Whale

This article is about a marine mammal. For other uses, see Whale (disambiguation).

For further information, see Cetacea.

Whales
Whales are not a taxon, they are an informal grouping of the infraorder Cetacea
Southern right whale
Information
Families considered whales	Parvorder Mysticeti Family Balaenidae Family Balaenopteridae Family Eschrichtiidae Family Cetotheriidae Parvorder Odontoceti (excluding dolphins and porpoises) Family Monodontidae Family Physeteridae Family Kogiidae Family Ziphiidae
Range map

Whale is the common name for a widely distributed and diverse group of fully aquatic marine mammals. They are an informal grouping within the infraorder Cetacea, excluding dolphins and porpoises, so to zoologists the grouping is paraphyletic. The whales comprise the extant families Cetotheriidae (whose only living member is the pygmy right whale), Balaenopteridae (the rorquals), Balaenidae (right whales), Eschrichtiidae (the gray whale), Monodontidae (belugas and narwhals), Physeteridae (the sperm whale), Kogiidae (the dwarf and pygmy sperm whale), and Ziphiidae (the beaked whales). There are 40 extant species of whales. The two parvorders of whales, Mysticeti and Odontoceti, are thought to have split apart around 34 million years ago. Whales, dolphins and porpoises belong to the order Cetartiodactyla with even-toed ungulates and their closest living relatives are the hippopotamuses, having diverged about 40 million years ago.

Whales range in size from the 2.6 metres (8.5 ft) and 135 kilograms (298 lb) dwarf sperm whale to the 34 metres (112 ft) and 190 metric tons (210 short tons) blue whale, which is the largest creature on earth. Several species exhibit sexual dimorphism, in that the females are larger than males. They have streamlined bodies and two limbs that are modified into flippers. Though not as flexible or agile as seals, whales can travel at up to 20 knots. Balaenopterids use their throat pleats to expand the mouth to take in gulps of water. Balaenids have heads that can make up 40% of their body mass to take in water. Odontocetes have conical teeth designed for catching fish or squid. Mysticetes have a well developed sense of "smell", whereas odontocetes have well-developed hearing − their hearing, that is adapted for both air and water, is so well developed that some can survive even if they are blind. Some species are well adapted for diving to great depths. They have a layer of fat, or blubber, under the skin to keep warm in the cold water.

Although whales are widespread, most species prefer the colder waters of the Northern and Southern Hemispheres, and migrate to the equator to give birth. Odontocetes feed largely on fish and squid. A few, like the sperm whale, feed on large invertebrates, such as giant squid. Grey whales are specialized for feeding on bottom-dwelling molluscs. Male whales typically mate with multiple females every year, but females only mate every two to three years. Calves are typically born in the spring and summer months and females bear all the responsibility for raising them. Mothers of some species fast and nurse their young for a relatively long period of time. Whales produce a variety of vocalizations, notably the songs of the humpback whale.

Once relentlessly hunted for their products, whales are now protected by international law. The North Atlantic right whales nearly became extinct in the twentieth century, with a population low of 450, and the North Pacific gray whale population is ranked Critically Endangered by the IUCN. Besides whaling, they also face threats from bycatch and marine pollution. The meat, blubber and baleen of whales have traditionally been used by indigenous peoples of the Arctic. Whales have been depicted in various cultures worldwide, notably by the Inuit and the coastal peoples of Vietnam and Ghana, who sometimes hold whale funerals. Whales occasionally feature in literature and film, as in the great white whale of Herman Melville's Moby Dick. Small whales, such as belugas, are sometimes kept in captivity and trained to perform tricks, but breeding success has been poor and the animals often die within a few months of capture. Whale watching has become a form of tourism around the world.

Taxonomy and evolution

See also: List of whale species and List of extinct cetaceans

The hippopotamuses are the sister clade to the Cetacea, which contains whales, dolphins and porpoises.

The whales are part of the largely terrestrial mammalian clade Laurasiatheria. Whales do not form a clade or order; the infraorder Cetacea includes dolphins and porpoises, which are not considered whales.

Whales are paraphyletic, consisting as they do of all cetaceans except dolphins and porpoises.

Laurasiatheria

Ferae (carnivores and allies) Perissodactyla (horses, rhinos, tapirs) Artiodactyla Tylopoda (camelids) A Pigs C Ruminants (cattle, sheep, antelopes) W Hippopotamuses Cetacea Archaeocetes (Ambulocetus, Protocetus, Basilosaurus) Mysticeti (baleen whales) (right, gray, rorquals) Odontoceti (toothed whales) Delphinoidea (dolphins, porpoises, belugas) Lipotoidea (river dolphins) Physeteroidea (sperm whales) Ziphoidea (beaked whales)

• Whales in green
• A=Artiofabula
• C=Cetruminantia
• W=Whippomorpha

Cetaceans are divided into two parvorders:

• The largest parvorder, Mysticeti (baleen whales), is characterized by the presence of baleen, a sieve-like structure in the upper jaw made of keratin, which it uses to filter plankton, among others, from the water.
• Odontocetes (toothed whales) are characterized by bearing sharp teeth for hunting, as opposed to their counterparts' baleen.

