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The Morrison Man/sandbox3
Temporal range: 53.5–0 Ma Early Eocene – Present
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Sperm whale
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Humpback whale
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Orca
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North Atlantic right whale
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Amazon river dolphin
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Harbor porpoise
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Clade: Cetaceamorpha
Infraorder: Cetacea
Brisson, 1762
Subgroups

(see text for families)

Diversity
Around 94 species

Cetacea (/sɪˈteɪʃə/; from Latin cetus 'whale', from Ancient Greek κῆτος (kêtos) 'huge fish, sea monster')[3] is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel themselves through the water with powerful up-and-down movement of their tail which ends in a paddle-like fluke, using their flipper-shaped forelimbs to maneuver.[4]

While the majority of cetaceans live in marine environments, a small number reside solely in brackish water or fresh water. Having a cosmopolitan distribution, they can be found in some rivers and all of Earth's oceans, and many species inhabit vast ranges where they migrate with the changing of the seasons.

Cetaceans are famous for their high intelligence, complex social behaviour, and the enormous size of some of the group's members. For example, the blue whale reaches a maximum confirmed length of 29.9 meters (98 feet) and a weight of 173 tonnes (190 short tons), making it the largest animal ever known to have existed.[5][6][7]

There are approximately 89[8] living species split into two parvorders: Odontoceti or toothed whales (containing porpoises, dolphins, other predatory whales like the beluga and the sperm whale, and the poorly understood beaked whales) and the filter feeding Mysticeti or baleen whales (which includes species like the blue whale, the humpback whale and the bowhead whale). Despite their highly modified bodies and carnivorous lifestyle, genetic and fossil evidence places cetaceans as nested within even-toed ungulates, most closely related to hippopotamus within the clade Whippomorpha.

Cetaceans have been extensively hunted for their meat, blubber and oil by commercial operations. Although the International Whaling Commission has agreed on putting a halt to commercial whaling, whale hunting is still going on, either under IWC quotas to assist the subsistence of Arctic native people or in the name of scientific research, although a large spectrum of non-lethal methods are now available to study marine mammals in the wild.[9] Cetaceans also face severe environmental hazards from underwater noise pollution, entanglement in abandoned ropes and nets, collisions with ships, plastic and heavy metals build-up, to accelerating climate change,[10][11] but how much they are affected varies widely from species to species, from minimally in the case of the southern bottlenose whale to the baiji (Chinese river dolphin) which is considered to be functionally extinct due to human activity.[12]

Anatomy

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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).[13]

The male genitals and the mammary glands of females are sunken into the body.[14][15] The male genitals are attached to a vestigial pelvis.[16]

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.[17] 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.[17][18] Hind legs are not present in cetaceans, nor are any other external body attachments such as a pinna and hair.[19]

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Whales have an elongated head, especially baleen whales, due to the wide overhanging jaw. Bowhead whale plates can be 9 metres (30 ft) long. Their nostril(s) make up the blowhole, with one in toothed whales and two in baleen whales.[20]

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.[21]

Brain

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Sperm whales have the largest brain mass of any animal on Earth, averaging 8,000 cm3 (490 in3) and 7.8 kg (17 lb) in mature males.[22] The brain to body mass ratio in some odontocetes, such as belugas and narwhals, is second only to humans.[23]

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.[24]

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. mbrain ∝ (mbody)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.[25] 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.[26]

The neocortex of many cetaceans is home to elongated spindle neurons that, prior to 2019, were known only in hominids.[27] In humans, these cells are thought to be involved in social conduct, emotions, judgment and theory of mind.[28] Cetacean spindle neurons are found in areas of the brain homologous to where they are found in humans, suggesting they perform a similar function.[29]

Fluke

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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.[30]

Physiology

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Abdominal organs

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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.[31]

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.[32] The urinary bladder is proportionally smaller in cetaceans than in land mammals.[33] The testes are located internally, without an external scrotum.[34][35][36][37] The uterus is bicornuate.[35]

Ears

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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.[38]

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 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.[39]

Cetaceans use sound to communicate, using groans, moans, whistles, clicks or the 'singing' of the humpback whale.[40]

Senses

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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. 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.[41] Cetaceans are known to possess excellent hearing.[42]

At least one species, the tucuxi or Guiana dolphin, is able to use electroreception to sense prey.[40]

Echolocation

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Odontoceti are generally capable of echolocation.[43] 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.[44]

Mysticeti have exceptionally thin, wide basilar membranes in their cochleae without stiffening agents, making their ears adapted for processing low to infrasonic frequencies.[45]

Chromosomes

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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.[46]

Ecology

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Range and habitat

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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.[47]

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.[48]

Migration

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Many species of whales migrate on a latitudinal basis to move between seasonal habitats. For example, the gray whale migrates 10,000 miles (16,000 km) round trip. The journey begins at winter birthing grounds in warm lagoons along Baja California, and traverses 5,000–7,000 miles (8,000–11,300 km) of coastline to summer feeding grounds in the Bering, Chuckchi and Beaufort seas off the coast of Alaska.[49]

Behaviour

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Sleep

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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.[50]

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.[51]

