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The toothed whales (also called odontocetes,
systematic name A systematic name is a name given in a systematic way to one unique group, organism, object or chemical substance, out of a specific population or collection. Systematic names are usually part of a nomenclature. A semisystematic name or semitrivial ...
Odontoceti) are a
parvorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
of
cetacea Cetacea (; , ) is an infraorder of aquatic mammals 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 them ...
ns that includes
dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (the ...
s,
porpoise Porpoises are a group of fully aquatic marine mammals, all of which are classified under the family Phocoenidae, parvorder Odontoceti (toothed whales). Although similar in appearance to dolphins, they are more closely related to narwhals an ...
s, and all other
whale Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s possessing
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
, such as the
beaked whale Beaked whales (systematic name Ziphiidae) are a family of cetaceans noted as being one of the least known groups of mammals because of their deep-sea habitat and apparent low abundance. Only three or four of the 24 species are reasonably well-k ...
s and
sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...
s. Seventy-three species of toothed whales are described. They are one of two living groups of cetaceans, the other being the
baleen whale Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...
s (Mysticeti), which have
baleen Baleen is a filter-feeding system inside the mouths of baleen whales. To use baleen, the whale first opens its mouth underwater to take in water. The whale then pushes the water out, and animals such as krill are filtered by the baleen and ...
instead of teeth. The two groups are thought to have diverged around 34
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
(mya). Toothed whales range in size from the and
vaquita The vaquita ( ; ''Phocoena sinus'') is a species of porpoise endemic to the northern end of the Gulf of California in Baja California, Mexico. Averaging (females) or (males) in length, it is the smallest of all living cetaceans. The species i ...
to the and
sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...
. Several species of odontocetes exhibit
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, in that there are size or other morphological differences between females and males. They have streamlined bodies and two limbs that are modified into flippers. Some can travel at up to 20 knots. Odontocetes have conical teeth designed for catching fish or squid. They have well-developed hearing, that is well adapted for both air and water, so much so that some can survive even if they are blind. Some species are well adapted for diving to great depths. Almost all have a layer of fat, or
blubber Blubber is a thick layer of vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians. Description Lipid-rich, collagen fiber-laced blubber comprises the hypodermis and covers the whole body, except for pa ...
, under the skin to keep warm in the cold water, with the exception of
river dolphin River dolphins are a polyphyletic group of fully aquatic mammals that reside exclusively in freshwater or brackish water. They are an informal grouping of dolphins, which itself is a paraphyletic group within the infraorder Cetacea. Extant riv ...
s. Toothed whales consist of some of the most widespread mammals, but some, as with the vaquita, are restricted to certain areas. Odontocetes feed largely on fish and squid, but a few, like the
orca The orca or killer whale (''Orcinus orca'') is a toothed whale belonging to the oceanic dolphin family, of which it is the largest member. It is the only Extant taxon, extant species in the genus ''Orcinus'' and is recognizable by its black ...
, feed on mammals, such as
pinnipeds Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammal, marine mammals. They comprise the extant taxon, extant family (biology ...
. Males typically mate with multiple females every year, making them
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
. Females mate every two to three years. Calves are typically born in the spring and summer, and females bear the responsibility for raising them, but more sociable species rely on the family group to care for calves. Many species, mainly dolphins, are highly sociable, with some pods reaching over a thousand individuals. Once hunted for their products, cetaceans are now protected by international law. Some species are attributed with high levels of intelligence. At the 2012 meeting of the
American Association for the Advancement of Science The American Association for the Advancement of Science (AAAS) is an American international non-profit organization with the stated goals of promoting cooperation among scientists, defending scientific freedom, encouraging scientific respons ...
, support was reiterated for a cetacean bill of rights, listing cetaceans as nonhuman persons. Besides
whaling Whaling is the process of hunting of whales for their usable products such as meat and blubber, which can be turned into a type of oil that became increasingly important in the Industrial Revolution. It was practiced as an organized industry ...
and drive hunting, they also face threats from bycatch and marine pollution. The
baiji The baiji (; IPA: ; ''Lipotes vexillifer'', ''Lipotes'' meaning "left behind" and ''vexillifer'' "flag bearer") is a possibly extinct species of freshwater dolphin native to the Yangtze river system in China. It is thought to be the first dolph ...
, for example, is considered functionally extinct by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
, with the last sighting in 2004, due to heavy pollution to the
Yangtze River The Yangtze or Yangzi ( or ; ) is the longest list of rivers of Asia, river in Asia, the list of rivers by length, third-longest in the world, and the longest in the world to flow entirely within one country. It rises at Jari Hill in th ...
. Whales occasionally feature in literature and film, as in the great white sperm whale of
Herman Melville Herman Melville (Name change, born Melvill; August 1, 1819 – September 28, 1891) was an American people, American novelist, short story writer, and poet of the American Renaissance (literature), American Renaissance period. Among his bes ...
's ''
Moby-Dick ''Moby-Dick; or, The Whale'' is an 1851 novel by American writer Herman Melville. The book is the sailor Ishmael (Moby-Dick), Ishmael's narrative of the obsessive quest of Captain Ahab, Ahab, captain of the whaler, whaling ship ''Pequod (Moby- ...
''. Small odontocetes, mainly dolphins, are kept in captivity and trained to perform tricks.
Whale watching Whale watching is the practice of observing whales and dolphins ( cetaceans) in their natural habitat. Whale watching is mostly a recreational activity (cf. birdwatching), but it can also serve scientific and/or educational purposes.Hoyt, E. 2 ...
has become a form of tourism around the world.


Taxonomy


Research history

In
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
's time, the fourth century BC, whales were regarded as fish due to their superficial similarity. Aristotle, however, could already see many physiological and anatomical similarities with the terrestrial vertebrates, such as blood (circulation), lungs, uterus, and fin anatomy. His detailed descriptions were assimilated by the Romans, but mixed with a more accurate knowledge of the dolphins, as mentioned by
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic '' ...
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 harbor porpoise is one of the most accessible species for early
cetologists Cetology (from Greek , ''kētos'', "whale"; and , ''-logia'') or whalelore (also known as whaleology) is the branch of marine mammal science that studies the approximately eighty species of whales, dolphins, and porpoises in the scientific order ...
, because it could be seen very close to land, inhabiting shallow coastal areas of Europe. Many of the findings that apply to all cetaceans were therefore first discovered in the porpoises. One of the first anatomical descriptions of the airways of the whales on the basis of a harbor porpoise dates from 1671 by John Ray. It nevertheless referred to the porpoise as a fish.