Cetaceans and artiodactyls now are classified under the order Cetartiodactyla, often still referred to as Artiodactyla, which includes both whales and hippopotamuses. The hippopotamus and pygmy hippopotamus are the whale's closest terrestrial living relatives.

Mysticetes

See also: Mysticeti

Mysticetes are also known as baleen whales. They have a pair of blowholes side-by-side and lack teeth, which renders them incapable of catching larger prey; they instead have baleen plates which is a sieve-like structure in the upper jaw made of keratin, which it uses to filter plankton and other food from the water; this forces them to follow krill or plankton migrations. Some whales, such as the humpback, reside in the polar regions where they feed on a reliable source of schooling fish and krill. These animals rely on their well-developed flippers and tail fin to propel themselves through the water; they swim by moving their fore-flippers and tail fin up and down. Whale ribs loosely articulate with their thoracic vertebrae at the proximal end, but do not form a rigid rib cage. This adaptation allows their chest to compress during deep dives as the pressure increases with depth. Mysticetes consist of four families: rorquals (balaenopterids), cetotheriids, right whales (balaenids), and gray whales (eschrichtiids).

The main difference between each family of mysticete is in their feeding adaptations and subsequent behaviour. Balaenopterids are the rorquals. These animals, along with the cetotheriids, rely on their throat pleats to gulp large amounts of water while feeding. The throat pleats extend from the mouth to the navel and allow the mouth to expand to a large volume for more efficient capture of the small animals they feed on. Balaenopterids consist of two genera and eight species. Balaenids are the right whales. These animals have very large heads, which can make up as much as 40% of their body mass, and much of the head is the mouth. This allows them to take in large amounts of water into their mouths, letting them feed more effectively. Eschrichtiids have one living member: the gray whale. They are bottom feeders, mainly eating crustaceans and benthic invertebrates. They feed by turning on their sides and taking in water mixed with sediment, which is then expelled through the baleen, leaving their prey trapped inside. This is an efficient method of hunting, in which the whale has no major competitors.

Odontocetes

See also: Odontoceti

Odontocetes are known as toothed whales; they have teeth and only one blowhole. They rely on their well-developed sonar to find their way in the water. Toothed whales send out ultrasonic clicks using the melon. Sound waves travel through the water. Upon striking an object in the water, the sound waves bounce back at the whale. These vibrations are received through fatty tissues in the jaw, which is then rerouted into the ear-bone and into the brain where the vibrations are interpreted. All toothed whales are opportunistic, meaning they will eat anything they can fit in their throat because they are unable to chew. These animals rely on their well-developed flippers and tail fin to propel themselves through the water; they swim by moving their fore-flippers and tail fin up and down. Whale ribs loosely articulate with their thoracic vertebrae at the proximal end, but they do not form a rigid rib cage. This adaptation allows the chest to compress during deep dives as opposed to resisting the force of water pressure. Excluding dolphins and porpoises, odontocetes consist of four families: belugas and narwhals (monodontids), sperm whales (physeterids), dwarf and pygmy sperm whales (kogiids), and beaked whales (ziphiids). There are six species, sometimes referred to as "blackfish", that are dolphins commonly misconceived as whales: the killer whale, the melon-headed whale, the pygmy killer whale, the false killer whale, and the two species of pilot whales, all of which are classified under the family Delphinidae (oceanic dolphins).

The differences between families of odontocetes include size, feeding adaptations and distribution. Monodontids consist of two species: the beluga and the narwhal. They both reside in the frigid arctic and both have large amounts of blubber. Belugas, being white, hunt in large pods near the surface and around pack ice, their coloration acting as camouflage. Narwhals, being black, hunt in large pods in the aphotic zone, but their underbelly still remains white to remain camouflaged when something is looking directly up or down at them. They have no dorsal fin to prevent collision with pack ice. Physeterids and Kogiids consist of sperm whales. Sperm whales consist the largest and smallest odontocetes, and spend a large portion of their life hunting squid. P. macrocephalus spends most of its life in search of squid in the depths; these animals do not require any degree of light at all, in fact, blind sperm whales have been caught in perfect health. The behaviour of Kogiids remains largely unknown, but, due to their small lungs, they are thought to hunt in the photic zone. Ziphiids consist of 22 species of beaked whale. These vary from size, to coloration, to distribution, but they all share a similar hunting style. They use a suction technique, aided by a pair of grooves on the underside of their head, not unlike the throat pleats on the rorquals, to feed.

Evolution

Basilosaurus skeleton

Main article: Evolution of cetaceans

Odontocetes and mysticetes split into two separate parvorders around 34 million years ago.

