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Sperm whale

January 01, 1970

"Cachalot" redirects here. For other uses, see Cachalot (disambiguation).
"Kashalot" redirects here. For the Soviet submarine, see Kashalot-class submarine.
For the 2015 film, see Sperm Whale (film).

The sperm whale or cachalot[a] (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 family, along with the pygmy sperm whale and dwarf sperm whale of the genus Kogia.

Sperm whale[1]
Temporal range: Pliocene – Recent[2]
↓
CITES Appendix I (CITES)[4]
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Infraorder: Cetacea
Family: Physeteridae
Genus: Physeter
Species:
P. macrocephalus
Binomial name
Physeter macrocephalus
Major sperm whale grounds
Synonyms
  • Physeter catodon Linnaeus, 1758
  • Physeter microps Linnaeus, 1758
  • Physeter tursio Linnaeus, 1758
  • Physeter australasianus Desmoulins, 1822

The sperm whale is a pelagic mammal with a worldwide range, and will migrate seasonally for feeding and breeding.[5] Females and young males live together in groups, while mature males (bulls) live solitary lives outside of the mating season. The females cooperate to protect and nurse their young. Females give birth every four to twenty years, and care for the calves for more than a decade. A mature sperm whale has no natural predators, although calves and weakened adults are sometimes killed by pods of killer whales (orcas).

Mature males average 16 metres (52 ft) in length, with the head representing up to one-third of the animal's length. Plunging to 2,250 metres (7,382 ft), it is the third deepest diving mammal, exceeded only by the southern elephant seal and Cuvier's beaked whale.[6][7] The sperm whale uses echolocation and vocalization with source level as loud as 236 decibels (re 1 µPa m) underwater.[8][9] It has the largest brain on Earth, more than five times heavier than a human's. Sperm whales can live 70 years or more.[10][11][12]

Sperm whales' heads are filled with a waxy substance called "spermaceti" (sperm oil), from which the whale derives its name. Spermaceti was a prime target of the whaling industry and was sought after for use in oil lamps, lubricants, and candles. Ambergris, a solid waxy waste product sometimes present in its digestive system, is still highly valued as a fixative in perfumes, among other uses. Beachcombers look out for ambergris as flotsam.[13] Sperm whaling was a major industry in the 19th century, depicted in the novel Moby-Dick. The species is protected by the International Whaling Commission moratorium, and is listed as vulnerable by the International Union for Conservation of Nature.

Taxonomy and naming

Etymology

The name "sperm whale" is a clipping of "spermaceti whale". Spermaceti, originally mistakenly identified as the whales' semen, is the semi-liquid, waxy substance found within the whale's head.[14] (See "Spermaceti organ and melon" below.)

The sperm whale is also known as the "cachalot", which is thought to derive from the archaic French for 'tooth' or 'big teeth', as preserved for example in the word caishau in the Gascon dialect (a word of either Romance[15] or Basque[16] origin).

The etymological dictionary of Corominas says the origin is uncertain, but it suggests that it comes from the Vulgar Latin cappula 'sword hilts'.[17] The word cachalot came to English via French from Spanish or Portuguese cachalote, perhaps from Galician/Portuguese cachola 'big head'.[18]

The term is retained in the Russian word for the animal, kashalot (кашалот), as well as in many other languages.[citation needed]

The scientific genus name Physeter comes from the Greek physētēr (φυσητήρ), meaning 'blowpipe, blowhole (of a whale)', or – as a pars pro toto – 'whale'.[citation needed]

The specific name macrocephalus is Latinized from the Greek makroképhalos (μακροκέφαλος 'big-headed'), from makros (μακρός) + kephalē (κεφαλή).[citation needed]

Its synonymous specific name catodon means 'down-tooth', from the Greek elements cat(a)- ('below') and odṓn ('tooth'); so named because it has visible teeth only in its lower jaw.[19] (See "Jaws and teeth" below.)

Another synonym australasianus ('Australasian') was applied to sperm whales in the Southern Hemisphere.[20]

Taxonomy

The sperm whale belongs to the order Cetartiodactyla,[21] the order containing all cetaceans and even-toed ungulates. It is a member of the unranked clade Cetacea, with all the whales, dolphins, and porpoises, and further classified into Odontoceti, containing all the toothed whales and dolphins. It is the sole extant species of its genus, Physeter, in the family Physeteridae. Two species of the related extant genus Kogia, the pygmy sperm whale Kogia breviceps and the dwarf sperm whale K. sima, are placed either in this family or in the family Kogiidae.[22] In some taxonomic schemes the families Kogiidae and Physeteridae are combined as the superfamily Physeteroidea (see the separate entry on the sperm whale family).[23]

Swedish ichthyologist Peter Artedi described it as Physeter catodon in his 1738 work Genera piscium, from the report of a beached specimen in the Orkneys in 1693 and two beached in the Netherlands in 1598 and 1601.[24] The 1598 specimen was near Berkhey.[citation needed]

The sperm whale is one of the species originally described by Carl Linnaeus in his landmark 1758 10th edition of Systema Naturae. He recognised four species in the genus Physeter.[25] Experts soon realised that just one such species exists, although there has been debate about whether this should be named P. catodon or P. macrocephalus, two of the names used by Linnaeus. Both names are still used, although most recent authors now accept macrocephalus as the valid name, limiting catodon's status to a lesser synonym. Until 1974, the species was generally known as P. catodon. In that year, however, Dutch zoologists Antonius M. Husson and Lipke Holthuis proposed that the correct name should be P. macrocephalus, the second name in the genus Physeter published by Linnaeus concurrently with P. catodon.

This proposition was based on the grounds that the names were synonyms published simultaneously, and, therefore, the ICZN Principle of the First Reviser should apply. In this instance, it led to the choice of P. macrocephalus over P. catodon, a view re-stated in Holthuis, 1987.[26] This has been adopted by most subsequent authors, although Schevill (1986[27] and 1987[28]) argued that macrocephalus was published with an inaccurate description and that therefore only the species catodon was valid, rendering the principle of "First Reviser" inapplicable. The most recent version of ITIS has altered its usage from P. catodon to P. macrocephalus,[29] following L. B. Holthuis and more recent (2008) discussions with relevant experts.[30][31] Furthermore, The Taxonomy Committee of the Society for Marine Mammalogy, the largest international association of marine mammal scientists in the world, officially uses Physeter macrocephalus when publishing their definitive list of marine mammal species.[32]

Biology

External appearance

Average sizes[10][33]
Length Weight
Male 16 metres (52 ft) 45 tonnes (50 short tons)
Female 11 metres (36 ft) 15 tonnes (17 short tons)
Newborn 4 metres (13 ft) 1 tonne (1.1 short tons)

The sperm whale is the largest toothed whale and is among the most sexually dimorphic of all cetaceans.[34] Both sexes are about the same size at birth,[10] but mature males are typically 30% to 50% longer and three times as massive as females.[35][36]

Newborn sperm whales are usually between 3.7 and 4.3 meters (12 and 14 ft) long.[37] Female sperm whales are sexually mature at 8 to 9 meters (26 to 30 ft) in length, whilst males are sexually mature at 11 to 12 meters (36 to 39 ft).[38] Female sperm whales are physically mature at about 10.6 to 11 meters (35 to 36 ft) in length and generally do not achieve lengths greater than 12 metres (39 ft).[34][36][38] The largest female sperm whale measured up to 12.3 meters (40 ft) long, and an individual of such size would have weighed about 17 tonnes (19 short tons).[39][40] Male sperm whales are physically mature at about 15 to 16 meters (49 to 52 ft) in length, and larger males can generally achieve 18 to 19 meters (59 to 62 ft).[38][41][34] An 18 meters (59 ft) long male sperm whale is estimated to have weighed 57 tonnes (56 long tons; 63 short tons).[33] By contrast, the second largest toothed whale (Baird's beaked whale) measures up to 12.8 meters (42 ft) and weighs up to 14 tonnes (15 short tons).[42]

There are occasional reports of individual sperm whales achieving even greater lengths, with some historical claims reaching or exceeding 80 feet (24 m). One example is the whale that sank the Essex (one of the incidents behind Moby-Dick), which was claimed to be 85 feet (26 m). However, there is disagreement as to the accuracy of some of these claims, which are often considered exaggerations or as being measured along the curves of the body.[43][34][41]

An individual measuring 20.7 metres (68 ft) was reported from a Soviet whaling fleet near the Kuril Islands in 1950 and is cited by some authors as the largest accurately measured.[34][44] It has been estimated to weigh 80 tonnes (79 long tons; 88 short tons).[43] In a review of size variation in marine megafauna, McClain and colleagues noted that the International Whaling Commission's data contained eight individuals larger than 20.7 metres (68 ft). The authors supported a 24-metre (79 ft) male from the South Pacific in 1933 as the largest recorded. However, sizes like these are rare, with 95% of recorded sperm whales below 15.85 metres (52.0 ft).[34]

In 1853, one sperm whale was reported at 62 feet (19 m) in length, with a head measuring 20 feet (6.1 m).[45] Large lower jawbones are held in the British Natural History Museum and the Oxford University Museum of Natural History, measuring 5 metres (16 ft) and 4.7 metres (15 ft), respectively.[43]

The average size of sperm whales has decreased over the years, probably due to pressure from whaling.[34] Another view holds that exploitation by overwhaling had virtually no effect on the size of the bull sperm whales, and their size may have actually increased in current times on the basis of density dependent effects.[46] Old males taken at Solander Islands were recorded to be extremely large and unusually rich in blubbers.[47]

 
Unusual among cetaceans, the sperm whale's blowhole is highly skewed to the left side of the head.