Evolution

Toothed whales, as well as baleen whales, are descendants of land-dwelling mammals of the
artiodactyl The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
(even-toed ungulates). They are closely related to the
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
, sharing a common ancestor that lived around 54
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
(mya). The primitive cetaceans, or
archaeocetes Archaeoceti ("ancient whales"), or Zeuglodontes in older literature, is a paraphyletic group of primitive cetaceans that lived from the Early Eocene to the late Oligocene (). Representing the earliest cetacean radiation, they include the initial ...
, first took to the sea approximately 49 mya and became fully aquatic by 5–10 million years later. The ancestors of toothed whales and baleen whales diverged in the early Oligocene. This was due to a change in the climate of the southern oceans that affected where the environment of the plankton that these whales ate. The adaptation of echolocation and enhanced fat synthesis in blubber occurred when toothed whales split apart from baleen whales, and distinguishes modern toothed whales from fully aquatic archaeocetes. This happened around 34 mya. Unlike toothed whales, baleen whales do not have wax ester deposits nor branched fatty chain acids in their blubber. Thus, more recent evolution of these complex blubber traits occurred after baleen whales and toothed whales split, and only in the toothed whale lineage. Modern toothed whales do not rely on their sense of sight, but rather on their sonar to hunt prey. Echolocation also allowed toothed whales to dive deeper in search of food, with light no longer necessary for navigation, which opened up new food sources. Toothed whales (
Odontocetes The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales possessing teeth, such as the beaked whales and sperm whales. Seventy-three species of t ...
) echolocate by creating a series of clicks emitted at various frequencies. Sound pulses are emitted through their melon-shaped foreheads, reflected off objects, and retrieved through the lower jaw. Skulls of ''
Squalodon ''Squalodon'' is an extinct genus of whales of the Oligocene and Miocene epochs, belonging to the family Squalodontidae. Named by Jean-Pierre Sylvestre de Grateloup in 1840, it was originally believed to be an iguanodontid dinosaur but has since ...
'' show evidence for the first hypothesized appearance of echolocation. ''Squalodon'' lived from the early to middle
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
to the middle
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
, around 33-14 mya. ''Squalodon'' featured several commonalities with modern Odontocetes. The cranium was well compressed, the rostrum telescoped outward (a characteristic of the modern parvorder
Odontoceti The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales possessing teeth, such as the beaked whales and sperm whales. Seventy-three species of ...
), giving ''Squalodon'' an appearance similar to that of modern toothed whales. However, it is thought unlikely that squalodontids are direct ancestors of living dolphins.


Biology


Anatomy

Toothed whales have torpedo-shaped bodies with inflexible necks, limbs modified into flippers, nonexistent external ear flaps, a large tail fin, and bulbous heads (with the exception of
sperm whales The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the Genus (biology), genus ''Physeter'' and one of three extant species in the s ...
). Their skulls have small eye orbits, long beaks (with the exception sperm whales), and eyes placed on the sides of their heads. Toothed whales range in size from the and
vaquita The vaquita ( ; ''Phocoena sinus'') is a species of porpoise endemic to the northern end of the Gulf of California in Baja California, Mexico. Averaging (females) or (males) in length, it is the smallest of all living cetaceans. The species i ...
to the and
sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...
. Overall, they tend to be dwarfed by their relatives, the
baleen whales Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...
(Mysticeti). Several species have 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 Cementum is a specialized calcified substance covering the root of a tooth. The cementum is the part of the periodontium that attaches the teeth to the alveolar bone by anchoring the periodontal ligament.Illustrated Dental Embryology, Histology, a ...
cells overlying
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
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. Except for the sperm whale, most toothed whales are smaller than the baleen whales. The teeth differ considerably among the species. They may be numerous, with some dolphins bearing over 100 teeth in their jaws. At the other extreme are the
narwhal The narwhal, also known as a narwhale (''Monodon monoceros''), is a medium-sized toothed whale that possesses a large "tusk" from a protruding canine tooth. It lives year-round in the Arctic waters around Greenland, Canada and Russia. It is o ...
s with their single long tusks and the almost toothless
beaked whales Beaked whales (systematic name Ziphiidae) are a family of cetaceans noted as being one of the least known groups of mammals because of their deep-sea habitat and apparent low abundance. Only three or four of the 24 species are reasonably well-kn ...
with tusk-like teeth only in males. Not all species are believed to use their teeth for feeding. For instance, the sperm whale likely uses its teeth for aggression and showmanship. Breathing involves expelling stale air from their one blowhole, forming an upward, steamy spout, followed by inhaling fresh air into the lungs. Spout shapes differ among species, which facilitates identification. The spout only forms when warm air from the lungs meets cold air, so it does not form in warmer climates, as with river dolphins. Almost all cetaceans have a thick layer of
blubber Blubber is a thick layer of vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians. Description Lipid-rich, collagen fiber-laced blubber comprises the hypodermis and covers the whole body, except for pa ...
, with the exception of river dolphins. In species that live near the poles, the blubber can be as thick as . This blubber can help with buoyancy, protection to some extent as predators would have a hard time getting through a thick layer of fat, energy for fasting during leaner times, and insulation from the harsh climates. Calves are born with only a thin layer of blubber, but some species compensate for this with thick
lanugo Lanugo is very thin, soft, usually unpigmented, downy hair that is sometimes found on the body of a fetus or newborn. It is the first hair to be produced by the fetal hair follicles, and it usually appears around sixteen weeks of gestation and is ...
s. Toothed whales have also evolved the ability to store large amounts of wax esters in their adipose tissue as an addition to or in complete replacement of other fats in their blubber. They can produce isovaleric acid from branched chain fatty acids (BCFA). These adaptations are unique, are only in more recent, derived lineages and were likely part of the transition for species to become deeper divers as the families of toothed whales (Physeteridae, Kogiidae, and Ziphiidae) that have the highest quantities of wax esters and BCFAs in their blubber are also the species that dive the deepest and for the longest amount of time. Toothed whales have a two-chambered stomach similar in structure to terrestrial carnivores. They have
fundic The gastric glands are glands in the lining of the stomach that play an essential role in the process of digestion. All of the glands have mucus-secreting foveolar cells. Mucus lines the entire stomach, and protects the stomach lining from the e ...
and
pyloric The pylorus ( or ), or pyloric part, connects the stomach to the duodenum. The pylorus is considered as having two parts, the ''pyloric antrum'' (opening to the body of the stomach) and the ''pyloric canal'' (opening to the duodenum). The ''pylori ...
chambers.