Whales are descendants of land-dwelling mammals of the artiodactyl order (even-toed ungulates). They are related to the Indohyus, an extinct chevrotain-like ungulate, from which they split approximately 48 million years ago. Primitive cetaceans, or archaeocetes, first took to the sea approximately 49 million years ago and became fully aquatic 5–10 million years later. Archaeoceti is a parvorder comprising ancient whales. These ancient whales are the predecessors of modern whales, stretching back to their first ancestor that spent their lives near (rarely in) the water. Likewise, the archaeocetes can be anywhere from near fully terrestrial, to semi-aquatic to fully aquatic, but what defines an archaeocete is the presence of anatomical features exclusive to cetaceans alongside other primitive features not found in modern cetaceans, like visible legs or asymmetrical teeth. Their features became adapted for living in the marine environment. Major anatomical changes include their hearing set-up that channels vibrations from the jaw to the earbone which occurred with Ambulocetus 49 million years ago, a streamlined body and the growth of flukes on the tail which occurred around 43 million years ago with Protocetus, the migration of the nasal openings toward the top of the cranium and the modification of the forelimbs into flippers which occurred with Basilosaurus 35 million years ago, and the shrinking and eventual disappearance of the hind limbs which took place with the first odontocetes and mysticetes 34 million years ago.

Today, the closest living relatives of cetaceans are the hippopotamuses; these share a semi-aquatic ancestor that branched off from other artiodactyls some 60 million years ago. Around 40 million years ago, a common ancestor between the two branched off into cetacea and anthracotheres; nearly all anthracotheres went extinct at the end of the Pleistocene two-and-a-half million years ago, eventually leaving only one surviving lineage: the hippo.

Biology

Anatomy

Features of a blue whale

Features of a sperm whale skeleton

Whales have torpedo shaped bodies with non-flexible necks, limbs modified into flippers, non-existent external ear flaps, a large tail fin, and flat heads (with the exception of monodontids and ziphiids). Whale skulls have small eye orbits, long snouts (with the exception of monodontids and ziphiids) and eyes placed on the sides of its head. Whales range in size from the 2.6 metres (8.5 ft) and 135 kilograms (298 lb) dwarf sperm whale to the 34 metres (112 ft) and 190 metric tons (210 short tons) blue whale. Overall, they tend to dwarf other cetartiodactyls; the blue whale is the largest creature on earth. Several species have female-biased sexual dimorphism, with the females being larger than the males. One exception is with the sperm whale, which has males larger than the females.

Odontocetes, such as the sperm whale, possess teeth with cementum cells overlying dentine cells. Unlike human teeth, which are composed mostly of enamel on the portion of the tooth outside of the gum, whale teeth have cementum outside the gum. Only in larger whales, where the cementum is worn away on the tip of the tooth, does enamel show. Mysticetes have large whalebone, as opposed to teeth, made of keratin. Mysticetes have two blowholes, whereas Odontocetes contain only one.

Breathing involves expelling stale air from the blowhole, forming an upward, steamy spout, followed by inhaling fresh air into the lungs; a humpback whale's lungs can hold about 5,000 litres of air. Spout shapes differ among species, which facilitates identification.

All whales have a thick layer of blubber. In species that live near the poles, the blubber can be as thick as 11 inches. This blubber can help with buoyancy (which is helpful for a 100-ton whale), protection to some extent as predators would have a hard time getting through a thick layer of fat, and energy for fasting when migrating to the equator; the primary usage for blubber is insulation from the harsh climate. It can constitute as much as 50% of a whales body weight. Calves are born with only a thin layer of blubber, but some species compensate for this with thick lanugos.

Whales have a two- to three-chambered stomach that is similar in structure to terrestrial carnivores. Mysticetes contain a proventriculus as an extension of the oesophagus; this contains stones that grind up food. They also have fundic and pyloric chambers.

Locomotion

Skeleton of a bowhead whale; notice the hind limb. Richard Lydekker, 1894

Whales have two flippers on the front, and a tail fin. These flippers contain four digits. Although whales do not possess fully developed hind limbs, some, such as the sperm whale and bowhead whale, possess discrete rudimentary appendages, which may contain feet and digits. Whales are fast swimmers in comparison to seals, which typically cruise at 5–15 kn, or 9–28 kilometres per hour (5.6–17.4 mph); the fin whale, in comparison, can travel at speeds up to 47 kilometres per hour (29 mph) and the sperm whale can reach speeds of 35 kilometres per hour (22 mph). The fusing of the neck vertebrae, while increasing stability when swimming at high speeds, decreases flexibility; whales are unable to turn their heads. When swimming, whales rely on their tail fin propel them through the water. Flipper movement is continuous. Whales swim by moving their tail fin and lower body up and down, propelling themselves through vertical movement, while their flippers are mainly used for steering. Some species log out of the water, which may allow then to travel faster. Their skeletal anatomy allows them to be fast swimmers. Most species have a dorsal fin.

Whales are adapted for diving to great depths. In addition to their streamlined bodies, they can slow their heart rate to conserve oxygen; blood is rerouted from tissue tolerant of water pressure to the heart and brain among other organs; haemoglobin and myoglobin store oxygen in body tissue; and they have twice the concentration of myoglobin than haemoglobin. Before going on long dives, many whales exhibit a behaviour known as sounding; they stay close to the surface for a series of short, shallow dives while building their oxygen reserves, and then make a sounding dive.

