, though exact numbers are hard to determine. Other species kept in captivity are spotted dolphins, false killer whales and common dolphins, Commerson's dolphins, as well as rough-toothed dolphins, but all in much lower numbers. There are also fewer than ten pilot whales, Amazon river dolphins, Risso's dolphins, spinner dolphins, or tucuxi in captivity. Two unusual and rare hybrid dolphins, known as wolphins, are kept at Sea Life Park in Hawaii, which is a cross between a bottlenose dolphin and a false killer whale. Also, two common/bottlenose hybrids reside in captivity at Discovery Cove and SeaWorld San Diego.{{citation needed|date=October 2024}}

In repeated attempts in the 1960s and 1970s, narwhals kept in captivity died within months. A breeding pair of pygmy right whales were retained in a netted area. They were eventually released in South Africa. In 1971, SeaWorld captured a California gray whale calf in Mexico at Scammon's Lagoon. The calf, later named Gigi, was separated from her mother using a form of lasso attached to her flukes. Gigi was displayed at SeaWorld San Diego for a year. She was then released with a radio beacon affixed to her back; however, contact was lost after three weeks. Gigi was the first captive baleen whale. JJ, another gray whale calf, was kept at SeaWorld San Diego. JJ was an orphaned calf that beached itself in April 1997 and was transported two miles to SeaWorld. The {{convert|680|kg|lbs}} calf was a popular attraction and behaved normally, despite separation from his mother. A year later, the then {{convert|8,164.7|kg|lbs}} whale though smaller than average, was too big to keep in captivity, and was released on April 1, 1998. A captive Amazon river dolphin housed at Acuario de Valencia is the only trained river dolphin in captivity.{{cite book|url=https://portals.iucn.org/library/sites/library/files/documents/RD-1991-001.pdf |archive-url=https://web.archive.org/web/20150509184324/https://portals.iucn.org/library/sites/library/files/documents/RD-1991-001.pdf |archive-date=2015-05-09 |url-status=live|last1=Klinowska|first1=Margaret|last2=Cooke|first2=Justin|year=1991| title=Dolphins, Porpoises, and Whales of the World: the IUCN Red Data Book|access-date=6 September 2015}}{{cite journal| url=http://www.aquaticmammalsjournal.org/share/AquaticMammalsIssueArchives/2001/AquaticMammals_27-03/27-03_Goff.pdf |archive-url=https://web.archive.org/web/20151015214719/http://www.aquaticmammalsjournal.org/share/AquaticMammalsIssueArchives/2001/AquaticMammals_27-03/27-03_Goff.pdf |archive-date=2015-10-15 |url-status=live |title=Growth of two gray whale calves|author1=J. L. Sumich|author2=T. Goff|author3=W. L. Perryman|journal=Aquatic Mammals|pages=231–233|year=2001|access-date=6 September 2015}}

{{reflist}}

{{Wiktionary|Cetacea}}

{{Wikispecies|Cetacea}}

{{Commons category|Cetacea}}

• {{Wikibooks inline|Dichotomous Key|Cetacea}}
• {{cite EB1911|wstitle=Cetacea |volume=5 |short=x}}
• {{cite web |url=http://www.eoearth.org/article/Cetaceans?topic=49540 |publisher=Encyclopedia of Earth |title=Cetaceans}}
• [http://www.crru.org.uk/ Scottish Cetacean Research & Rescue] – see page on [http://www.crru.org.uk/taxonomy.asp Taxonomy]
• {{cite web |url=https://www.sciencedaily.com/news/plants_animals/dolphins_and_whales/ |website=Science Daily |title=Dolphin and Whale News}}
• {{cite web |first=Douglas J. |last=Futuyma |year=1998 |url=http://www.stephenjaygould.org/library/futuyma_cetacea.html |title=Cetacea Evolution |access-date=2007-03-23 |archive-date=2008-05-29 |archive-url=https://web.archive.org/web/20080529032651/http://www.stephenjaygould.org/library/futuyma_cetacea.html |url-status=dead}}
• [https://web.archive.org/web/20110227051045/http://www.eia-international.org/campaigns/species/cetaceans/ EIA Cetacean campaign]: Reports and latest info.
• [https://web.archive.org/web/20101121062046/http://www.eia-global.org/species_in_peril/ EIA in USA]: reports etc.

{{Mammals}}

{{Cetacea}}

{{Portal bar|Cetaceans|Mammals|Marine life|Oceans}}

{{Taxonbar|from1=Q160|from2=Q37496661|from3=Q21075082}}

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Category:Extant Ypresian first appearances

Category:Mammal infraorders

Category:Taxa named by Mathurin Jacques Brisson

Category:Taxa described in 1762