Locomotion

Cetaceans have two flippers on the front, and a tail fin. These flippers contain four digits. Although toothed whales do not possess fully developed hind limbs, some, such as the sperm whale, possess discrete rudimentary appendages, which may contain feet and digits. Toothed whales are fast swimmers in comparison to seals, which typically cruise at 5–15 knots, or ; the sperm whale, in comparison, can travel at speeds of up to . The fusing of the neck vertebrae, while increasing stability when swimming at high speeds, decreases flexibility, rendering them incapable of turning their heads; river dolphins, however, have unfused neck vertebrae and can turn their heads. When swimming, toothed whales rely on their tail fins to propel them through the water. Flipper movement is continuous. They 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 Log most often refers to: * Trunk (botany), the stem and main wooden axis of a tree, called logs when cut ** Logging, cutting down trees for logs ** Firewood, logs used for fuel ** Lumber or timber, converted from wood logs * Logarithm, in mathe ...
out of the water, which may allow them to travel faster. Their skeletal anatomy allows them to be fast swimmers. Most species have a
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through conv ...
. Most toothed whales are adapted for diving to great depths, porpoises are one exception. 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 Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
and
myoglobin Myoglobin (symbol Mb or MB) is an iron- and oxygen-binding protein found in the cardiac and skeletal muscle tissue of vertebrates in general and in almost all mammals. Myoglobin is distantly related to hemoglobin. Compared to hemoglobin, myoglobi ...
store oxygen in body tissue; and they have twice the concentration of myoglobin than haemoglobin. Before going on long dives, many toothed 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

Toothed whale eyes are relatively small for their size, yet they do retain a good degree of eyesight. As well as this, the eyes are placed on the sides of its head, so their vision consists of two fields, rather than a binocular view as humans have. When a
beluga The beluga whale (/bɪˈluːɡə/) (Delphinapterus leucas) is an Arctic and sub-Arctic cetacean. It is one of two members of the family Monodontidae, along with the narwhal, and the only member of the genus Delphinapterus. It is also known as the wh ...
surfaces, its lenses and corneas correct the nearsightedness that results from the refraction of light; they contain both rod and
cone A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines con ...
cells, meaning they can see in both dim and bright light. They do, however, lack short wavelength-sensitive visual pigments in their cone cells, indicating a more limited capacity for colour vision than most mammals. Most toothed whales have slightly flattened eyeballs, enlarged pupils (which shrink as they surface to prevent damage), slightly flattened corneas, and a
tapetum lucidum The ''tapetum lucidum'' ( ; ; ) is a layer of tissue in the eye of many vertebrates and some other animals. Lying immediately behind the retina, it is a retroreflector. It reflects visible light back through the retina, increasing the light a ...
; 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 ahead of itself, but 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 The olfactory bulb (Latin: ''bulbus olfactorius'') is a grey matter, neural structure of the vertebrate forebrain involved in olfaction, the sense of odor, smell. It sends olfactory information to be further processed in the amygdala, the orbitof ...
are absent in toothed whales, and unlike baleen whales, they lack the
vomeronasal organ The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. ...
, suggesting they have no sense of smell. Toothed whales are not thought to have a good sense of taste, as their taste buds are atrophied or missing altogether. However, some dolphins have preferences between different kinds of fish, indicating some sort of attachment to taste.


Sonar

Toothed whales are capable of making a broad range of sounds using nasal airsacs located just below the blowhole. Roughly three categories of sounds can be identified:
frequency-modulated Frequency modulation (FM) is the encoding of information in a carrier wave by varying the instantaneous frequency of the wave. The technology is used in telecommunications, radio broadcasting, signal processing, and computing. In analog freque ...
whistles, burst-pulsed sounds, and clicks. Dolphins communicate with whistle-like sounds produced by vibrating connective tissue, similar to the way human vocal cords function, and through burst-pulsed sounds, though the nature and extent of that ability is not known. The clicks are directional and are used for echolocation, often occurring in a short series called a click train. The click rate increases when approaching an object of interest. Toothed whale
biosonar Echolocation, also called bio sonar, is a biological sonar used by several animal species. Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these ...
clicks are amongst the loudest sounds made by marine animals. The cetacean ear has specific adaptations to the
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
environment. In humans, the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
works as an impedance equalizer between the outside air's low impedance and the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory org ...
r fluid's high impedance. In whales, and other marine mammals, no great difference exists 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 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 A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. The word "melon" can refer to either the plant or specifically to the fruit. Botanically, a melon is a kind of berry, specifically a "pepo". The ...
. 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 it is, the more dependent they are on it. A beaked whale, for example, has a small bulge sitting on top of its skull, whereas a sperm whale's head is filled mainly with the melon. Odontocetes are well adapted to hear sounds at
ultrasonic Ultrasound is sound waves with frequencies higher than the upper audible limit of human hearing. Ultrasound is not different from "normal" (audible) sound in its physical properties, except that humans cannot hear it. This limit varies fr ...
frequencies, as opposed to
mysticetes Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in the ...
who generally hear sounds within the range of
infrasonic Infrasound, sometimes referred to as low status sound, describes sound waves with a frequency below the lower limit of human audibility (generally 20 Hz). Hearing becomes gradually less sensitive as frequency decreases, so for humans to perce ...
frequencies. Bottlenose dolphins have been found to have signature whistles unique to a specific individual. These whistles are used for dolphins to communicate with one another by identifying an individual. It can be seen as the dolphin equivalent of a
name A name is a term used for identification by an external observer. They can identify a class or category of things, or a single thing, either uniquely, or within a given context. The entity identified by a name is called its referent. A personal ...
for humans. Because dolphins are generally associated in groups, communication is necessary. Signal masking is when other similar sounds (conspecific sounds) interfere with the original acoustic sound. In larger groups, individual whistle sounds are less prominent. Dolphins tend to travel in pods, in some instances including up to 600 members.


Life history and behaviour


Intelligence

Cetaceans are known to communicate and therefore are able to teach, learn, cooperate, scheme, and grieve. The
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
of many species of dolphins is home to elongated
spindle neurons Von Economo neurons (VENs), also called spindle neurons, are a specific class of mammalian cortical neurons characterized by a large spindle-shaped soma (or body) gradually tapering into a single apical axon (the ramification that ''transmits' ...
that, prior to 2007, were known only in hominids. In humans, these cells are involved in social conduct, emotions, judgement, and theory of mind. Dolphin spindle neurons are found in areas of the brain homologous to where they are found in humans, suggesting they perform a similar function.
Brain size The size of the brain is a frequent topic of study within the fields of anatomy, biological anthropology, animal science and evolution. Brain size is sometimes measured by weight and sometimes by volume (via MRI scans or by skull volume). Neur ...
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 Allometry is the study of the relationship of body size to shape, anatomy, physiology and finally behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in ''On Growth and Form'' and by Julian Huxley in 1932. Overview Allom ...
analysis indicates that mammalian brain size scales around the two-thirds or three-quarters 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 Encephalization quotient (EQ), encephalization level (EL), or just encephalization is a relative brain size measure that is defined as the ratio between observed to predicted brain mass for an animal of a given size, based on nonlinear regressi ...
that can be used as another indication of animal intelligence. Sperm whales have the largest brain mass of any animal on earth, averaging and in mature males, in comparison to the average human brain which averages in mature males. The
brain to body mass ratio A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
in some odontocetes, such as belugas and narwhals, is second only to humans. Dolphins are known to engage in complex
play behaviour Play is a range of intrinsically motivated activities done for recreational pleasure and enjoyment. Play is commonly associated with children and juvenile-level activities, but may be engaged in at any life stage, and among other higher-functio ...
, which includes such things as producing stable underwater
toroid In mathematics, a toroid is a surface of revolution with a hole in the middle. The axis of revolution passes through the hole and so does not intersect the surface. For example, when a rectangle is rotated around an axis parallel to one of its ...
al air-core
vortex In fluid dynamics, a vortex ( : vortices or vortexes) is a region in a fluid in which the flow revolves around an axis line, which may be straight or curved. Vortices form in stirred fluids, and may be observed in smoke rings, whirlpools in th ...
rings or "
bubble ring A bubble ring, or toroidal bubble, is an underwater vortex ring where an air bubble occupies the core of the vortex, forming a ring shape. The ring of air as well as the nearby water spins poloidally as it travels through the water, much like a ...
s". Two main methods of bubble ring production are: 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 Helical may refer to: * Helix, the mathematical concept for the shape * Helical engine, a proposed spacecraft propulsion drive * Helical spring, a coilspring * Helical plc, a British property company, once a maker of steel bar stock * Helicoil A t ...
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. Dolphins are known to use this method during hunting. Dolphins have also been known to use tools. In
Shark Bay Shark Bay (Malgana: ''Gathaagudu'', "two waters") is a World Heritage Site in the Gascoyne region of Western Australia. The http://www.environment.gov.au/heritage/places/world/shark-bay area is located approximately north of Perth, on the ...
, a population of
Indo-Pacific bottlenose dolphin The Indo-Pacific bottlenose dolphin (''Tursiops aduncus'') is a species of bottlenose dolphin. This dolphin grows to long, and weighs up to . It lives in the waters around India, northern Australia, South China, the Red Sea, and the eastern co ...
s put sponges on their beak to protect them from abrasions and sting ray barbs while foraging in the seafloor. This behaviour is passed on from mother to daughter, and it is only observed in 54 female individuals.
Self-awareness In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifesty ...
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 metacognitive 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 The mirror test—sometimes called the mark test, mirror self-recognition (MSR) test, red spot technique, or rouge test—is a behavioral technique developed in 1970 by American psychologist Gordon Gallup Jr. as an attempt to determine whether an ...
, in which a temporary dye is placed on an animal's body, and the animal is then presented with a mirror; then whether the animal shows signs of self-recognition is determined. In 1995, Marten and Psarakos used television to test dolphin self-awareness. 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 repeated since then, dolphins have since "passed" the mirror test.