Senses

Biosonar by cetaceans

Sperm whale skeleton. Richard Lydekker, 1894.

The whale ear has specific adaptations to the marine environment. In humans, the middle ear works as an impedance equalizer between the outside air's low impedance and the cochlear fluid's high impedance. In whales, and other marine mammals, there is no great difference between the outer and inner environments. Instead of sound passing through the outer ear to the middle ear, whales receive sound through the throat, from which it passes through a low-impedance fat-filled cavity to the inner ear. The whale ear is acoustically isolated from the skull by air-filled sinus pockets, which allow for greater directional hearing underwater. Odontocetes send out high frequency clicks from an organ known as a melon. This melon consists of fat, and the skull of any such creature containing a melon will have a large depression. The melon size varies between species, the bigger the more dependent they are of it. A beaked whale for example has a small bulge sitting on top of its skull, whereas a sperm whale's head is filled up mainly with the melon.

The whale eye is relatively small for its size, yet they do retain a good degree of eyesight. As well as this, the eyes of a whale are placed on the sides of its head, so their vision consists of two fields, rather than a binocular view like humans have. When belugas surface, their lens and cornea correct the nearsightedness that results from the refraction of light; they contain both rod and cone cells, meaning they can see in both dim and bright light, but they have far more rod cells than they do cone cells. Whales do, however, lack short wavelength sensitive visual pigments in their cone cells indicating a more limited capacity for colour vision than most mammals. Most whales have slightly flattened eyeballs, enlarged pupils (which shrink as they surface to prevent damage), slightly flattened corneas and a tapetum lucidum; these adaptations allow for large amounts of light to pass through the eye and, therefore, a very clear image of the surrounding area. In water, a whale can see around 10.7 metres (35 ft) ahead of itself, but, of course, they have a smaller range above water. They also have glands on the eyelids and outer corneal layer that act as protection for the cornea.

The olfactory lobes are absent in toothed whales, suggesting that they have no sense of smell. Some whales, such as the bowhead whale, possess a vomeronasal organ, which does mean that they can "sniff out" krill.

Whales are not thought to have a good sense of taste, as their taste buds are atrophied or missing altogether. However, some toothed whales have preferences between different kinds of fish, indicating some sort of attachment to taste. The presence of the Jacobson's organ indicates that whales can smell food once inside their mouth, which might be similar to the sensation of taste.

Communication

Main article: Whale vocalization

Whale vocalization is likely to serve several purposes. Some species, such as the humpback whale, communicate using melodic sounds, known as whale song. These sounds may be extremely loud, depending on the species. Humpback whales only have been heard making clicks, while toothed whales use sonar that may generate up to 20,000 watts of sound (+73 dBm or +43 dBw) and be heard for many miles.

Captive whales have occasionally been known to mimic human speech. Scientists have suggested this indicates a strong desire on behalf of the whales to communicate with humans, as whales have a very different vocal mechanism, so imitate human speech likely takes considerable effort.

Whales emit two distinct kinds of acoustic signals, which are called whistles and clicks:

Clicks are quick broadband burst pulses, used for sonar, although some lower-frequency broadband vocalizations may serve a non-echolocative purpose such as communication; for example, the pulsed calls of belugas. Pulses in a click train are emitted at intervals of ~35–50 milliseconds, and in general these inter-click intervals are slightly greater than the round-trip time of sound to the target.

Whistles are narrow-band frequency modulated (FM) signals, used for communicative purposes, such as contact calls.

Intelligence

Main article: Cetacean intelligence

See also: Cetacean surfacing behaviour

Whales are known to teach, learn, cooperate, scheme, and grieve. The neocortex of many species of whale is home to elongated spindle neurons that, prior to 2007, were known only in hominids. In humans, these cells are involved in social conduct, emotions, judgement, and theory of mind. Whale spindle neurons are found in areas of the brain that are homologous to where they are found in humans, suggesting that they perform a similar function.

Bubble net feeding

Brain size was previously considered a major indicator of the intelligence of an animal. 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 more complex cognitive tasks. Allometric analysis indicates that mammalian brain size scales at approximately the ⅔ or ¾ exponent of the body mass. Comparison of a particular animal's brain size with the expected brain size based on such allometric analysis provides an encephalisation quotient that can be used as another indication of animal intelligence. Sperm whales have the largest brain mass of any animal on earth, averaging 8,000 cubic centimetres (490 in³) and 7.8 kilograms (17 lb) in mature males, in comparison to the average human brain which averages 1,450 cubic centimetres (88 in³) in mature males. The brain to body mass ratio in some odontocetes, such as belugas and narwhals, is second only to humans. In some whales, however, it is less than half that of humans: 0.9% versus 2.1%. This comparison seems more favourable if the large amount of blubber that some whales require for insulation is omitted.