Diving

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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 100 m (330 ft). Species such as sperm whales can dive to 3,000 m (9,800 ft), although more commonly 1,200 metres (3,900 ft).[52][53]

Social relations

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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.[54]

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.[54]

Whale song

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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.[54] In baleen species such as humpbacks, blues and fins, male-specific song is believed to be used to attract and display fitness to females.[55]

Decision-making

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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). [56]

Hunting

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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.[54]

Intelligence

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Main article: Cetacean intelligence

Cetacea are known to teach, learn, cooperate, scheme and grieve.[57]

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.[58]

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.[59]

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.[60] 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.[61]

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.[61]

In 1995, Marten and Psarakos used video to test dolphin self-awareness.[62] 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.[61]

Life history

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Reproduction and development

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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.[54] 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.[54]

Lifespan

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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.[63]

Death

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Main article: 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.[64]

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.[64][65]

Disease

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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.[66]

Evolution

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Main article: Evolution of cetaceans

Fossil history

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Transition from land to sea

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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.[67]

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 3 m (9.8 ft) 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.[68] 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.[69]

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.[67]

Marine animals

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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.[70]

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.[67]

The two parvorders, baleen whales (Mysticeti) and toothed whales (Odontoceti), are thought to have diverged around thirty-four million years ago.[71]

Phylogeny

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Further information: List of cetaceans and List of extinct cetaceans

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.[67] 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.[67] 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.[72][73][74]

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:[75][76][77][78][79]

Artiodactyla 

Tylopoda (camels) 

 Artiofabula 

 Suina (pigs) 

 Cetruminantia 
 Ruminantia (ruminants) 

 Tragulidae (mouse deer) 

 Pecora (horn bearers) 

 Cetancodonta/Whippomorpha 

 Hippopotamidae (hippopotamuses) 

 Cetacea (whales) 

54 mya
Relationship of extinct and extant cetaceans[80]:
 

 Raoellidae †

 Cetacea 
† Extinct taxa

The terms whale and dolphin are informal:

The term 'great whales' covers those currently regulated by the International Whaling Commission:[81] 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).[82]

Threats

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The primary threats to cetaceans come from people, both directly from whaling or drive hunting and indirect threats from fishing and pollution.[83]

Whaling

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Main articles: Whaling, History of Whaling, and Dolphin drive hunting

The first efforts to protect whales came in 1931. Some particularly endangered species, such as the humpback whale (which then numbered about 100 animals).[84]

Aboriginal whaling is still permitted. About 1,200 pilot whales were taken in the Faroe Islands in 2017.[85]

Bycatch

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See also: Cetacean bycatch

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.[83]

Strandings

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Main article: Cetacean stranding

The causes are not clear. Possible reasons for mass beachings are:[83]

  • 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.[86]

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.[87]

Environmental hazards

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See also: Marine mammals and sonar

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.

—Rear Admiral Lawrence Rice

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.[83]

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.[83]

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.[83]

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.[88] 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.[89]

Relationship with humans

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See also: Cryptid whale and Whale § In myth, literature and art

Antiquity

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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.[90] 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.[91] 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.[92][93]

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 Bible especially, the leviathan plays a role as a sea monster. The essence, which features a giant crocodile or a dragon and a whale, was created according to the Bible by God[94] and should again be destroyed by him.[95][96] In the Book of Job, the leviathan is described in more detail.[97][98]

Middle Ages to the 19th century

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See also: History of Whaling and Cryptid whales

In the Canadian Arctic (east coast) in Punuk and Thule culture (1000–1600 C.E.),[99] baleen was used to construct houses in place of wood as roof support for winter houses, with half of the building buried under the ground. The actual roof was probably made of animal skins that were covered with soil and moss.[100]

Modern culture

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Further information: Whale § Interactions with humans

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.[101]

Captivity

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Belugas

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Main article: 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.[102] For most of the 20th century, Canada was the predominant source.[103] 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.[103] Russia then became the largest provider.[103] 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.[103] They have not been domesticated.[104]

As of 2006, 30 belugas lived in Canada and 28 in the United States. 42 deaths in captivity had been reported.[103] 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.[105]

Orcas

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Main article: Captive killer whales
See also: SeaWorld § Criticism and resulting impact, and Incidents at SeaWorld parks

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.[106]

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.[107]

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 160 kilometres (100 mi) in a day and critics say the animals are too big and intelligent to be suitable for captivity.[108] Captives occasionally act aggressively towards themselves, their tankmates, or humans, which critics say is a result of stress.[109] 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.[110]

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.[111]

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.[112] 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 after various unsuccessful attempts to restore its revenues, reputation, and stock price.[113]

Others

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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 680 kilograms (1,500 lb) calf was a popular attraction and behaved normally, despite separation from his mother. A year later, the then 8,164.7 kilograms (18,000 lb) 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.[114][115]

Images

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References

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[[Category:Cetaceans| [[Category:Extant Ypresian first appearances [[Category:Mammal infraorders [[Category:Taxa named by Mathurin Jacques Brisson [[Category:Taxa described in 1762

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  NODES
Association 3
Idea 6
idea 6
INTERN 10
Note 1
todo 1