Vocalisations

Dolphins are capable of making a broad range of sounds using nasal airsacs located just below the blowhole. Roughly three categories of sounds can be identified:
frequency modulated Frequency modulation (FM) is the encoding of information in a carrier wave by varying the instantaneous frequency of the wave. The technology is used in telecommunications, radio broadcasting, signal processing, and computing. In analog freque ...
whistles, burst-pulsed sounds and clicks. Dolphins communicate with whistle-like sounds produced by vibrating connective tissue, similar to the way human vocal cords function, and through burst-pulsed sounds, though the nature and extent of that ability is not known. The clicks are directional and are for echolocation, often occurring in a short series called a click train. The click rate increases when approaching an object of interest. Dolphin echolocation clicks are amongst the loudest sounds made by marine animals. Bottlenose dolphins have been found to have signature whistles, a whistle that is unique to a specific individual. These whistles are used in order for dolphins to communicate with one another by identifying an individual. It can be seen as the dolphin equivalent of a name for humans. These signature whistles are developed during a dolphin's first year; it continues to maintain the same sound throughout its lifetime. An auditory experience influences the whistle development of each dolphin. Dolphins are able to communicate to one another by addressing another dolphin through mimicking their whistle. The signature whistle of a male bottlenose dolphin tends to be similar to that of his mother, while the signature whistle of a female bottlenose dolphin tends to be more identifying. Bottlenose dolphins have a strong memory when it comes to these signature whistles, as they are able to relate to a signature whistle of an individual they have not encountered for over twenty years. Research done on signature whistle usage by other dolphin species is relatively limited. The research on other species done so far has yielded varied outcomes and inconclusive results. Sperm whales can produce three specific vocalisations: creaks, codas, and slow clicks. A creak is a rapid series of high-frequency clicks that sounds somewhat like a creaky door hinge. It is typically used when homing in on prey. A coda is a short pattern of 3 to 20 clicks that is used in social situations to identify one another (like a signature whistle), but it is still unknown whether sperm whales possess individually specific coda repertoires or whether individuals make codas at different rates. Slow clicks are heard only in the presence of males (it is not certain whether females occasionally make them). Males make a lot of slow clicks in breeding grounds (74% of the time), both near the surface and at depth, which suggests they are primarily mating signals. Outside breeding grounds, slow clicks are rarely heard, and usually near the surface.