Small whales are known to engage in complex play behaviour, which includes such things as producing stable underwater toroidal air-core vortex rings or "bubble rings". There are two main methods of bubble ring production: rapid puffing of a burst 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 are also known to produce bubble-nets for the purpose of foraging.

A southern right whale sailing

Larger whales are also thought, to some degree, to engage in play. The southern right whale, for example, elevates their tail fluke above the water, remaining in the same position for a considerable amount of 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. Humpback whales, among others, are also known to display this behaviour.

Self-awareness is seen, by some, to be a sign of highly developed, abstract thinking. Self-awareness, though not well-defined scientifically, is believed to be the precursor to more advanced processes like meta-cognitive reasoning (thinking about thinking) that are typical of humans. Research in this field has suggested that cetaceans, among others, possess self-awareness. 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; they then see if the animal shows signs of self-recognition.

Some disagree with these findings, arguing that the results of these tests are open to human interpretation and susceptible to the Clever Hans effect. This test is much less definitive than when used for primates, because primates can touch the mark or the mirror, while cetaceans cannot, making their alleged self-recognition behaviour less certain. Sceptics argue that behaviours that are said to identify self-awareness resemble existing social behaviours, and so researchers could be misinterpreting self-awareness for social responses to another individual. The researchers counter-argue that the behaviours shown are evidence of self-awareness, as they are very different from normal responses to another individual. Whereas apes can merely touch the mark on themselves with their fingers, cetaceans show less definitive behaviour of self-awareness; they can only twist and turn themselves to observe the mark.

Life cycle

See also: Right whale § Courtship and reproduction and Fin whale § Breeding

Whales are fully aquatic creatures, which means that birth and courtship behaviours are very different from terrestrial and semi-aquatic creatures. Since they are unable to go onto land to calve, they deliver the baby with the fetus positioned for tail-first delivery. This prevents the baby from drowning either upon or during delivery. To feed the new-born, whales, being aquatic, must squirt the milk into the mouth of the calf. Being mammals, they, of course, have mammary glands used for nursing calves; they are weaned off at about 11 months of age. This milk contains high amounts of fat which is meant to hasten the development of blubber; it contains so much fat that it has the consistency of toothpaste. Females deliver a single calf with gestation lasting about a year, dependency until one to two years, and maturity around seven to ten years, all varying between the species. This mode of reproduction produces few offspring, but increases the survival probability of each one. Females, referred to as "cows", carry the responsibility of childcare as males, referred to as "bulls", play no part in raising calves.

Most mysticetes reside at the poles. So, to prevent the unborn calf from dying of frostbite, they migrate to calving/mating grounds. They will then stay there for a matter of months until the calf has developed enough blubber to survive the bitter temperatures of the poles. Until then, the calves will feed on the mother's fatty milk. Migration times are basically uniform between the species of the suborder, but the males of some species will not undertake the migration; one example is the North Atlantic right whale. Most will travel from the Arctic or Antarctic into the tropics to mate, calve, and raise during the winter and spring; they will migrate back to the poles in the warmer summer months so the calf can continue growing while the mother can continue eating, as they fast in the breeding grounds. One exception to this is the southern right whale, which migrates to Patagonia and western New Zealand to calve; both are well out of the tropic zone.

Sleep

Unlike most animals, whales are conscious breathers. All mammals sleep, but whales cannot afford to become unconscious for long because they may drown. While knowledge of sleep in wild cetaceans is limited, toothed cetaceans in captivity have been recorded to 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.

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.

Ecology

Foraging and predation

See also: Beluga whale § Predation

Polar bear with the remains of a beluga

All whales are carnivorous and predatory. Odontocetes, as a whole, mostly feed on fish and cephalopods, and then followed by crustaceans and bivalves. All species are generalist and opportunistic feeders. Mysticetes, as a whole, mostly feed on krill and plankton, followed by crustaceans and other invertebrates. A few are specialists. Examples include the blue whale, which eats almost exclusively krill, the minke whale, which eats mainly schooling fish, the sperm whale, which specialize on squid, and the gray whale which feed on bottom-dwelling invertebrates. The elaborate baleen "teeth" of filter-feeding species, mysticetes, allow them to remove water before they swallow their planktonic food by using the teeth as a sieve. Usually whales hunt solitarily, but they do sometimes hunt cooperatively in small groups. The former behaviour is typical when hunting non-schooling fish, slow-moving or immobile invertebrates or endothermic prey. When large amounts of prey are available, whales such as certain mysticetes hunt cooperatively in small groups. Some cetaceans may forage with other kinds of animals, such as other species of whales or certain species of pinnipeds.

Large whales, such as mysticetes, are not usually subject to predation, but smaller whales, such as monodontids or ziphiids, are. These species are preyed on by the killer whale or orca. To subdue and kill whales, orcas continuously ram them with their heads; this can sometimes kill bowhead whales, or severely injure them. Other times they corral the narwhals or belugas before striking. They are typically hunted by groups of 10 or fewer orcas, but they are seldom attacked by an individual. Calves are more commonly taken by orcas, but adults can be targeted as well.