The sperm whale's unique body is unlikely to be confused with any other species. The sperm whale's distinctive shape comes from its very large, block-shaped head, which can be one-quarter to one-third of the animal's length. The S-shaped blowhole is located very close to the front of the head and shifted to the whale's left.[35] This gives rise to a distinctive bushy, forward-angled spray.[citation needed]

The sperm whale's flukes (tail lobes) are triangular and very thick. Proportionally, they are larger than that of any other cetacean, and are very flexible.[48] The whale lifts its flukes high out of the water as it begins a feeding dive.[35] It has a series of ridges on the back's caudal third instead of a dorsal fin. The largest ridge was called the 'hump' by whalers, and can be mistaken for a dorsal fin because of its shape and size.[10]

In contrast to the smooth skin of most large whales, its back skin is usually wrinkly and has been likened to a prune by whale-watching enthusiasts.[49] Albinos have been reported.[11][50]

Skeleton

 
A sperm whale skeleton

The ribs are bound to the spine by flexible cartilage, which allows the ribcage to collapse rather than snap under high pressure.[51] While sperm whales are well adapted to diving, repeated dives to great depths have long-term effects. Bones show the same avascular necrosis that signals decompression sickness in humans. Older skeletons showed the most extensive damage, whereas calves showed no damage. This damage may indicate that sperm whales are susceptible to decompression sickness, and sudden surfacing could be lethal to them.[52]

Like that of all cetaceans, the spine of the sperm whale has reduced zygapophysial joints, of which the remnants are modified and are positioned higher on the vertebral dorsal spinous process, hugging it laterally, to prevent extensive lateral bending and facilitate more dorso-ventral bending. These evolutionary modifications make the spine more flexible but weaker than the spines of terrestrial vertebrates.[53]

 
Labeled sperm whale skeleton

Like many cetaceans, the sperm whale has a vestigial pelvis that is not connected to the spine.[citation needed]

Like that of other toothed whales, the skull of the sperm whale is asymmetrical so as to aid echolocation. Sound waves that strike the whale from different directions will not be channeled in the same way.[54] Within the basin of the cranium, the openings of the bony narial tubes (from which the nasal passages spring) are skewed towards the left side of the skull.[citation needed]

Jaws and teeth

 
Sperm whale tooth
 
The lower jaw is long and narrow. The teeth fit into sockets along the upper jaw. (lifelike sculpture)

The sperm whale's lower jaw is very narrow and underslung.[55] The sperm whale has 18 to 26 teeth on each side of its lower jaw which fit into sockets in the upper jaw.[55] The teeth are cone-shaped and weigh up to 1 kilogram (2.2 lb) each.[56] The teeth are functional, but do not appear to be necessary for capturing or eating squid, as well-fed animals have been found without teeth or even with deformed jaws. One hypothesis is that the teeth are used in aggression between males.[57] Mature males often show scars which seem to be caused by the teeth[citation needed]. Rudimentary teeth are also present in the upper jaw, but these rarely emerge into the mouth.[58] Analyzing the teeth is the preferred method for determining a whale's age. Like the age-rings in a tree, the teeth build distinct layers of cementum and dentine as they grow.[59]

Brain

 
The sperm whale's brain is the largest in the world, five times heavier than a human brain.

The sperm whale brain is the largest known of any modern or extinct animal, weighing on average about 7.8 kilograms (17 lb)[60][61] (with the smallest known weighing 6.4 kilograms (14 lb) and the largest known weighing 9.2 kilograms (20 lb)),[43][44] more than five times heavier than a human brain, and has a volume of about 8,000 cm3.[62] Although larger brains generally correlate with higher intelligence, it is not the only factor. Elephants and dolphins also have larger brains than humans.[63] The sperm whale has a lower encephalization quotient than many other whale and dolphin species, lower than that of non-human anthropoid apes, and much lower than that of humans.[61][64]

The sperm whale's cerebrum is the largest in all mammalia, both in absolute and relative terms. The olfactory system is reduced, suggesting that the sperm whale has a poor sense of taste and smell. By contrast, the auditory system is enlarged. The pyramidal tract is poorly developed, reflecting the reduction of its limbs.[65]

Biological systems

See also: Physiology of underwater diving § Marine mammals

The sperm whale respiratory system has adapted to cope with drastic pressure changes when diving. The flexible ribcage allows lung collapse, reducing nitrogen intake, and metabolism can decrease to conserve oxygen.[66][67] Between dives, the sperm whale surfaces to breathe for about eight minutes before diving again.[35] Odontoceti (toothed whales) breathe air at the surface through a single, S-shaped blowhole, which is extremely skewed to the left. Sperm whales spout (breathe) 3–5 times per minute at rest, increasing to 6–7 times per minute after a dive. The blow is a noisy, single stream that rises up to 2 metres (6.6 ft) or more above the surface and points forward and left at a 45° angle.[68] On average, females and juveniles blow every 12.5 seconds before dives, while large males blow every 17.5 seconds before dives.[69] A sperm whale killed 160 km (100 mi) south of Durban, South Africa, after a 1-hour, 50-minute dive was found with two dogfish (Scymnodon sp.), usually found at the sea floor, in its belly.[70]

The sperm whale has the longest intestinal system in the world,[71] exceeding 300 m in larger specimens.[72][73] The sperm whale has a four-chambered stomach that is similar to ruminants. The first secretes no gastric juices and has very thick muscular walls to crush the food (since whales cannot chew) and resist the claw and sucker attacks of swallowed squid. The second chamber is larger and is where digestion takes place. Undigested squid beaks accumulate in the second chamber – as many as 18,000 have been found in some dissected specimens.[72][74][75] Most squid beaks are vomited by the whale, but some occasionally make it to the hindgut. Such beaks precipitate the formation of ambergris.[75]

 
The arterial system of a sperm whale foetus

In 1959, the heart of a 22 metric-ton (24 short-ton) male taken by whalers was measured to be 116 kilograms (256 lb), about 0.5% of its total mass.[76] The circulatory system has a number of specific adaptations for the aquatic environment. The diameter of the aortic arch increases as it leaves the heart. This bulbous expansion acts as a windkessel, ensuring a steady blood flow as the heart rate slows during diving.[77] The arteries that leave the aortic arch are positioned symmetrically. There is no costocervical artery. There is no direct connection between the internal carotid artery and the vessels of the brain.[78] Their circulatory system has adapted to dive at great depths, as much as 2,250 metres (7,382 ft)[6][7][79][80][81][excessive citations] for up to 120 minutes.[82] More typical dives are around 400 metres (1,310 ft) and 35 minutes in duration.[35] Myoglobin, which stores oxygen in muscle tissue, is much more abundant than in terrestrial animals.[83] The blood has a high density of red blood cells, which contain oxygen-carrying haemoglobin. The oxygenated blood can be directed towards only the brain and other essential organs when oxygen levels deplete.[84][85][86] The spermaceti organ may also play a role by adjusting buoyancy (see below).[87] The arterial retia mirabilia are extraordinarily well-developed. The complex arterial retia mirabilia of the sperm whale are more extensive and larger than those of any other cetacean.[78]

Senses

Spermaceti organ and melon

 
Anatomy of the sperm whale's head. The organs above the jaw are devoted to sound generation.

Atop the whale's skull is positioned a large complex of organs filled with a liquid mixture of fats and waxes called spermaceti. The purpose of this complex is to generate powerful and focused clicking sounds, the existence of which was proven by Valentine Worthington and William Schevill when a recording was produced on a research vessel in May 1959.[88] The sperm whale uses these sounds for echolocation and communication.[89][90][91][92][93][94][95][96][excessive citations]

The spermaceti organ is like a large barrel of spermaceti. Its surrounding wall, known as the case, is extremely tough and fibrous. The case can hold within it up to 1,900 litres of spermaceti.[97] It is proportionately larger in males.[98] This oil is a mixture of triglycerides and wax esters. The proportion of wax esters in the spermaceti organ increases with the age of the whale: 38–51% in calves, 58–87% in adult females, and 71–94% in adult males.[99] The spermaceti at the core of the organ has a higher wax content than the outer areas.[100] The speed of sound in spermaceti is 2,684 m/s (at 40 kHz, 36 °C), making it nearly twice as fast as in the oil in a dolphin's melon.[101]

Below the spermaceti organ lies the "junk" which consists of compartments of spermaceti separated by cartilage. It is analogous to the melon found in other toothed whales.[102] The structure of the junk redistributes physical stress across the skull and may have evolved to protect the head during ramming.[103][104][102]

Running through the head are two air passages. The left passage runs alongside the spermaceti organ and goes directly to the blowhole, whilst the right passage runs underneath the spermaceti organ and passes air through a pair of phonic lips and into the distal sac at the very front of the nose. The distal sac is connected to the blowhole and the terminus of the left passage. When the whale is submerged, it can close the blowhole, and air that passes through the phonic lips can circulate back to the lungs. The sperm whale, unlike other odontocetes, has only one pair of phonic lips, whereas all other toothed whales have two,[105] and it is located at the front of the nose instead of behind the melon.