Foraging and predation

All whales are
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
and
predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
. Odontocetes, as a whole, mostly feed on fish and
cephalopods A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, an ...
, and then followed by
crustaceans Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
and
bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
. All species are generalist and opportunistic feeders. Some may forage with other kinds of animals, such as other species of whales or certain species of
pinnipeds Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammal, marine mammals. They comprise the extant taxon, extant family (biology ...
. One common feeding method is herding, where a pod squeezes a school of fish into a small volume, known as a
bait ball A bait ball, or baitball, occurs when small fish swarm in a tightly packed spherical formation about a common centre. It is a last-ditch defensive measure adopted by small schooling fish when they are threatened by predators. Small schooling f ...
. Individual members then take turns plowing through the ball, feeding on the stunned fish. Coralling is a method where dolphins chase fish into shallow water to catch them more easily. Orcas and bottlenose dolphins have also been known to drive their prey onto a beach to feed on it, a behaviour known as beach or strand feeding. The shape of the snout may correlate with tooth number and thus feeding mechanisms. The narwhal, with its blunt snout and reduced dentition, relies on
suction feeding Aquatic feeding mechanisms face a special difficulty as compared to feeding on land, because the density of water is about the same as that of the prey, so the prey tends to be pushed away when the mouth is closed. This problem was first identifi ...
. Sperm whales usually dive between , and sometimes , in search of food. Such dives can last more than an hour. They feed on several species, notably the
giant squid The giant squid (''Architeuthis dux'') is a species of deep-ocean dwelling squid in the family Architeuthidae. It can grow to a tremendous size, offering an example of abyssal gigantism: recent estimates put the maximum size at around Trace ...
, but also the
colossal squid The colossal squid (''Mesonychoteuthis hamiltoni'') is part of the family Cranchiidae. It is sometimes called the Antarctic squid or giant cranch squid and is believed to be the largest squid species in terms of mass. It is the only recognize ...
,
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttle ...
es, and fish like
demersal The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer of ...
ray Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (g ...
s, but their diet is mainly medium-sized
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
. Some prey may be taken accidentally while eating other items. A study in the Galápagos found that squid from the genera ''
Histioteuthis ''Histioteuthis'' is a genus of squid in the family Histioteuthidae. It goes by the common name cock-eyed squid, because in all species the right eye is normal-sized, round, blue and sunken; whereas the left eye is at least twice the diameter of ...
'' (62%), ''
Ancistrocheirus ''Ancistrocheirus lesueurii'', the sharpear enope squid, is the only species in the genus ''Ancistrocheirus'' and family Ancistrocheiridae. With a mantle length of 25 cm, this moderately sized squid may be found throughout the tropical and ...
'' (16%), and ''Octopoteuthis'' (7%) weighing between were the most commonly taken. Battles between sperm whales and giant squid or colossal squid have never been observed by humans; however, white scars are believed to be caused by the large squid. A 2010 study suggests that female sperm whales may collaborate when hunting
Humboldt squid The Humboldt squid (''Dosidicus gigas''), also known as jumbo squid or jumbo flying squid (EN), and Pota in Peru or Jibia in Chile (ES) is a large, predatory squid living in the eastern Pacific Ocean. It is the only known species of the genus ''D ...
. The orca is known to prey on numerous other toothed whale species. One example is the
false killer whale The false killer whale (''Pseudorca crassidens'') is a species of oceanic dolphin that is the only extant representative of the genus ''Pseudorca''. It is found in oceans worldwide but mainly in tropical regions. It was first described in 1846 ...
. To subdue and kill whales, orcas continually ram them with their heads; this can sometimes kill
bowhead whale The bowhead whale (''Balaena mysticetus'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus ''Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters, ...
s, or severely injure them. Other times, they corral their prey before striking. They are typically hunted by groups of 10 or fewer orca, but they are seldom attacked by an individual. Calves are more commonly taken by orca, but adults can be targeted, as well. Groups even attack larger cetaceans such as
minke whale The minke whale (), or lesser rorqual, is a species complex of baleen whale. The two species of minke whale are the common (or northern) minke whale and the Antarctic (or southern) minke whale. The minke whale was first described by the Danish n ...
s,
gray whale The gray whale (''Eschrichtius robustus''), also known as the grey whale,Britannica Micro.: v. IV, p. 693. gray back whale, Pacific gray whale, Korean gray whale, or California gray whale, is a baleen whale that migrates between feeding and bree ...
s, and rarely sperm whales or
blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can ...
s. Other marine mammal prey species include nearly 20 species of
seal Seal may refer to any of the following: Common uses * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impr ...
,
sea lion Sea lions are pinnipeds characterized by external ear flaps, long foreflippers, the ability to walk on all fours, short and thick hair, and a big chest and belly. Together with the fur seals, they make up the family Otariidae, eared seals. ...
and
fur seal Fur seals are any of nine species of pinnipeds belonging to the subfamily Arctocephalinae in the family '' Otariidae''. They are much more closely related to sea lions than true seals, and share with them external ears (pinnae), relatively lon ...
. These cetaceans are targeted by terrestrial and pagophilic 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 orca. Polar bear attacks on belugas and narwhals are usually successful in winter, but rarely inflict any damage in summer. For most of the smaller species of dolphins, only a few of the larger sharks, such as the
bull shark The bull shark (''Carcharhinus leucas''), also known as the Zambezi shark (informally zambi) in Africa and Lake Nicaragua shark in Nicaragua, is a species of requiem shark commonly found worldwide in warm, shallow waters along coasts and in riv ...
, dusky shark,
tiger shark The tiger shark (''Galeocerdo cuvier'') is a species of requiem shark and the last extant member of the family Galeocerdonidae. It is a large macropredator, capable of attaining a length over . Populations are found in many tropical and tempera ...
, and
great white shark The great white shark (''Carcharodon carcharias''), also known as the white shark, white pointer, or simply great white, is a species of large mackerel shark which can be found in the coastal surface waters of all the major oceans. It is nota ...
, are a potential risk, especially for calves. Dolphins can tolerate and recover from extreme injuries (including shark bites) although the exact methods used to achieve this are not known. The
healing With physical trauma or disease suffered by an organism, healing involves the repairing of damaged tissue(s), organs and the biological system as a whole and resumption of (normal) functioning. Medicine includes the process by which the cells i ...
process is rapid and even very deep wounds do not cause dolphins to
hemorrhage Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, vag ...
to death. Even gaping wounds restore in such a way that the animal's body shape is restored, and infection of such large wounds are rare.


Life cycle

Toothed whales are fully aquatic creatures, which means their birth and courtship behaviours are very different from terrestrial and semiaquatic creatures. Since they are unable to go onto land to calve, they deliver their young with the fetus positioned for tail-first delivery. This prevents the calf from drowning either upon or during delivery. To feed the newborn, toothed whales, being aquatic, must squirt the milk into the mouth of the calf. Being mammals, they have
mammary gland A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primat ...
s used for nursing calves; they are weaned around 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, 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 10 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. In orcas,
false killer whale The false killer whale (''Pseudorca crassidens'') is a species of oceanic dolphin that is the only extant representative of the genus ''Pseudorca''. It is found in oceans worldwide but mainly in tropical regions. It was first described in 1846 ...
s,
short-finned pilot whale The short-finned pilot whale (''Globicephala macrorhynchus'') is one of the two species of cetaceans in the genus ''Pilot whale, Globicephala'', which it shares with the long-finned pilot whale (''G. melas''). It is part of the oceanic dolphin ...
s,
narwhal The narwhal, also known as a narwhale (''Monodon monoceros''), is a medium-sized toothed whale that possesses a large "tusk" from a protruding canine tooth. It lives year-round in the Arctic waters around Greenland, Canada and Russia. It is o ...
s, and belugas, there is an unusually long post-reproductive lifespan (
menopause Menopause, also known as the climacteric, is the time in women's lives when menstrual periods stop permanently, and they are no longer able to bear children. Menopause usually occurs between the age of 47 and 54. Medical professionals often d ...
) in females. Older females, though unable to have their own children, play a key role in the rearing of other calves in the pod, and in this sense, given the costs of pregnancy especially at an advanced age, extended menopause is advantageous.