These small whales are also targeted by terrestrial predators. The polar bear is well adapted for hunting Arctic whales and calves. Bears are known to use sit-and-wait tactics as well as active stalking and pursuit of prey on ice or water. Whales lessen the chance of predation by gathering in groups. This however means less room around the breathing hole as the ice slowly closes the gap. When out at sea, whales dive out of the reach of surface-hunting orcas. Polar bear attacks on belugas and narwhals are usually successful in winter, but rarely inflict any damage in summer.

Whale pump

"Whale pump" – the role played by whales in recycling ocean nutrients

A 2010 study considered whales to be a positive influence to the productivity of ocean fisheries, in what has been termed a "whale pump." Whales carry nutrients such as nitrogen from the depths back to the surface. This functions as an upward biological pump, reversing an earlier presumption that whales accelerate the loss of nutrients to the bottom. This nitrogen input in the Gulf of Maine is "more than the input of all rivers combined" emptying into the gulf, some 23,000 metric tons (25,000 short tons) each year. Whales defecate at the ocean's surface; their excrement is important for fisheries because it is rich in iron and nitrogen. The whale faeces are liquid and instead of sinking, they stay at the surface where phytoplankton feed off it.

Whale fall

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.

Deterioration of whale carcasses happens though a series of three stages. Initially, moving 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 sulphide enabling the growth of chemoautotrophic organisms, which in turn, support other organisms such as mussels, clams, limpets, and sea snails. This stage may last for decades and supports a rich assemblage of species, averaging 185 species per site.

Interaction with humans

Whaling

Main articles: Whaling and History of whaling

Whale Fishing: Woodcut by Thevet, Paris, 1574

Dutch whalers near Spitsbergen, their most successful port. Abraham Storck, 1690

World population graph of blue whales

Whaling by humans has existed since the Stone Age. Ancient whalers used harpoons to spear the bigger animals from boats out at sea. People from Norway started hunting whales around 2000 B.C., and people from Japan began hunting whales in the Pacific at least as early. Whales are typically hunted for their meat and blubber by aboriginal groups; they used baleen for baskets or roofing, and made tools and masks out of bones. The Inuit hunted whales in the Arctic Ocean. The Basques started whaling as early as the 11th century, sailing as far as Newfoundland in the 16th century in search of right whales. 18th and 19th century whalers hunted down whales mainly for their oil, which was used as lamp fuel and a lubricant, baleen or whalebone, which was used for items such as corsets and skirt hoops, and ambergris, which was used as a fixative for perfumes. The most successful whaling nations at this time were Holland, Japan, and the United States.

Commercial whaling was historically important as an industry well throughout the 17th, 18th and 19th centuries. Whaling was at that time a sizeable European industry with ships from Britain, France, Spain, Denmark, the Netherlands and Germany, sometimes collaborating to hunt whales in the Arctic, sometimes in competition leading even to war. By the early 1790s, whalers, namely the Americans and Australians, mainly focused efforts in the South Pacific where they mainly hunted sperm whales and right whales, with catches of up to 39,000 right whales by Americans alone. By 1853, U.S. profits turned to US$11,000,000 (UK£6.5m), equivalent to US$348,000,000 (UK£230m) today, the most profitable year for the American whaling industry. Commonly exploited species included North Atlantic right whales, sperm whales, which were mainly hunted by Americans, bowhead whales, which were mainly hunted by the Dutch, common minke whales, blue whales, and gray whales. The scale of whale harvesting decreased substantially after 1982 when the International Whaling Commission (IWC) placed a moratorium which set a catch limit for each country, excluding aboriginal groups until 2004.

Current whaling nations are Norway, Iceland, and Japan, despite their joining to the IWC, as well as the aboriginal communities of Siberia, Alaska, and northern Canada. Subsistence hunters typically use whale products for themselves and depend on them for survival. National and international authorities have given special treatment to aboriginal hunters since their methods of hunting are seen as less destructive and wasteful. This distinction is being questioned as these aboriginal groups are using more modern weaponry and mechanized transport to hunt with, and are selling whale products in the marketplace. Some anthropologists argue that the term "subsistence" should also apply to these cash-based exchanges as long as they take place within local production and consumption. In 1946, the IWC placed a moratorium, limiting the annual whale catch. Since then, yearly profits for these "subsistence" hunters have been close to US$31 million (UK£20m) per year.

Other threats

Further information: Cetacean bycatch and Cetacean stranding

Whales can also be threatened by humans more indirectly. They are unintentionally caught in fishing nets by commercial fisheries as bycatch and accidentally swallow fishing hooks. Gillnetting and Seine netting is a significant cause of mortality in whales and other marine mammals. Species commonly entangled include beaked whales. Whales are also affected by marine pollution. High levels of organic chemicals accumulate in these animals since they are high in the food chain. They have large reserves of blubber, more so for toothed whales as they are higher up the food chain than baleen whales. Lactating mothers can pass the toxins on to their young. These pollutants can cause gastrointestinal cancers and greater vulnerability to infectious diseases. They can also be poisoned by swallowing litter, such as plastic bags. Environmentalists speculate that advanced naval sonar endangers some whales. Some scientists suggest that sonar may trigger whale beachings, and they point to signs that such whales have experienced decompression sickness.