At the posterior end of this spermaceti complex is the frontal sac, which covers the concave surface of the cranium. The posterior wall of the frontal sac is covered with fluid-filled knobs, which are about 4–13 mm in diameter and separated by narrow grooves. The anterior wall is smooth. The knobbly surface reflects sound waves that come through the spermaceti organ from the phonic lips. The grooves between the knobs trap a film of air that is consistent whatever the orientation or depth of the whale, making it an excellent sound mirror.[101]

The spermaceti organs may also help adjust the whale's buoyancy. It is hypothesized that before the whale dives, cold water enters the organ, and it is likely that the blood vessels constrict, reducing blood flow, and, hence, temperature. The wax therefore solidifies and reduces in volume.[87][106] The increase in specific density generates a down force of about 392 newtons (88 lbf) and allows the whale to dive with less effort.[citation needed] During the hunt, oxygen consumption, together with blood vessel dilation, produces heat and melts the spermaceti, increasing its buoyancy and enabling easy surfacing.[107] However, more recent work[91] has found many problems with this theory including the lack of anatomical structures for the actual heat exchange.[108] Another issue is that if the spermaceti does indeed cool and solidify, it would affect the whale's echolocation ability just when it needs it to hunt in the depths.[citation needed]

Herman Melville's fictional story Moby-Dick suggests that the "case" containing the spermaceti serves as a battering ram for use in fights between males.[109] A few famous instances include the well-documented sinking of the ships Essex and Ann Alexander by attackers estimated to weigh only one-fifth as much as the ships.[102]

  •  
    The phonic lips.
  •  
    The frontal sac, exposed. Its surface is covered with fluid-filled knobs.
  •  
    A piece of the posterior wall of the frontal sac. The grooves between the knobs trap a consistent film of air, making it an excellent sound mirror.[101]

Eyes and vision

Like other toothed whales, the sperm whale can retract its eyes.

The sperm whale's eye does not differ greatly from those of other toothed whales except in size. It is the largest among the toothed whales, weighing about 170 g. It is overall ellipsoid in shape, compressed along the visual axis, measuring about 7×7×3 cm. The cornea is elliptical and the lens is spherical. The sclera is very hard and thick, roughly 1 cm anteriorly and 3 cm posteriorly. There are no ciliary muscles. The choroid is very thick and contains a fibrous tapetum lucidum. Like other toothed whales, the sperm whale can retract and protrude its eyes, thanks to a 2-cm-thick retractor muscle attached around the eye at the equator,[110] but are unable to roll the eyes in their sockets.[111]

According to Fristrup and Harbison (2002),[112] sperm whale's eyes afford good vision and sensitivity to light. They conjectured that sperm whales use vision to hunt squid, either by detecting silhouettes from below or by detecting bioluminescence. If sperm whales detect silhouettes, Fristrup and Harbison suggested that they hunt upside down, allowing them to use the forward parts of the ventral visual fields for binocular vision.[citation needed]

Sleeping

For some time researchers have been aware that pods of sperm whales may sleep for short periods, assuming a vertical position with their heads just below or at the surface, or head down.[113] A 2008 study published in Current Biology recorded evidence that whales may sleep with both sides of the brain. It appears that some whales may fall into a deep sleep for about 7 percent of the time, most often between 6 p.m. and midnight.[114]

Genetics

Sperm whales have 21 pairs of chromosomes (2n=42).[115] The genome of live whales can be examined by recovering shed skin.[116]

Vocalization complex

Further information on the sonar operator slang term: Carpenter fish
Sperm whale vocalization

After Valentine Worthington and William E. Schevill confirmed the existence of sperm whale vocalization,[88] further studies found that sperm whales are capable of emitting sounds at a volume of 230 decibels – more than an aircraft jet engine at takeoff – making the sperm whale the loudest animal in the world.[117]

Mechanism

When echolocating, the sperm whale emits a directionally focused beam of broadband clicks. Clicks are generated by forcing air through a pair of phonic lips (also known as "monkey lips" or "museau de singe") at the front end of the nose, just below the blowhole. The sound then travels backwards along the length of the nose through the spermaceti organ. Most of the sound energy is then reflected off the frontal sac at the cranium and into the melon, whose lens-like structure focuses it.[89] Some of the sound will reflect back into the spermaceti organ and back towards the front of the whale's nose, where it will be reflected through the spermaceti organ a third time. This back and forth reflection which happens on the scale of a few milliseconds creates a multi-pulse click structure.[118]

This multi-pulse click structure allows researchers to measure the whale's spermaceti organ using only the sound of its clicks.[119][120] Because the interval between pulses of a sperm whale's click is related to the length of the sound producing organ, an individual whale's click is unique to that individual. However, if the whale matures and the size of the spermaceti organ increases, the tone of the whale's click will also change.[120] The lower jaw is the primary reception path for the echoes. A continuous fat-filled canal transmits received sounds to the inner ear.[121]

The source of the air forced through the phonic lips is the right nasal passage. While the left nasal passage opens to the blow hole, the right nasal passage has evolved to supply air to the phonic lips. It is thought that the nostrils of the land-based ancestor of the sperm whale migrated through evolution to their current functions, the left nostril becoming the blowhole and the right nostril becoming the phonic lips.[122]

Air that passes through the phonic lips passes into the distal sac, then back down through the left nasal passage. This recycling of air allows the whale to continuously generate clicks for as long as it is submerged.[123]

Vocalization types

The sperm whale's vocalizations are all based on clicking, described in four types: the usual echolocation, creaks, codas, and slow clicks.[124]

The usual echolocation click type is used in searching for prey.[124] 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.[124]

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.[125]

Characteristics of sperm whale clicks[124]
Click type Apparent
source level
(dB re 1 µPa m) 		Directionality Centroid
frequency
(kHz) 		Inter-click
interval
(s) 		Duration
of click
(ms) 		Duration
of pulse
(ms) 		Range audible
to sperm whale
(km) 		Inferred function Audio sample
Usual 230 High 15 0.5–1.0 15–30 0.1 16 Searching for prey
Creak 205 High 15 0.005–0.1 0.1–5 0.1 6 Homing in on prey
Coda 180 Low 5 0.1–0.5 35 0.5 ~2 Social communication
Slow 190 Low 0.5 5–8 30 5 60 Communication by males

Codas

The most distinctive vocalizations are codas, which are short rhythmic sequences of clicks, mostly numbering 3–12 clicks, in stereotyped patterns.[126] They are classified using variations in the number of clicks, rhythm, and tempo.[127]

Codas are the result of vocal learning within a stable social group,[128] and are made in the context of the whales' social unit.[126] “The foundation of sperm whale society is the matrilineally based social unit of ten or so females and their offspring. The members of the unit travel together, suckle each others' infants, and babysit them while mothers make long deep dives to feed.”[126] Over 70% of a sperm whale's time is spent independently foraging; codas “could help whales reunite and reaffirm their social ties in between long foraging dives.”[127][129]

While nonidentity codas are commonly used in multiple different clans,[130] some codas express clan identity, and denote different patterns of travel, foraging, and socializing or avoidance among clans.[129][131] In particular, whales will not group with whales of another clan even though they share the same geographical area.[126] Statistically, as the clans' ranges become more overlapped, the distinction in clan identity coda usage becomes more pronounced.[130] Distinctive codas identify seven clans described among the approximately 150,000 female sperm whales in the Pacific Ocean, and there are another four clans in the Atlantic.[126] As “arbitrary traits that function as reliable indicators of cultural group membership,” clan identity codas act as symbolic markers that modulate interactions between individuals.[127]

Individual identity in sperm whale vocalizations is an ongoing scientific issue, however. A distinction needs to be made between cues and signals. Human acoustic tools can distinguish individual whales by analyzing micro-characteristics of their vocalizations,[132] and the whales can probably do the same. This does not prove that the whales deliberately use some vocalizations to signal individual identity in the manner of the signature whistles that bottlenose dolphins use as individual labels.[128][127]

Ecology

Distribution

 
Global concentrations of sperm whales

Sperm whales are among the most cosmopolitan species. They prefer ice-free waters over 1,000 metres (3,300 ft) deep.[3] Although both sexes range through temperate and tropical oceans and seas, only adult males populate higher latitudes.[11] Among several regions, such as along coastal waters of southern Australia, sperm whales have been considered to be locally extinct.[133]

They are relatively abundant from the poles to the equator and are found in all the oceans. They inhabit the Mediterranean Sea, but not the Black Sea,[10] while their presence in the Red Sea is uncertain.[3] The shallow entrances to both the Black Sea and the Red Sea may account for their absence.[134] The Black Sea's lower layers are also anoxic and contain high concentrations of sulphur compounds such as hydrogen sulphide.[135][What does being anoxic have to do with whales that don't breath under water? Lack of prey?] The first ever sighting off the coast of Pakistan was made in 2017.[136][137] The first ever record off the west coast of the Korean Peninsula (Yellow Sea) was made in 2005.[138][139] followed by one near Ganghwa Island in 2009.[140]