Interaction with humans


Threats


Sperm whaling

The head of the sperm whale is filled with a waxy liquid called
spermaceti Spermaceti is a waxy substance found in the head cavities of the sperm whale (and, in smaller quantities, in the oils of other whales). Spermaceti is created in the spermaceti organ inside the whale's head. This organ may contain as much as of ...
. This liquid can be refined into spermaceti wax and
sperm oil Sperm oil is a waxy liquid obtained from sperm whales. It is a clear, yellowish liquid with a very faint odor. Sperm oil has a different composition from common whale oil, obtained from rendered blubber. Although it is traditionally called an " ...
. These were much sought after by 18th-, 19th-, and 20th-century whalers. These substances found a variety of commercial applications, such as
candle A candle is an ignitable wick embedded in wax, or another flammable solid substance such as tallow, that provides light, and in some cases, a fragrance. A candle can also provide heat or a method of keeping time. A person who makes candles i ...
s,
soap Soap is a salt of a fatty acid used in a variety of cleansing and lubricating products. In a domestic setting, soaps are surfactants usually used for washing, bathing, and other types of housekeeping. In industrial settings, soaps are use ...
,
cosmetics Cosmetics are constituted mixtures of chemical compounds derived from either natural sources, or synthetically created ones. Cosmetics have various purposes. Those designed for personal care and skin care can be used to cleanse or protect ...
, machine oil, other specialized lubricants, lamp oil, pencils, crayons, leather waterproofing, rustproofing materials, and many pharmaceutical compounds.
Ambergris Ambergris ( or , la, ambra grisea, fro, ambre gris), ''ambergrease'', or grey amber is a solid, waxy, flammable substance of a dull grey or blackish colour produced in the digestive system of sperm whales. Freshly produced ambergris has a mari ...
, a solid, waxy, flammable substance produced in the digestive system of sperm whales, was also sought as a fixative in
perfumery Perfume (, ; french: parfum) is a mixture of fragrant essential oils or aroma compounds (fragrances), fixatives and solvents, usually in liquid form, used to give the human body, animals, food, objects, and living-spaces an agreeable scent. Th ...
. Sperm whaling in the 18th century began with small sloops carrying only a pair of whaleboats (sometimes only one). As the scope and size of the fleet increased, so did the rig of the vessels change, as
brigs A brig is a type of sailing vessel defined by its rig: two masts which are both square rig, square-rigged. Brigs originated in the second half of the 18th century and were a common type of smaller merchant vessel or warship from then until the ...
,
schooners A schooner () is a type of sailing vessel defined by its rig: fore-and-aft rigged on all of two or more masts and, in the case of a two-masted schooner, the foremast generally being shorter than the mainmast. A common variant, the topsail schoon ...
, and finally ships and barks were introduced. In the 19th-century stubby, square-rigged ships (and later barks) dominated the fleet, being sent to the Pacific (the first being the British whaleship ''Emilia'', in 1788), the Indian Ocean (1780s), and as far away as the Japan grounds (1820) and the coast of Arabia (1820s), as well as Australia (1790s) and New Zealand (1790s). Hunting for sperm whales during this period was a notoriously dangerous affair for the crews of the 19th-century whaleboats. Although a properly harpooned sperm whale generally exhibited a fairly consistent pattern of attempting to flee underwater to the point of exhaustion (at which point it would surface and offer no further resistance), it was not uncommon for bull whales to become enraged and turn to attack pursuing whaleboats on the surface, particularly if it had already been wounded by repeated harpooning attempts. A commonly reported tactic was for the whale to invert itself and violently thrash the surface of the water with its
fluke Fluke may refer to: Biology * Fluke (fish), a species of marine flatfish * Fluke (tail), the lobes of the tail of a cetacean, such as dolphins or whales, ichthyosaurs, mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ...
, flipping and crushing nearby boats. The estimated historic worldwide sperm whale population numbered 1,100,000 before commercial sperm whaling began in the early 18th century. By 1880, it had declined an estimated 29%. From that date until 1946, the population appears to have recovered somewhat as whaling pressure lessened, but after the Second World War, with the industry's focus again on sperm whales, the population declined even further to only 33%. In the 19th century, between 184,000 and 236,000 sperm whales were estimated to have been killed by the various whaling nations, while in the modern era, at least 770,000 were taken, the majority between 1946 and 1980. Remaining sperm whale populations are large enough so that the species' conservation status is vulnerable, rather than endangered. However, the recovery from the whaling years is a slow process, particularly in the South Pacific, where the toll on males of breeding age was severe.


Drive hunting

Dolphins and porpoises are hunted in an activity known as dolphin drive hunting. This is accomplished by driving a pod together with boats and 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 Solomon Islands is an island country consisting of six major islands and over 900 smaller islands in Oceania, to the east of Papua New Guinea and north-west of Vanuatu. It has a land area of , and a population of approx. 700,000. Its capita ...
, the
Faroe Islands The Faroe Islands ( ), or simply the Faroes ( fo, Føroyar ; da, Færøerne ), are a North Atlantic island group and an autonomous territory of the Kingdom of Denmark. They are located north-northwest of Scotland, and about halfway bet ...
,
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
, and
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
, the most well-known practitioner of this method. By numbers, dolphins are mostly hunted for their
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
, though some end up in
dolphinarium A dolphinarium is an aquarium for dolphins. The dolphins are usually kept in a pool, though occasionally they may be kept in pens in the open sea, either for research or public performances. Some dolphinariums consist of one pool where dolphins pe ...
s. Despite the controversial nature of the hunt resulting in international criticism, and the possible health risk that the often polluted meat causes, thousands of dolphins are caught in drive hunts each year. In Japan, the hunting is done by a select group of fishermen. When a pod of dolphins has been spotted, they are driven into a bay by the fishermen while banging on metal rods in the water to scare and confuse the dolphins. When the dolphins are in the bay, it is quickly closed off with nets so the dolphins cannot escape. The dolphins are usually not caught and killed immediately, but instead left to calm down over night. The following day, the dolphins are caught one by one and killed. The killing of the animals used to be done by slitting their throats, but the Japanese government banned this method, and now dolphins may officially only be killed by driving a metal pin into the neck of the dolphin, which causes them to die within seconds according to a memo from Senzo Uchida, the executive secretary of the
Japan Cetacean Conference on Zoological Gardens and Aquariums Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. A veterinary team's analysis of a 2011 video footage of Japanese hunters killing
striped dolphin The striped dolphin (''Stenella coeruleoalba'') is an extensively studied dolphin found in temperate and tropical waters of all the world's oceans. It is a member of the oceanic dolphin family (biology), family, Delphinidae. Taxonomy The stripe ...
s using this method suggested that, in one case, death took over four minutes. Since much of the criticism is the result of photos and videos taken during the hunt and slaughter, it is now common for the final capture and slaughter to take place on site inside a tent or under a plastic cover, out of sight from the public. The most circulated footage is probably that of the drive and subsequent capture and slaughter process taken in Futo, Japan, in October 1999, shot by the Japanese animal welfare organization Elsa Nature Conservancy. Part of this footage was, amongst others, shown on
CNN CNN (Cable News Network) is a multinational cable news channel headquartered in Atlanta, Georgia, U.S. Founded in 1980 by American media proprietor Ted Turner and Reese Schonfeld as a 24-hour cable news channel, and presently owned by the M ...
. In recent years, the video has also become widespread on the internet and was featured in the animal welfare documentary ''
Earthlings Earthling or Earthlings may refer to: Film and television * ''Earthling'' (film), a 2010 sci-fi film * ''Earthlings'' (film), a 2005 animal rights documentary * ''The Earthling'', a 1980 drama film * "Earthling" (''Fringe''), a 2009 TV episode ...
'', though the method of killing dolphins as shown in this video is now officially banned. In 2009, a critical documentary on the hunts in Japan titled '' The Cove'' was released and shown amongst others at the
Sundance Film Festival The Sundance Film Festival (formerly Utah/US Film Festival, then US Film and Video Festival) is an annual film festival organized by the Sundance Institute. It is the largest independent film festival in the United States, with more than 46,66 ...
.