Conservation

Further information: Whaling controversy

World map showing International Whaling Commission (IWC) members in blue

The scale of whale harvesting decreased substantially after 1946 when, in response to the steep decline in whale populations, the International Whaling Commission placed a moratorium which set a catch limit for each country; this excluded aboriginal groups up until 2004. As of 2015, aboriginal communities are allowed to take 280 bowhead whales off of Alaska and two from the western coast of Greenland, 620 gray whales off of Washington state, three common minke whales off of the eastern coast of Greenland and 178 on their western coast, 10 fin whales from the west coast of Greenland, nine humpback whales from the west coast of Greenland and 20 off of St. Vincent and the Grenadines each year. Several species that were commercially exploited have rebounded in numbers; for example, Grey whales may be as numerous as they were prior to harvesting, but the North Atlantic population is functionally extinct. Conversely, the North Atlantic right whale was extirpated from much of its former range, which stretched across the North Atlantic, and only remains in small fragments along the coast of Canada, Greenland, and is considered functionally extinct along the European coastline.

The IWC has designated two whale sanctuaries: the Southern Ocean Whale Sanctuary, and the Indian Ocean Whale Sanctuary. The Southern Ocean whale sanctuary spans 30,560,860 square kilometres (11,799,610 sq mi) and envelopes Antarctica. The Indian Ocean whale sanctuary takes up all of the Indian Ocean south of 55°S. The IWC is a voluntary organization, with no treaty. Any nation may leave as they wish; the IWC cannot enforce any law it makes.

As of 2013, the International Union for Conservation of Nature (IUCN) recognized 86 cetacean species, 40 of which are considered whales. Six are considered at risk, as they are ranked Critically Endangered (the North Atlantic right whale), "Endangered" (blue whale, fin whale, North Pacific right whale, and sei whale), and "Vulnerable" (sperm whale). Twenty-one species have a "Data Deficient" ranking. Species that live in polar habitats are vulnerable to the effects of recent and ongoing climate change, particularly the time when pack ice forms and melts.

Whale watching

Main article: Whale watching

Whale watching off Bar Harbour, Maine

An estimated 13 million people went whale watching globally in 2008, in all oceans except the Arctic. Rules and codes of conduct have been created to minimize harassment of the whales. Iceland, Japan and Norway have both whaling and whale watching industries. Whale watching lobbyists are concerned that the most inquisitive whales, which approach boats closely and provide much of the entertainment on whale-watching trips, will be the first to be taken if whaling is resumed in the same areas. Whale watching generated US$2.1 billion (UK£1.4 billion) per annum in tourism revenue worldwide, employing around 13,000 workers. In contrast, the whaling industry, with the moratorium in place, generates US$31 million (UK£20 million) per year. The size and rapid growth of the industry has led to complex and continuing debates with the whaling industry about the best use of whales as a natural resource.

In myth, literature and art

Engraving by William van der Gouwen depicting a stranded sperm whale being butchered on the Dutch coast, 1598

Whalers off Twofold Bay, New South Wales. Watercolour by Oswald Brierly, 1867

As marine creatures that reside in either the depths or the poles, humans knew very little about whales over the course of history; many feared or revered them. The Nords and various arctic tribes revered the whale as they were important pieces of their lives. In Inuit creation myths, when 'Big Raven', a deity in human form, found a stranded whale, he was told by the Great Spirit where to find special mushrooms that would give him the strength to drag the whale back to the sea and thus, return order to the world. In an Icelandic legend, a man threw a stone at a fin whale and hit the blowhole, causing the whale to burst. The man was told not to go to sea for twenty years, but during the nineteenth year he went fishing and a whale came and killed him.

Whales played a major part in shaping the art forms of many coastal civilizations, such as the Norse, with some dating to the Stone Age. Petroglyphs off a cliff face in Bangudae, South Korea show 300 depictions of various animals, a third of which are whales. Some show particular detail in which there are throat pleats, typical of rorquals. These petroglyphs show these people, of around 7,000 to 3,500 B.C.E. in South Korea, had a very high dependency on whales.

The Pacific Islanders and Australian Aborigines viewed whales as bringers of good and joy. One exception is French Polynesia, where in many parts, cetaceans are met with great brutality.

In Vietnam and Ghana, among other places, whales hold a sense of divinity. They are so respected in their cultures that they occasionally hold funerals for beached whales, a throwback to Vietnam's ancient sea-based Austro-Asiatic culture. The god of the seas, according to Chinese folklore, was a large whale with human limbs.