Populations are denser close to continental shelves and canyons.[11] Sperm whales are usually found in deep, off-shore waters, but may be seen closer to shore, in areas where the continental shelf is small and drops quickly to depths of 310 to 920 metres (1,020 to 3,020 ft).[10] Coastal areas with significant sperm whale populations include the Azores and Dominica.[141] In east Asian waters, whales are also observed regularly in coastal waters in places such as the Commander and Kuril Islands, Shiretoko Peninsula which is one of few locations where sperm whales can be observed from shores,[142] off Kinkasan, vicinity to Tokyo Bay[143] and the Bōsō Peninsula to the Izu[144][145] and the Izu Islands, the Volcano Islands, Yakushima and the Tokara Islands to the Ryukyu Islands,[146][147] Taiwan, the Northern Mariana Islands,[148] and so forth. Historical catch records suggest there could have been smaller aggression grounds in the Sea of Japan as well.[149] Along the Korean Peninsula, the first confirmed observation within the Sea of Japan, eight animals off Guryongpo, was made in 2004 since after the last catches of five whales off Ulsan in 1911,[139][150] while nine whales were observed in the East China Sea side of the peninsula in 1999.[151]

Grown males are known to enter surprisingly shallow bays to rest (whales will be in a state of rest during these occasions). Unique, coastal groups have been reported from various areas around the globe, such as near Scotland's coastal waters,[152] and the Shiretoko Peninsula, off Kaikōura, in Davao Gulf. Such coastal groups were more abundant in pre-whaling days.[153]

Genetic analysis indicates that the world population of sperm whales originated in the Pacific Ocean from a population of about 10,000 animals around 100,000 years ago, when expanding ice caps blocked off their access to other seas. In particular, colonization of the Atlantic was revealed to have occurred multiple times during this expansion of their range.[154]

Diet

 
A piece of sperm whale skin with giant squid sucker scars

Sperm whales usually dive between 300 and 800 metres (980 and 2,620 ft), and sometimes 1 to 2 kilometres (3,300 to 6,600 ft), in search of food.[155] Such dives can last more than an hour.[155] They feed on several species, notably the giant squid, but also the colossal squid, octopuses, and fish such as demersal rays and sharks,[citation needed] but their diet is mainly medium-sized squid.[156] Some prey may be taken accidentally while eating other items.[156] Most of what is known about deep-sea squid has been learned from specimens in captured sperm whale stomachs, although more recent studies analysed faeces.

One study, carried out around the Galápagos, found that squid from the genera Histioteuthis (62%), Ancistrocheirus (16%), and Octopoteuthis (7%) weighing between 12 and 650 grams (0.026 and 1.433 lb) were the most commonly taken.[157] 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. One study published in 2010 collected evidence that suggests that female sperm whales may collaborate when hunting Humboldt squid.[158] Tagging studies have shown that sperm whales hunt upside down at the bottom of their deep dives. It is suggested that the whales can see the squid silhouetted above them against the dim surface light.[159]

An older study, examining whales captured by the New Zealand whaling fleet in the Cook Strait region, found a 1.69:1 ratio of squid to fish by weight.[160] Sperm whales sometimes take sablefish and toothfish from long lines. Long-line fishing operations in the Gulf of Alaska complain that sperm whales take advantage of their fishing operations to eat desirable species straight off the line, sparing the whales the need to hunt.[161] However, the amount of fish taken is very little compared to what the sperm whale needs per day. Video footage has been captured of a large male sperm whale "bouncing" a long line, to gain the fish.[162] Sperm whales are believed to prey on the megamouth shark, a rare and large deep-sea species discovered in the 1970s.[163] In one case, three sperm whales were observed attacking or playing with a megamouth.[164]

Sperm whales have also been noted to feed on bioluminescent pyrosomes such as Pyrosoma atlanticum.[165][166][167] It is thought that the foraging strategy of sperm whales for bioluminescent squids may also explain the presence of these light-emitting pyrosomes in the diet of the sperm whale.[167]

 
Ambergris

The sharp beak of a consumed squid lodged in the whale's intestine may lead to the production of ambergris, analogous to the production of pearls in oysters.[168] The irritation of the intestines caused by squid beaks stimulates the secretion of this lubricant-like substance. Sperm whales are prodigious feeders and eat around 3% of their body weight per day. The total annual consumption of prey by sperm whales worldwide is estimated to be about 272 million tonnes (300 million short tons).[169][170] In comparison, human consumption of seafood is estimated to be 157 million tonnes (173 million short tons).[171]

Sperm whales hunt through echolocation. Their clicks are among the most powerful sounds in the animal kingdom (see above). It has been hypothesised that it can stun prey with its clicks. Experimental studies attempting to duplicate this effect have been unable to replicate the supposed injuries, casting doubt on this idea.[172] One study showing that sound pressure levels on the squid are more than an order of magnitude below levels required for debilitation, and therefore, precluding acoustic stunning to facilitate prey capture.[173]

Sperm whales, as well as other large cetaceans, help fertilise the surface of the ocean by consuming nutrients in the depths and transporting those nutrients to the oceans' surface when they defecate, an effect known as the whale pump.[174] This fertilises phytoplankton and other plants on the surface of the ocean and contributes to ocean productivity and the drawdown of atmospheric carbon.[175]

Life cycle

See also: Whale reproduction

Sperm whales can live 70 years or more.[10][11][12] They are a prime example of a species that has been K-selected, meaning their reproductive strategy is associated with stable environmental conditions and comprises a low birth rate, significant parental aid to offspring, slow maturation, and high longevity.[35]

How they choose mates has not been definitively determined. Bulls will fight with each other over females, and males will mate with multiple females, making them polygynous, but they do not dominate the group as in a harem.[176][177] Bulls do not provide paternal care to their offspring but rather play a fatherly role to younger bulls to show dominance.[178]

Females become fertile at around 9 years of age.[179] The oldest pregnant female ever recorded was 41 years old.[180] Gestation requires 14 to 16 months, producing a single calf.[10] Sexually mature females give birth once every 4 to 20 years (pregnancy rates were higher during the whaling era).[179] Birth is a social event, as the mother and calf need others to protect them from predators. The other adults may jostle and bite the newborn in its first hours.[181]

Lactation proceeds for 19 to 42 months, but calves, rarely, may suckle up to 13 years.[10] Like that of other whales, the sperm whale's milk has a higher fat content than that of terrestrial mammals: about 36%,[182] compared to 4% in cow milk. This gives it a consistency similar to cottage cheese,[183] which prevents it from dissolving in the water before the calf can drink it.[184] It has an energy content of roughly 3,840 kcal/kg,[182] compared to just 640 kcal/kg in cow milk.[185] Calves may be allowed to suckle from females other than their mothers.[10]

Males become sexually mature at 18 years. Upon reaching sexual maturity, males move to higher latitudes, where the water is colder and feeding is more productive. Females remain at lower latitudes.[10] Males reach their full size at about age 50.[35]

Social behaviour

Relations within the species

 
Sperm whales adopt the "marguerite formation" to defend a vulnerable pod member.

Like elephants, females and their young live in matriarchal groups called pods, while bulls live apart. Bulls sometimes form loose bachelor groups with other males of similar age and size. As they grow older, they typically live solitary lives, only returning to the pod to socialize or to breed.[35] Bulls have beached themselves together, suggesting a degree of cooperation which is not yet fully understood.[35] The whales rarely, if ever, leave their group.[186]

A social unit is a group of sperm whales who live and travel together over a period of years. Individuals rarely, if ever, join or leave a social unit. There is a huge variance in the size of social units. They are most commonly between six and nine individuals in size but can have more than twenty.[187] Unlike orcas, sperm whales within a social unit show no significant tendency to associate with their genetic relatives.[188] Females and calves spend about three-quarters of their time foraging and a quarter of their time socializing. Socializing usually takes place in the afternoon.[189]

When sperm whales socialize, they emit complex patterns of clicks called codas. They will spend much of the time rubbing against each other. Tracking of diving whales suggests that groups engage in herding of prey, similar to bait balls created by other species, though the research needs to be confirmed by tracking the prey.[190][191]

Relations with other species

The most common natural predator of sperm whales is the orca (killer whale), but pilot whales and false killer whales sometimes harass them.[192][193] Orcas prey on target groups of females with young, usually making an effort to extract and kill a calf. The females will protect their calves or an injured adult by encircling them. They may face inwards with their tails out (the 'marguerite formation', named after the flower). The heavy and powerful tail of an adult whale is potentially capable of delivering lethal blows.[194] Alternatively, they may face outwards (the 'heads-out formation'). Other than sperm whales, southern right whales had been observed to perform similar formations.[195] However, formations in non-dangerous situations have been recorded as well.[195] Early whalers exploited this behaviour, attracting a whole unit by injuring one of its members.[196] Such a tactic is described in Moby-Dick:

"Say you strike a Forty-barrel-bull—poor devil! all his comrades quit him. But strike a member of the harem school, and her companions swim around her with every token of concern, sometimes lingering so near her and so long, as themselves to fall a prey."[197]

If the killer whale pod is large, its members may sometimes be able to kill adult female sperm whales and can at least injure an entire pod of sperm whales.[198][199] Bulls have no predators, and are believed to be too large, powerful and aggressive to be threatened by killer whales.[200] Solitary bulls are known to interfere and come to the aid of vulnerable groups nearby.[201] However, the bull sperm whale, when accompanying pods of female sperm whales and their calves as such, may be reportedly unable to effectively dissuade killer whales from their attacks on the group, although the killer whales may end the attack sooner when a bull is present.[202][203]