Other threats

Toothed whales can also be threatened by humans more indirectly. They are unintentionally caught in fishing nets by commercial fisheries as
bycatch Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
and accidentally swallow fishing hooks.
Gillnetting Gillnetting is a fishing method that uses gillnets: vertical panels of netting that hang from a line with regularly spaced floaters that hold the line on the surface of the water. The floats are sometimes called "corks" and the line with corks is ...
and Seine netting are significant causes of mortality in cetaceans and other
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s. Porpoises are commonly entangled in fishing nets. Whales are also affected by
marine pollution Marine pollution occurs when substances used or spread by humans, such as industrial waste, industrial, agricultural pollution, agricultural and municipal solid waste, residential waste, particle (ecology), particles, noise, excess carbon dioxid ...
. High levels of
organic chemical In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. The st ...
s 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 cancer Gastrointestinal cancer refers to malignant conditions of the gastrointestinal tract (GI tract) and accessory organs of digestion, including the esophagus, stomach, biliary system, pancreas, small intestine, large intestine, rectum and anus. The ...
s and greater vulnerability to infectious diseases. They can also be poisoned by swallowing litter, such as plastic bags. Pollution of the Yangtze river has led to the extinction of the
baiji The baiji (; IPA: ; ''Lipotes vexillifer'', ''Lipotes'' meaning "left behind" and ''vexillifer'' "flag bearer") is a possibly extinct species of freshwater dolphin native to the Yangtze river system in China. It is thought to be the first dolph ...
. Environmentalists speculate that advanced naval
sonar Sonar (sound navigation and ranging or sonic navigation and ranging) is a technique that uses sound propagation (usually underwater, as in submarine navigation) to navigation, navigate, measure distances (ranging), communicate with or detect o ...
endangers some whales. Some scientists suggest that sonar may trigger
whale beaching Cetacean stranding, commonly known as beaching, is a phenomenon in which whales and dolphins strand themselves on land, usually on a beach. Beached whales often die due to dehydration, collapsing under their own weight, or drowning when high tide ...
s, and they point to signs that such whales have experienced
decompression sickness Decompression sickness (abbreviated DCS; also called divers' disease, the bends, aerobullosis, and caisson disease) is a medical condition caused by dissolved gases emerging from solution as bubbles inside the body tissues during decompressio ...
.


Conservation

Currently, no international convention gives universal coverage to all small whales, although the
International Whaling Commission The International Whaling Commission (IWC) is a specialised regional fishery management organisation, established under the terms of the 1946 International Convention for the Regulation of Whaling (ICRW) to "provide for the proper conservation of ...
has attempted to extend its jurisdiction over them.
ASCOBANS Agreement on the Conservation of Small Cetaceans of the Baltic, North East Atlantic, Irish and North Seas, often abbreviated to ASCOBANS, is a regional agreement on the protection of small cetaceans that was concluded as the Agreement on the Con ...
was negotiated to protect all small whales in the North and Baltic Seas and in the northeast Atlantic. ACCOBAMS protects all whales in the Mediterranean and Black Seas. The global
UNEP The United Nations Environment Programme (UNEP) is responsible for coordinating responses to environmental issues within the United Nations system. It was established by Maurice Strong, its first director, after the United Nations Conference on th ...
Convention on Migratory Species The Convention on the Conservation of Migratory Species of Wild Animals, also known as the Convention on Migratory Species (CMS) or the Bonn Convention, is an international agreement that aims to conserve migratory species throughout their r ...
currently covers seven toothed whale species or populations on its Appendix I, and 37 species or populations on Appendix II. All oceanic cetaceans are listed in
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
appendices, meaning international trade in them and products derived from them is very limited. Numerous organisation are dedicated to protecting certain species that do not fall under any international treaty, such as the Committee for the Recovery of the Vaquita, and the Wuhan Institute of Hydrobiology (for the
Yangtze finless porpoise The Yangtze finless porpoise (''Neophocaena asiaeorientalis'') is a species of toothed whale in the family Phocoenidae. It is endemic to the Yangtze River in China, making it the country's only known freshwater cetacean following the possible ext ...
).


In captivity


Species

Various species of toothed whales, mainly dolphins, are kept in captivity, as well as several other species of porpoise such as
harbour porpoise The harbour porpoise (''Phocoena phocoena'') is one of eight extant species of porpoise. It is one of the smallest species of cetacean. As its name implies, it stays close to coastal areas or river estuaries, and as such, is the most familiar ...
s and
finless porpoise ''Neophocaena'' is a genus of porpoise native to the Indian and Pacific oceans, as well as the freshwater habitats of the Yangtze River basin in China. They are commonly known as finless porpoises. Genetic studies indicate that ''Neophocaena'' is ...
s. These small cetaceans are more often than not kept in theme parks, such as
SeaWorld SeaWorld is an American theme park chain with headquarters in Orlando, Florida. It is a proprietor of marine mammal parks, oceanariums, animal theme parks, and rehabilitation centers owned by SeaWorld Parks & Entertainment (one park will be ...
, commonly known as a dolphinarium.
Bottlenose dolphin Bottlenose dolphins are aquatic mammals in the genus ''Tursiops.'' They are common, cosmopolitan members of the family Delphinidae, the family of oceanic dolphins. Molecular studies show the genus definitively contains two species: the common ...
s are the most common species kept in dolphinariums, as they are relatively easy to train, have a long lifespan in captivity, and have a friendly appearance. Hundreds if not thousands of bottlenose dolphins live in captivity across the world, though exact numbers are hard to determine. Orca are well known for their performances in shows, but the number kept in captivity is very small, especially when compared to the number of bottlenose dolphins, with only 44
captives ''Captives'' is a 1994 British romantic crime drama film directed by Angela Pope and written by the Dublin screenwriter Frank Deasy. It stars Julia Ormond, Tim Roth and Keith Allen (actor), Keith Allen. The picture was selected as the opening fil ...
being held in aquaria as of 2012. Other species kept in captivity are spotted dolphins, false killer whales, and
common dolphin The common dolphin (''Delphinus delphis'') is the most abundant cetacean in the world, with a global population of about six million. Despite this fact and its vernacular name, the common dolphin is not thought of as the archetypal dolphin, with ...
s,
Commerson's dolphin Commerson's dolphin (''Cephalorhynchus commersonii''), also referred to by the common names jacobita, skunk dolphin, piebald dolphin, panda dolphin, or tonina overa (in South America), is a small oceanic dolphin of the genus ''Cephalorhynchus'' ...
s, as well as
rough-toothed dolphin The rough-toothed dolphin (''Steno bredanensis'') is a species of dolphin that can be found in deep warm and tropical waters around the world. The species was first described by Georges Cuvier in 1823. The genus name ''Steno'', of which this spe ...
s, but all in much lower numbers than the bottlenose dolphin. Also, fewer than ten
pilot whale Pilot whales are cetaceans belonging to the genus ''Globicephala''. The two extant species are the long-finned pilot whale (''G. melas'') and the short-finned pilot whale (''G. macrorhynchus''). The two are not readily distinguishable at sea, a ...
s,
Amazon river dolphin The Amazon river dolphin (''Inia geoffrensis''), also known as the boto, bufeo or pink river dolphin, is a species of toothed whale classified in the family Iniidae. Three subspecies are currently recognized: ''I. g. geoffrensis'' (Amazon river ...
s,
Risso's dolphin Risso's dolphin (''Grampus griseus'') is a dolphin, the only species of the genus ''Grampus''. Some of the closest related species to these dolphins include: pilot whales (''Globicephala'' spp.), pygmy killer whales (''Feresa attenuata''), melon ...
s,
spinner dolphin The spinner dolphin (''Stenella longirostris'') is a small dolphin found in off-shore tropical waters around the world. It is famous for its acrobatic displays in which it rotates around its longitudinal axis as it leaps through the air. It is a ...
s, or
tucuxi The tucuxi (''Sotalia fluviatilis''), alternatively known in Peru ''bufeo gris'' or ''bufeo negro'', is a species of freshwater dolphin found in the rivers of the Amazon basin. The word ''tucuxi'' is derived from the Tupi language word ''tuchuc ...
are in captivity. Two unusual and very rare
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...
dolphins, known as
wolphins A wholphin (portmanteau of whale + dolphin) is an extremely rare cetacean hybrid born from a mating of a female common bottlenose dolphin (''Tursiops truncatus'') with a male false killer whale (''Pseudorca crassidens''). The name implies a hy ...
, are kept at the Sea Life Park in
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
, which is a cross between a bottlenose dolphin and a false killer whale. Also, two common/bottlenose hybrids reside in captivity: one at
Discovery Cove Discovery Cove is a theme park owned and operated by SeaWorld Parks & Entertainment, and located in Orlando, Florida. It is the sister park of SeaWorld Orlando and Aquatica Orlando. Visitors to the park can interact with a range of marine animals ...
and the other at
SeaWorld San Diego SeaWorld San Diego is an animal theme park, oceanarium, outside aquarium and marine mammal park, in San Diego, California, United States, inside Mission Bay Park. It is owned and operated by SeaWorld Parks & Entertainment. SeaWorld San Diego ...
.