Whales have also played a role in sacred texts such as the Bible. It mentions whales in Genesis 1:21, Job 7:12, and Ezekiel 32:2. The "leviathan" described at length in Job 41:1-34 is generally understood to refer to a whale. The "sea monsters" in Lamentations 4:3 have been taken by some to refer to marine mammals, in particular whales, although most modern versions use the word "jackals" instead. The story of Jonah being swallowed by a great fish is told both in the Qur'an and in the Bible. A medieval column capital sculpture depicting this was made in the 12th century in the abbey church in Mozac, France. The Old Testament contains the Book of Jonah and in the New Testament, Jesus mentions this story in Matthew 12:40.

In 1585, Alessandro Farnese, 1585, and Francois, Duke of Anjou, 1582, were greeted on his ceremonial entry into the port city of Antwerp by floats including "Neptune and the Whale", indicating at least the city's dependence on the sea for its wealth.

Whales continue to be prevalent in modern literature. For example, Herman Melville's Moby Dick features a "great white whale" as the main antagonist for Ahab, who eventually is killed by it. The whale is an albino sperm whale, considered by Melville to be the largest type of whale, and is partly based on the historically attested bull whale Mocha Dick. Rudyard Kipling's Just So Stories includes the story of "How the Whale got in his Throat". Niki Caro's film the Whale Rider has a Māori girl ride a whale in her journey to be a suitable heir to the chief-ship. Walt Disney's film Pinocchio features a giant whale named Monstro as the final antagonist. Alan Hovhaness' orchestra And God Created Great Whales including the recorded sounds of humpback and bowhead whales. Léo Ferré's song "Il n'y a plus rien" is an example of biomusic that begins and ends with recorded whale songs mixed with a symphonic orchestra and his voice.

In captivity

Main article: Cetaceans in captivity

Further information: Beluga whale § Captivity

Beluga whales and trainers in an aquarium

Belugas were the first whales to be kept in captivity. Other species were too rare, too shy, or too big. The first beluga was shown at Barnum's Museum in New York City in 1861. For most of the 20th century, Canada was the predominant source of wild belugas. 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. Russia has become the largest provider since it had been banned in Canada. Belugas are caught in the Amur River delta and their eastern coast, and then are either transported domestically to aquariums or dolphinariums in Moscow, St. Petersburg, and Sochi, or exported to other countries, such as Canada. Most captive belugas are caught in the wild, since captive-breeding programs are not very successful.

As of 2006, 30 belugas were in Canada and 28 in the United States, and 42 deaths in captivity had been reported up to that time. A single specimen can reportedly fetch up to US$100,000 (UK£64,160) on the market. The beluga's popularity is due to its unique colour 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.

Between 1960 and 1992, the Navy carried out a program that included the study of marine mammals' abilities with sonar, with the objective of improving the detection of underwater objects. A large number of belugas were used from 1975 on, the first being dolphins. The program also included training them to carry equipment and material to divers working underwater by holding cameras in their mouths to locate lost objects, survey ships and submarines, and underwater monitoring. A similar program was used by the Russian Navy during the Cold War, in which belugas were also trained for antimining operations in the Arctic.

Aquariums have tried housing other species of whales in captivity. The success of belugas turned attention to maintaining their relative, the narwhal, in captivity. However, in repeated attempts in the 1960s and 1970s, all narwhals kept in captivity died within months. A breeding pair of pygmy right whales were retained in an enclosed area (with nets); they were eventually released in South Africa. Gigi, a gray whale calf, was kept at SeaWorld San Diego. Gigi was an orphaned calf that beached itself, and was transported two miles to SeaWorld. The 680 kilograms (1,500 lb) calf was a popular attraction, and behaved normally, despite being separated from his mother. A year later, the 8,164.7 kilograms (18,000 lb) whale grew too big to keep in captivity and was released; it was the first of two baleen whales, the other being another gray whale calf named JJ, to be kept in captivity.

Cryptid whales

Main article: Cryptid whale

Over the last few hundred years of human history, sailors and whalers have reported seeing whales they cannot identify. Some of the most well-known of these purported whales are Giglioli's whale, the rhinoceros dolphin, Trunko and the Alula whale. Giglioli's whale is a purported species of baleen whale observed by Enrico Hillyer Giglioli. The rhinoceros dolphin, or Delphinus rhinoceros or Cetodipteros rhinoceros, is a cryptid species of dolphin-oid said to have two dorsal fins, much like Giglioli's whale, but one of the dorsal fins is on the head (hence the name "rhinoceros dolphin"), allegedly sighted off the coast of the Sandwich Islands and New South Wales by Jean René Constant Quoy and Joseph Gaimard. Trunko is the nickname for a whale-like creature reportedly sighted in Margate, South Africa in 1924. The high-finned sperm whale, or Physeter tursio, is a supposed variant or relative of the known sperm whale, Physeter macrocephalus, said to live in the seas around the Shetland Islands, the Southern Ocean, and Nova Scotia. The Alula whale, or the Alula Killer, is a cryptid that resembles a sepia brown killer whale with a well-rounded forehead and white, star-like scars on the body. The dorsal fin, supposedly 60 centimetres (24 in) high, is prominent and often protrudes well above the surface of the water.