However, male sperm whales have been observed to attack and intimidate killer whale pods in competitive feeding instances. An incident was filmed from a long-line trawler: a killer whale pod was systematically taking fish caught on the trawler's long lines (as the lines were being pulled into the ship) when a male sperm whale appeared to repeatedly charge the killer whale pod in an attempt to drive them away; it was speculated by the film crew that the sperm whale was attempting to access the same fish. The killer whales employed a tail outward and tail-slapping defensive position against the bull sperm whale similar to that used by female sperm whales against attacking killer whales.[204] However, at some potential feeding sites, the killer whales may prevail over sperm whales even when outnumbered by the sperm whales. Some authors consider the killer whales "usually" behaviorally dominant over sperm whales but express that the two species are "fairly evenly matched", with the killer whales' greater aggression, more considerable biting force for their size and predatory prowess more than compensating for their smaller size.[202][205]

Sperm whales are not known for forging bonds with other species, but it was observed that a bottlenose dolphin with a spinal deformity had been accepted into a pod of sperm whales.[206] They are known to swim alongside other cetaceans such as humpback,[207] fin, minke, pilot,[208] and killer whales on occasion.[209]

Parasites

Sperm whales can suffer from parasites. Out of 35 sperm whales caught during the 1976–1977 Antarctic whaling season, all of them were infected by Anisakis physeteris (in their stomachs) and Phyllobothrium delphini (in their blubber). Both whales with a placenta were infected with Placentonema gigantissima,[210] potentially the largest nematode worm ever described.[citation needed]

Evolutionary history

See also: Sperm whale family
Evolutionary family tree of sperm whales,[211]
including simplified summary of extinct groups (†)[212]

Fossil record

Although the fossil record is poor,[213] several extinct genera have been assigned to the clade Physeteroidea, which includes the last common ancestor of the modern sperm whale, pygmy sperm whales, dwarf sperm whales, and extinct physeteroids. These fossils include Ferecetotherium, Idiorophus, Diaphorocetus, Aulophyseter, Orycterocetus, Scaldicetus, Placoziphius, Zygophyseter and Acrophyseter.[23][212][214] Ferecetotherium, found in Azerbaijan and dated to the late Oligocene (about 28 to 23 million years ago), is the most primitive fossil that has been found, which possesses sperm whale-specific features, such as an asymmetric rostrum ("beak" or "snout").[215] Most sperm whale fossils date from the Miocene period, 23 to 5 million years ago. Diaphorocetus, from Argentina, has been dated to the early Miocene. Fossil sperm whales from the Middle Miocene include Aulophyseter, Idiorophus and Orycterocetus, all of which were found on the West Coast of the United States, and Scaldicetus, found in Europe and Japan.[215][216] Orycterocetus fossils have also been found in the North Atlantic Ocean and the Mediterranean Sea, in addition to the west coast of the United States.[217] Placoziphius, found in Europe, and Acrophyseter, from Peru, are dated to the late Miocene.[23][215]

Fossil sperm whales differ from modern sperm whales in tooth count and the shape of the face and jaws.[215] For example, Scaldicetus had a tapered rostrum.[216] Genera from the Oligocene and early and middle Miocene, with the possible exception of Aulophyseter, had teeth in their upper jaws.[215] Acrophyseter, from the late Miocene, also had teeth in both the upper and lower jaws as well as a short rostrum and an upward curving mandible (lower jaw).[23] These anatomical differences suggest that fossil species may not have necessarily been deep-sea squid eaters such as the modern sperm whale, but that some genera mainly ate fish.[215] Zygophyseter, dated from the middle to late Miocene and found in southern Italy, had teeth in both jaws and appears to have been adapted to feed on large prey, rather like the modern killer whale (orca). Other fossil sperm whales with adaptations similar to this are collectively known as killer sperm whales.[212]

Two poorly known fossil species belonging to the modern genus Physeter have been recognized so far: P. antiquus (Neogene of France)[218] and P. vetus (Neogene of eastern North America).[219] Physeter vetus is very likely an invalid species, as the few teeth that were used to identify this species appear to be identical to those of another toothed whale, Orycterocetus quadratidens.[220]

Phylogeny

The traditional view has been that Mysticeti (baleen whales) and Odontoceti (toothed whales) arose from more primitive whales early in the Oligocene period, and that the super-family Physeteroidea, which contains the sperm whale, dwarf sperm whale, and pygmy sperm whale, diverged from other toothed whales soon after that, over 23 million years ago.[213][215] From 1993 to 1996, molecular phylogenetics analyses by Milinkovitch and colleagues, based on comparing the genes of various modern whales, suggested that the sperm whales are more closely related to the baleen whales than they are to other toothed whales, which would have meant that Odontoceti were not monophyletic; in other words, it did not consist of a single ancestral toothed whale species and all its descendants.[211] However, more recent studies, based on various combinations of comparative anatomy and molecular phylogenetics, criticised Milinkovitch's analysis on technical grounds and reaffirmed that the Odontoceti are monophyletic.[211][221][222]

These analyses also confirm that there was a rapid evolutionary radiation (diversification) of the Physeteroidea in the Miocene period.[212] The Kogiidae (dwarf and pygmy sperm whales) diverged from the Physeteridae (true sperm whales) at least 8 million years ago.[221]

Relationship with humans

Sperm whaling

Main articles: Whaling and Sperm whaling
 
In the 19th century, sperm whales were hunted using rowboats and hand-thrown harpoons, a rather dangerous method, as the whales sometimes fought back.

Spermaceti, obtained primarily from the spermaceti organ, and sperm oil, obtained primarily from the blubber in the body, were much sought after by 18th, 19th, and 20th century whalers. These substances found a variety of commercial applications, such as candles, soap, cosmetics, machine oil, other specialised lubricants, lamp oil, pencils, crayons, leather waterproofing, rust-proofing materials and many pharmaceutical compounds.[223][224][225][226] Ambergris, a highly expensive, solid, waxy, flammable substance produced in the digestive system of sperm whales, was also sought as a fixative in perfumery.[citation needed]

Prior to the early eighteenth century, hunting was mostly by indigenous Indonesians.[227] Legend has it that sometime in the early 18th century, around 1712, Captain Christopher Hussey, while cruising for right whales near shore, was blown offshore by a northerly wind, where he encountered a sperm whale pod and killed one.[228] Although the story may not be true, sperm whales were indeed soon exploited by American whalers. Judge Paul Dudley, in his Essay upon the Natural History of Whales (1725), states that a certain Atkins, 10 or 12 years in the trade, was among the first to catch sperm whales sometime around 1720 off the New England coast.[229]

There were only a few recorded instances during the first few decades (1709–1730s) of offshore sperm whaling. Instead, sloops concentrated on the Nantucket Shoals, where they would have taken right whales or went to the Davis Strait region to catch bowhead whales. By the early 1740s, with the advent of spermaceti candles (before 1743), American vessels began to focus on sperm whales. The diary of Benjamin Bangs (1721–1769) shows that, along with the bumpkin sloop he sailed, he found three other sloops flensing sperm whales off the coast of North Carolina in late May 1743.[230] On returning to Nantucket in the summer 1744 on a subsequent voyage, he noted that "45 spermacetes are brought in here this day," another indication that American sperm whaling was in full swing.[230]

American sperm whaling soon spread from the east coast of the American colonies to the Gulf Stream, the Grand Banks, West Africa (1763), the Azores (1765), and the South Atlantic (1770s). From 1770 to 1775 Massachusetts, New York, Connecticut, and Rhode Island ports produced 45,000 barrels of sperm oil annually, compared to 8,500 of whale oil.[231] In the same decade, the British began sperm whaling, employing American ships and personnel.[232] By the following decade, the French had entered the trade, also employing American expertise.[232] Sperm whaling increased until the mid-nineteenth century. Spermaceti oil was important in public lighting (for example, in lighthouses, where it was used in the United States until 1862, when it was replaced by lard oil, in turn replaced by petroleum) and for lubricating the machines (such as those used in cotton mills) of the Industrial Revolution. Sperm whaling declined in the second half of the nineteenth century, as petroleum came into broader use. In that sense, petroleum use may be said to have protected whale populations from even greater exploitation.[233][234] Sperm whaling in the 18th century began with small sloops carrying only one or two whaleboats. The fleet's scope and size increased over time, and larger ships entered the fishery. In the late 18th century and early 19th century, sperm whaling ships sailed to the equatorial Pacific, the Indian Ocean, Japan, the coast of Arabia, Australia and New Zealand.[232][235][236] Hunting could be dangerous to the crew, since sperm whales (especially bulls) will readily fight to defend themselves against attack, unlike most baleen whales. When dealing with a threat, sperm whales will use their huge head effectively as a battering ram.[102] Arguably the most famous sperm whale counter-attack occurred on 20 November 1820, when a whale claimed to be about 25.9 metres (85 ft) long rammed and sank the Nantucket whaleship Essex. Only 8 out of 21 sailors survived to be rescued by other ships.[237]

 
Scrimshaw was the art of engraving on the teeth of sperm whales. It was a way for whalers to pass the time between hunts.