Controversy

Organisations such as
Animal Welfare Institute The Animal Welfare Institute (AWI) is an American non-profit charitable organization founded by Christine Stevens in 1951 with the goal of reducing suffering inflicted on animals by humans. It is one of the oldest animal welfare organizations in ...
and the
Whale and Dolphin Conservation Whale and Dolphin Conservation (WDC), formerly Whale and Dolphin Conservation Society in the UK, is a wildlife charity that is dedicated solely to the worldwide conservation and welfare of all whales, dolphins and porpoises (cetaceans). It has ...
campaign against the captivity of dolphins and orca. SeaWorld faced a lot of criticism after the documentary ''
Blackfish Blackfish is a common name for the following species of fish, dolphins, and whales: Fish * Alaska blackfish, (''Dallia pectoralis''), an Esocidae from Alaska, Siberia and the Bering Sea islands * Black fish (''Carassioides acuminatus'') a cyprin ...
'' was released in 2013. Aggression among captive orca is common. In August 1989, a dominant female orca,
Kandu V Orcas, or killer whales, are large predatory cetaceans that were first captured live and displayed in exhibitions in the 1960s. They soon became popular attractions at public aquariums and aquatic theme parks due to their intelligence, traina ...
, attempted to rake a newcomer whale, Corky II, with her mouth during a live show, and smashed her head into a wall. Kandu V broke her jaw, which severed an artery, and then bled to death. In November 2006, a dominant female killer whale,
Kasatka Kasatka (1976 – August 15, 2017) was a wild caught female orca who lived at SeaWorld San Diego. Life Kasatka was captured off the southeastern coast of Iceland on October 26, 1978 with another young female whale named Kahana. Both were estima ...
, repeatedly dragged experienced trainer Ken Peters to the bottom of the stadium pool during a show after hearing her calf crying for her in the back pools. In February 2010, an experienced female trainer at SeaWorld Orlando, Dawn Brancheau, was killed by orca
Tilikum Tillicum or Tilikum is a word in Chinook Jargon that means people, family, tribe, and relatives, and may refer to: Places * Tilikum Crossing, a bridge in Portland, Oregon * Tillicum, Lakewood, a neighborhood in Lakewood, Washington ** Tillicum st ...
shortly after a show in Shamu Stadium. Tilikum had been associated with the deaths of two people previously. In May 2012,
Occupational Safety and Health Administration The Occupational Safety and Health Administration'' (OSHA ) is a large regulatory agency of the United States Department of Labor that originally had federal visitorial powers to inspect and examine workplaces. Congress established the agenc ...
administrative law judge Ken Welsch cited SeaWorld for two violations in the death of Dawn Brancheau and fined the company a total of US$12,000. Trainers were banned from making close contact with the orca. In April 2014, the
US Court of Appeals for the District of Columbia The United States Court of Appeals for the District of Columbia Circuit (in case citations, D.C. Cir.) is one of the thirteen United States Courts of Appeals. It has the smallest geographical jurisdiction of any of the U.S. federal appellate cou ...
denied an appeal by SeaWorld. In 2013, SeaWorld's treatment of orca in captivity was the basis of the movie ''Blackfish'', which documents the history of Tilikum, an orca captured by SeaLand of the Pacific, later transported to SeaWorld Orlando, which has been involved in the deaths of three people. In the aftermath of the release of the film,
Martina McBride Martina Mariea McBride (née Schiff, born July 29, 1966) is an American country music singer-songwriter and record producer. She is known for her soprano singing range and her country pop material. McBride was born in Sharon, Kansas, and relocat ...
,
38 Special 38 Special may refer to: * .38 Special The .38 Special, also commonly known as .38 S&W Special (not to be confused with .38 S&W), .38 Smith & Wesson Special, .38 Spl, .38 Spc, (pronounced "thirty-eight special"), or 9x29mmR is a rimmed, cent ...
,
REO Speedwagon REO Speedwagon (originally stylized as R.E.O. Speedwagon) is an American rock band from Champaign, Illinois. Formed in 1967, the band cultivated a following during the 1970s and achieved significant commercial success throughout the 1980s. The ...
,
Cheap Trick Cheap Trick is an American rock band from Rockford, Illinois, formed in 1973 by guitarist Rick Nielsen, bassist Tom Petersson, lead vocalist Robin Zander and drummer Bun E. Carlos. The current lineup of the band consists of Zander, Nielsen and ...
,
Heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
,
Trisha Yearwood Patricia Lynn Yearwood (born September 19, 1964) is an American singer, actress, author and television personality. She rose to fame with her 1991 debut single " She's in Love with the Boy," which became a number one hit on the ''Billboard'' c ...
, and
Willie Nelson Willie Hugh Nelson (born April 29, 1933) is an American country musician. The critical success of the album ''Shotgun Willie'' (1973), combined with the critical and commercial success of ''Red Headed Stranger'' (1975) and '' Stardust'' (197 ...
cancelled scheduled concerts at SeaWorld parks. SeaWorld disputes the accuracy of the film, and in December 2013 released an ad countering the allegations and emphasizing its contributions to the study of cetaceans and their conservation.


References


External links

* * {{DEFAULTSORT:Toothed Whale Toothed whales Articles containing video clips Animals that use echolocation Taxa named by William Henry Flower