The sperm whale's ivory-like teeth were often sought by 18th- and 19th-century whalers, who used them to produce inked carvings known as scrimshaw. 30 teeth of the sperm whale can be used for ivory. Each of these teeth, up to 20 cm (8 in) and 8 cm (3 in) across, are hollow for the first half of their length. Like walrus ivory, sperm whale ivory has two distinct layers. However, sperm whale ivory contains a much thicker inner layer. Though a widely practised art in the 19th century, scrimshaw using genuine sperm whale ivory declined substantially after the retirement of the whaling fleets in the 1880s.[citation needed]

Modern whaling was more efficient than open-boat whaling, employing steam-powered ships and exploding harpoons. Initially, modern whaling activity focused on large baleen whales, but as these populations were taken, sperm whaling increased. Spermaceti, the fine waxy oil produced by sperm whales, was in high demand. In both the 1941–1942 and 1942–1943 seasons, Norwegian expeditions took over 3,000 sperm whales off the coast of Peru alone. After World War II, whaling continued unabated to obtain oil for cosmetics and high-performance machinery, such as automobile transmissions.[citation needed]

The hunting led to the near-extinction of large whales, including sperm whales, until bans on whale oil use were instituted in 1972. The International Whaling Commission gave the species full protection in 1985, but hunting by Japan in the northern Pacific Ocean continued until 1988.[234]

It is estimated that the historic worldwide population numbered 1,100,000 before commercial sperm whaling began in the early 18th century.[3] By 1880, it had declined by an estimated 29 percent.[3] From that date until 1946, the population appears to have partially recovered as whaling activity decreased, but after the Second World War, the population declined even further, to 33 per cent of the pre-whaling population.[citation needed] Between 184,000 and 236,000 sperm whales were killed by the various whaling nations in the 19th century,[238] while in the 20th century, at least 770,000 were taken, the majority between 1946 and 1980.[239]

 
Sperm whaling peaked in the 1830s and 1960s.

Sperm whales increase levels of primary production and carbon export by depositing iron-rich faeces into surface waters of the Southern Ocean. The iron-rich faeces cause phytoplankton to grow and take up more carbon from the atmosphere. When the phytoplankton dies, it sinks to the deep ocean and takes the atmospheric carbon with it. By reducing the abundance of sperm whales in the Southern Ocean, whaling has resulted in an extra 2 million tonnes of carbon remaining in the atmosphere each year.[240]

Remaining sperm whale populations are large enough that the species' conservation status is rated as vulnerable rather than endangered.[3] However, the recovery from centuries of commercial whaling is a slow process, particularly in the South Pacific, where the toll on breeding-age males was severe.[241]

Current conservation status

The total number of sperm whales in the world is unknown, but is thought to be in the hundreds of thousands.[3] The conservation outlook is brighter than for many other whales. Commercial whaling has ceased,[3] and the species is protected almost worldwide, though records indicate that in the 11-year period starting from 2000, Japanese vessels have caught 51 sperm whales.[needs update] Fishermen do not target sperm whales to eat,[3] but long-line fishing operations in the Gulf of Alaska have complained about sperm whales stealing fish from their lines.[161]

Currently, entanglement in fishing nets and collisions with ships represent the greatest threats to the sperm whale population.[11] Other threats include ingestion of marine debris, ocean noise, and chemical pollution.[242] The International Union for Conservation of Nature (IUCN) regards the sperm whale as being "vulnerable".[3] The species is listed as endangered on the United States Endangered Species Act.[243]

Sperm whales are listed on Appendix I[244] and Appendix II[244] of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). It is listed on Appendix I[244] as this species has been categorized as being in danger of extinction throughout all or a significant proportion of their range and CMS Parties strive towards strictly protecting these animals, conserving or restoring the places where they live, mitigating obstacles to migration and controlling other factors that might endanger them. It is listed on Appendix II[244] as it has an unfavourable conservation status or would benefit significantly from international co-operation organised by tailored agreements. It is also covered by the Agreement on the Conservation of Cetaceans in the Black Sea, Mediterranean Sea and Contiguous Atlantic Area (ACCOBAMS) and the Memorandum of Understanding for the Conservation of Cetaceans and Their Habitats in the Pacific Islands Region (Pacific Cetaceans MOU).[citation needed]

The species is protected under Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). This makes commercial international trade (including in parts and derivatives) prohibited, with all other international trade strictly regulated through a system of permits and certificates.[4]

Cultural importance

 
Sperm whale teeth necklace from Fiji

Rope-mounted teeth are important cultural objects throughout the Pacific. In New Zealand, the Māori know them as "rei puta"; such whale tooth pendants were rare objects because sperm whales were not actively hunted in traditional Māori society.[245] Whale ivory and bone were taken from beached whales. In Fiji the teeth are known as tabua, traditionally given as gifts for atonement or esteem (called sevusevu), and were important in negotiations between rival chiefs.[246] Friedrich Ratzel in The History of Mankind reported in 1896 that, in Fiji, whales' or cachalots' teeth were the most-demanded article of ornament or value. They occurred often in necklaces.[247] Today the tabua remains an important item in Fijian life. The teeth were originally rare in Fiji and Tonga, which exported teeth, but with the Europeans' arrival, teeth flooded the market and this "currency" collapsed. The oversupply led in turn to the development of the European art of scrimshaw.[248]

Herman Melville's novel Moby-Dick is based on a true story about a sperm whale that attacked and sank the whaleship Essex.[249][250] Melville associated the sperm whale with the Bible's Leviathan.[250][251] The fearsome reputation perpetuated by Melville was based on bull whales' ability to fiercely defend themselves from attacks by early whalers, smashing whaling boats and, occasionally, attacking and destroying whaling ships.[citation needed]

In Jules Verne's Twenty Thousand Leagues Under the Sea, the Nautilus fights a group of "cachalots" (sperm whales) to protect a pod of southern right whales from their attacks. Verne portrays them as being savage hunters ("nothing but mouth and teeth").[citation needed]

The sperm whale was designated as the Connecticut state animal by the General Assembly in 1975.[252] It was selected because of its specific contribution to the state's history and because of its present-day plight as an endangered species.[253]

Watching sperm whales

See also: Whale watching

Sperm whales are not the easiest of whales to watch, due to their long dive times and ability to travel long distances underwater. However, due to the distinctive look and large size of the whale, watching is increasingly popular.[citation needed] Sperm whale watchers often use hydrophones to listen to the clicks of the whales and locate them before they surface.[254] Popular locations for sperm whale watching include the town of Kaikōura on New Zealand's South Island, Andenes and Tromsø in Arctic Norway; as well as the Azores, where the continental shelf is so narrow that whales can be observed from the shore,[141][255] and Dominica[256] where a long-term scientific research program, The Dominica Sperm Whale Project, has been in operation since 2005.[257]

Plastic waste

The introduction of plastic waste to the ocean environment by humans is relatively new. From the 1970s, sperm whales have occasionally been found with pieces of plastic in their stomachs.[167][258][259][260]

See also

Notes

  1. ^ /ˈkæʃəlɒt, ˈkæʃəloʊ/ – "cachalot". Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)

References

  1. ^ Mead, J. G.; Brownell, R. L. Jr. (2005). "Order Cetacea". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 737. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ^ "Physeter macrocephalus Linnaeus 1758 (sperm whale)". Fossilworks: Gateway to the Paleobiology Database. Retrieved 17 December 2021.
  3. ^ a b c d e f g h i j Taylor, B.L.; Baird, R.; Barlow, J.; Dawson, S.M.; Ford, J.; Mead, J.G.; Notarbartolo di Sciara, G.; Wade, P.; Pitman, R.L. (2019). "Physeter macrocephalus". IUCN Red List of Threatened Species. 2019: e.T41755A160983555. doi:10.2305/IUCN.UK.2008.RLTS.T41755A160983555.en. Retrieved 19 November 2021.
  4. ^ a b "Appendices | CITES". cites.org. Retrieved 14 January 2022.
  5. ^ . acsonline.org. Archived from the original on 22 April 2017. Retrieved 13 May 2017.
  6. ^ a b Gregory S. Schorr; Erin A. Falcone; David J. Moretti; Russel D. Andrews (2014). "First long-term behavioral records from Cuvier's beaked whales (Ziphius cavirostris) reveal record-breaking dives". PLOS One. 9 (3): e92633. Bibcode:2014PLoSO...992633S. doi:10.1371/journal.pone.0092633. PMC 3966784. PMID 24670984.
  7. ^ a b "Census of Marine Life – From the Edge of Darkness to the Black Abyss" (PDF). Coml.org. Retrieved 15 December 2009.
  8. ^ Møhl, Bertel; Wahlberg, Magnus; Peter T. Madsen (2003). "The monopulsed nature of sperm whale clicks". The Journal of the Acoustical Society of America. 114 (2): 1143–1154. Bibcode:2003ASAJ..114.1143M. doi:10.1121/1.1586258. PMID 12942991.
  9. ^ Trivedi, Bijal P. (3 November 2003). . National Geographic. Archived from the original on 6 November 2003.
  10. ^ a b c d e f g h i j k Shirihai, H. & Jarrett, B. (2006). Whales, Dolphins, and Other Marine Mammals of the World. Princeton: Princeton Univ. Press. pp. 21–24. ISBN 978-0-691-12757-6.
  11. ^ a b c d e f Reeves, R.; Stewart, B.; Clapham, P. & Powell, J. (2003). Guide to Marine Mammals of the World. New York: A.A. Knopf. pp. 240–243. ISBN 978-0-375-41141-0.
  12. ^ a b Whitehead, H. & Weilgart, L. (2000). "The Sperm Whale". In Mann, J.; Connor, R.; Tyack, P. & Whitehead, H. (eds.). Cetacean Societies. The University of Chicago Press. p. 169. ISBN 978-0-226-50341-7.
  13. ^ Spitznagel, Eric (12 January 2012). "Ambergris, Treasure of the Deep". Bloomberg L.P. Retrieved 25 May 2017.
  14. ^ Wahlberg, Magnus; Frantzis, Alexandros; Alexiadou, Paraskevi; Madsen, Peter T.; Møhl, Bertel (2005). "Click production during breathing in a sperm whale (Physeter macrocephalus)". The Journal of the Acoustical Society of America. 118 (6): 3404–7. Bibcode:2005ASAJ..118.3404W. doi:10.1121/1.2126930. PMID 16419786.
  15. ^ Haupt, Paul (1907). "Jonah's Whale". Proceedings of the American Philosophical Society. 46 (185): 151–164. JSTOR 983449.
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Further reading

  • Whitehead, H. (2003). Sperm Whales: Social Evolution in the Ocean. Chicago: University of Chicago Press. p. 4. ISBN 978-0-226-89518-5.
  • Perrin, William F.; Würsig, Bernd; Thewissen, J.G.M., eds. (2002). Encyclopedia of Marine Mammals. San Diego, Calif.: Academic Press. ISBN 978-0-12-551340-1.
  • Carwardine, Hoyt; Fordyce & Gill (1998). Whales & Dolphins: The Ultimate Guide to Marine Mammals. London: HarperCollins. ISBN 978-0-00-220105-6.
  • Heptner, V. G.; Nasimovich, A. A; Bannikov, Andrei Grigorevich; Hoffmann, Robert S, Mammals of the Soviet Union, Volume II, part 3 (1996). Washington, D.C.: Smithsonian Institution Libraries and National Science Foundation

External links

 
Wikimedia Commons has media related to Physeter macrocephalus.
 
Wikispecies has information related to Physeter macrocephalus.
 
Wikisource has the text of the 1911 Encyclopædia Britannica article "Sperm-Whale".
  • The Dominica Sperm Whale Project- a long-term scientific research program focusing on the behaviour of sperm whale units.
  • 22 July 2007
  • Society for Marine Mammalogy Sperm Whale Fact Sheet
  • US National Marine Fisheries Service Sperm Whale web page
  • —information on the sperm whale
  • —Photographs, video.
  • —An online documentary film exploring the sperm whales in the Mediterranean Sea.
  • Website of the Memorandum of Understanding for the Conservation of Cetaceans and Their Habitats in the Pacific Islands Region
  • Official website of the Agreement on the Conservation of Cetaceans in the Black Sea, Mediterranean Sea and Contiguous Atlantic Area
  • Retroposon analysis of major cetacean lineages: The monophyly of toothed whales and the paraphyly of river dolphins 19 June 2001
  • Voices in the Sea – sounds of the sperm whale
  • Sperm whales quickly learned to avoid humans who were hunting them in the 19th century, scientists say. ABC News. 16 March 2021.

January 01, 1970
sperm, whale, cachalot, redirects, here, other, uses, cachalot, disambiguation, kashalot, redirects, here, soviet, submarine, kashalot, class, submarine, 2015, film, sperm, whale, film, sperm, whale, cachalot, physeter, macrocephalus, largest, toothed, whales,. Cachalot redirects here For other uses see Cachalot disambiguation Kashalot redirects here For the Soviet submarine see Kashalot class submarine For the 2015 film see Sperm Whale film The sperm whale or cachalot a 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 family along with the pygmy sperm whale and dwarf sperm whale of the genus Kogia Sperm whale 1 Temporal range Pliocene Recent 2 PreꞒ Ꞓ O S D C P T J K Pg N Conservation status Vulnerable IUCN 3 1 3 CITES Appendix I CITES 4 Scientific classification Domain Eukaryota Kingdom Animalia Phylum Chordata Class Mammalia Order Artiodactyla Infraorder Cetacea Family Physeteridae Genus Physeter Species P macrocephalus Binomial name Physeter macrocephalusLinnaeus 1758 Major sperm whale grounds Synonyms Physeter catodon Linnaeus 1758 Physeter microps Linnaeus 1758 Physeter tursio Linnaeus 1758 Physeter australasianus Desmoulins 1822 The sperm whale is a pelagic mammal with a worldwide range and will migrate seasonally for feeding and breeding 5 Females and young males live together in groups while mature males bulls live solitary lives outside of the mating season The females cooperate to protect and nurse their young Females give birth every four to twenty years and care for the calves for more than a decade A mature sperm whale has no natural predators although calves and weakened adults are sometimes killed by pods of killer whales orcas Mature males average 16 metres 52 ft in length with the head representing up to one third of the animal s length Plunging to 2 250 metres 7 382 ft it is the third deepest diving mammal exceeded only by the southern elephant seal and Cuvier s beaked whale 6 7 The sperm whale uses echolocation and vocalization with source level as loud as 236 decibels re 1 µPa m underwater 8 9 It has the largest brain on Earth more than five times heavier than a human s Sperm whales can live 70 years or more 10 11 12 Sperm whales heads are filled with a waxy substance called spermaceti sperm oil from which the whale derives its name Spermaceti was a prime target of the whaling industry and was sought after for use in oil lamps lubricants and candles Ambergris a solid waxy waste product sometimes present in its digestive system is still highly valued as a fixative in perfumes among other uses Beachcombers look out for ambergris as flotsam 13 Sperm whaling was a major industry in the 19th century depicted in the novel Moby Dick The species is protected by the International Whaling Commission moratorium and is listed as vulnerable by the International Union for Conservation of Nature Contents 1 Taxonomy and naming 1 1 Etymology 1 2 Taxonomy 2 Biology 2 1 External appearance 2 2 Skeleton 2 3 Jaws and teeth 2 4 Brain 2 5 Biological systems 2 6 Senses 2 6 1 Spermaceti organ and melon 2 6 2 Eyes and vision 2 7 Sleeping 2 8 Genetics 3 Vocalization complex 3 1 Mechanism 3 2 Vocalization types 3 2 1 Codas 4 Ecology 4 1 Distribution 4 2 Diet 5 Life cycle 6 Social behaviour 6 1 Relations within the species 6 2 Relations with other species 6 3 Parasites 7 Evolutionary history 7 1 Fossil record 7 2 Phylogeny 8 Relationship with humans 8 1 Sperm whaling 8 2 Current conservation status 8 3 Cultural importance 8 4 Watching sperm whales 8 5 Plastic waste 9 See also 10 Notes 11 References 12 Further reading 13 External linksTaxonomy and namingEtymology The name sperm whale is a clipping of spermaceti whale Spermaceti originally mistakenly identified as the whales semen is the semi liquid waxy substance found within the whale s head 14 See Spermaceti organ and melon below The sperm whale is also known as the cachalot which is thought to derive from the archaic French for tooth or big teeth as preserved for example in the word caishau in the Gascon dialect a word of either Romance 15 or Basque 16 origin The etymological dictionary of Corominas says the origin is uncertain but it suggests that it comes from the Vulgar Latin cappula sword hilts 17 The word cachalot came to English via French from Spanish or Portuguese cachalote perhaps from Galician Portuguese cachola big head 18 The term is retained in the Russian word for the animal kashalot kashalot as well as in many other languages citation needed The scientific genus name Physeter comes from the Greek physeter fyshthr meaning blowpipe blowhole of a whale or as a pars pro toto whale citation needed The specific name macrocephalus is Latinized from the Greek makrokephalos makrokefalos big headed from makros makros kephale kefalh citation needed Its synonymous specific name catodon means down tooth from the Greek elements cat a below and odṓn tooth so named because it has visible teeth only in its lower jaw 19 See Jaws and teeth below Another synonym australasianus Australasian was applied to sperm whales in the Southern Hemisphere 20 Taxonomy The sperm whale belongs to the order Cetartiodactyla 21 the order containing all cetaceans and even toed ungulates It is a member of the unranked clade Cetacea with all the whales dolphins and porpoises and further classified into Odontoceti containing all the toothed whales and dolphins It is the sole extant species of its genus Physeter in the family Physeteridae Two species of the related extant genus Kogia the pygmy sperm whale Kogia breviceps and the dwarf sperm whale K sima are placed either in this family or in the family Kogiidae 22 In some taxonomic schemes the families Kogiidae and Physeteridae are combined as the superfamily Physeteroidea see the separate entry on the sperm whale family 23 Swedish ichthyologist Peter Artedi described it as Physeter catodon in his 1738 work Genera piscium from the report of a beached specimen in the Orkneys in 1693 and two beached in the Netherlands in 1598 and 1601 24 The 1598 specimen was near Berkhey citation needed The sperm whale is one of the species originally described by Carl Linnaeus in his landmark 1758 10th edition of Systema Naturae He recognised four species in the genus Physeter 25 Experts soon realised that just one such species exists although there has been debate about whether this should be named P catodon or P macrocephalus two of the names used by Linnaeus Both names are still used although most recent authors now accept macrocephalus as the valid name limiting catodon s status to a lesser synonym Until 1974 the species was generally known as P catodon In that year however Dutch zoologists Antonius M Husson and Lipke Holthuis proposed that the correct name should be P macrocephalus the second name in the genus Physeter published by Linnaeus concurrently with P catodon This proposition was based on the grounds that the names were synonyms published simultaneously and therefore the ICZN Principle of the First Reviser should apply In this instance it led to the choice of P macrocephalus over P catodon a view re stated in Holthuis 1987 26 This has been adopted by most subsequent authors although Schevill 1986 27 and 1987 28 argued that macrocephalus was published with an inaccurate description and that therefore only the species catodon was valid rendering the principle of First Reviser inapplicable The most recent version of ITIS has altered its usage from P catodon to P macrocephalus 29 following L B Holthuis and more recent 2008 discussions with relevant experts 30 31 Furthermore The Taxonomy Committee of the Society for Marine Mammalogy the largest international association of marine mammal scientists in the world officially uses Physeter macrocephalus when publishing their definitive list of marine mammal species 32 BiologyExternal appearance Average sizes 10 33 Length Weight Male 16 metres 52 ft 45 tonnes 50 short tons Female 11 metres 36 ft 15 tonnes 17 short tons Newborn 4 metres 13 ft 1 tonne 1 1 short tons The sperm whale is the largest toothed whale and is among the most sexually dimorphic of all cetaceans 34 Both sexes are about the same size at birth 10 but mature males are typically 30 to 50 longer and three times as massive as females 35 36 Newborn sperm whales are usually between 3 7 and 4 3 meters 12 and 14 ft long 37 Female sperm whales are sexually mature at 8 to 9 meters 26 to 30 ft in length whilst males are sexually mature at 11 to 12 meters 36 to 39 ft 38 Female sperm whales are physically mature at about 10 6 to 11 meters 35 to 36 ft in length and generally do not achieve lengths greater than 12 metres 39 ft 34 36 38 The largest female sperm whale measured up to 12 3 meters 40 ft long and an individual of such size would have weighed about 17 tonnes 19 short tons 39 40 Male sperm whales are physically mature at about 15 to 16 meters 49 to 52 ft in length and larger males can generally achieve 18 to 19 meters 59 to 62 ft 38 41 34 An 18 meters 59 ft long male sperm whale is estimated to have weighed 57 tonnes 56 long tons 63 short tons 33 By contrast the second largest toothed whale Baird s beaked whale measures up to 12 8 meters 42 ft and weighs up to 14 tonnes 15 short tons 42 There are occasional reports of individual sperm whales achieving even greater lengths with some historical claims reaching or exceeding 80 feet 24 m One example is the whale that sank the Essex one of the incidents behind Moby Dick which was claimed to be 85 feet 26 m However there is disagreement as to the accuracy of some of these claims which are often considered exaggerations or as being measured along the curves of the body 43 34 41 An individual measuring 20 7 metres 68 ft was reported from a Soviet whaling fleet near the Kuril Islands in 1950 and is cited by some authors as the largest accurately measured 34 44 It has been estimated to weigh 80 tonnes 79 long tons 88 short tons 43 In a review of size variation in marine megafauna McClain and colleagues noted that the International Whaling Commission s data contained eight individuals larger than 20 7 metres 68 ft The authors supported a 24 metre 79 ft male from the South Pacific in 1933 as the largest recorded However sizes like these are rare with 95 of recorded sperm whales below 15 85 metres 52 0 ft 34 In 1853 one sperm whale was reported at 62 feet 19 m in length with a head measuring 20 feet 6 1 m 45 Large lower jawbones are held in the British Natural History Museum and the Oxford University Museum of Natural History measuring 5 metres 16 ft and 4 7 metres 15 ft respectively 43 The average size of sperm whales has decreased over the years probably due to pressure from whaling 34 Another view holds that exploitation by overwhaling had virtually no effect on the size of the bull sperm whales and their size may have actually increased in current times on the basis of density dependent effects 46 Old males taken at Solander Islands were recorded to be extremely large and unusually rich in blubbers 47 nbsp Unusual among cetaceans the sperm whale s blowhole is highly skewed to the left side of the head The sperm whale s unique body is unlikely to be confused with any other species The sperm whale s distinctive shape comes from its very large block shaped head which can be one quarter to one third of the animal s length The S shaped blowhole is located very close to the front of the head and shifted to the whale s left 35 This gives rise to a distinctive bushy forward angled spray citation needed The sperm whale s flukes tail lobes are triangular and very thick Proportionally they are larger than that of any other cetacean and are very flexible 48 The whale lifts its flukes high out of the water as it begins a feeding dive 35 It has a series of ridges on the back s caudal third instead of a dorsal fin The largest ridge was called the hump by whalers and can be mistaken for a dorsal fin because of its shape and size 10 In contrast to the smooth skin of most large whales its back skin is usually wrinkly and has been likened to a prune by whale watching enthusiasts 49 Albinos have been reported 11 50 Skeleton nbsp A sperm whale skeleton The ribs are bound to the spine by flexible cartilage which allows the ribcage to collapse rather than snap under high pressure 51 While sperm whales are well adapted to diving repeated dives to great depths have long term effects Bones show the same avascular necrosis that signals decompression sickness in humans Older skeletons showed the most extensive damage whereas calves showed no damage This damage may indicate that sperm whales are susceptible to decompression sickness and sudden surfacing could be lethal to them 52 Like that of all cetaceans the spine of the sperm whale has reduced zygapophysial joints of which the remnants are modified and are positioned higher on the vertebral dorsal spinous process hugging it laterally to prevent extensive lateral bending and facilitate more dorso ventral bending These evolutionary modifications make the spine more flexible but weaker than the spines of terrestrial vertebrates 53 nbsp Labeled sperm whale skeleton Like many cetaceans the sperm whale has a vestigial pelvis that is not connected to the spine citation needed Like that of other toothed whales the skull of the sperm whale is asymmetrical so as to aid echolocation Sound waves that strike the whale from different directions will not be channeled in the same way 54 Within the basin of the cranium the openings of the bony narial tubes from which the nasal passages spring are skewed towards the left side of the skull citation needed Jaws and teeth nbsp Sperm whale tooth nbsp The lower jaw is long and narrow The teeth fit into sockets along the upper jaw lifelike sculpture The sperm whale s lower jaw is very narrow and underslung 55 The sperm whale has 18 to 26 teeth on each side of its lower jaw which fit into sockets in the upper jaw 55 The teeth are cone shaped and weigh up to 1 kilogram 2 2 lb each 56 The teeth are functional but do not appear to be necessary for capturing or eating squid as well fed animals have been found without teeth or even with deformed jaws One hypothesis is that the teeth are used in aggression between males 57 Mature males often show scars which seem to be caused by the teeth citation needed Rudimentary teeth are also present in the upper jaw but these rarely emerge into the mouth 58 Analyzing the teeth is the preferred method for determining a whale s age Like the age rings in a tree the teeth build distinct layers of cementum and dentine as they grow 59 Brain nbsp The sperm whale s brain is the largest in the world five times heavier than a human brain The sperm whale brain is the largest known of any modern or extinct animal weighing on average about 7 8 kilograms 17 lb 60 61 with the smallest known weighing 6 4 kilograms 14 lb and the largest known weighing 9 2 kilograms 20 lb 43 44 more than five times heavier than a human brain and has a volume of about 8 000 cm3 62 Although larger brains generally correlate with higher intelligence it is not the only factor Elephants and dolphins also have larger brains than humans 63 The sperm whale has a lower encephalization quotient than many other whale and dolphin species lower than that of non human anthropoid apes and much lower than that of humans 61 64 The sperm whale s cerebrum is the largest in all mammalia both in absolute and relative terms The olfactory system is reduced suggesting that the sperm whale has a poor sense of taste and smell By contrast the auditory system is enlarged The pyramidal tract is poorly developed reflecting the reduction of its limbs 65 Biological systems See also Physiology of underwater diving Marine mammals The sperm whale respiratory system has adapted to cope with drastic pressure changes when diving The flexible ribcage allows lung collapse reducing nitrogen intake and metabolism can decrease to conserve oxygen 66 67 Between dives the sperm whale surfaces to breathe for about eight minutes before diving again 35 Odontoceti toothed whales breathe air at the surface through a single S shaped blowhole which is extremely skewed to the left Sperm whales spout breathe 3 5 times per minute at rest increasing to 6 7 times per minute after a dive The blow is a noisy single stream that rises up to 2 metres 6 6 ft or more above the surface and points forward and left at a 45 angle 68 On average females and juveniles blow every 12 5 seconds before dives while large males blow every 17 5 seconds before dives 69 A sperm whale killed 160 km 100 mi south of Durban South Africa after a 1 hour 50 minute dive was found with two dogfish Scymnodon sp usually found at the sea floor in its belly 70 The sperm whale has the longest intestinal system in the world 71 exceeding 300 m in larger specimens 72 73 The sperm whale has a four chambered stomach that is similar to ruminants The first secretes no gastric juices and has very thick muscular walls to crush the food since whales cannot chew and resist the claw and sucker attacks of swallowed squid The second chamber is larger and is where digestion takes place Undigested squid beaks accumulate in the second chamber as many as 18 000 have been found in some dissected specimens 72 74 75 Most squid beaks are vomited by the whale but some occasionally make it to the hindgut Such beaks precipitate the formation of ambergris 75 nbsp The arterial system of a sperm whale foetus In 1959 the heart of a 22 metric ton 24 short ton male taken by whalers was measured to be 116 kilograms 256 lb about 0 5 of its total mass 76 The circulatory system has a number of specific adaptations for the aquatic environment The diameter of the aortic arch increases as it leaves the heart This bulbous expansion acts as a windkessel ensuring a steady blood flow as the heart rate slows during diving 77 The arteries that leave the aortic arch are positioned symmetrically There is no costocervical artery There is no direct connection between the internal carotid artery and the vessels of the brain 78 Their circulatory system has adapted to dive at great depths as much as 2 250 metres 7 382 ft 6 7 79 80 81 excessive citations for up to 120 minutes 82 More typical dives are around 400 metres 1 310 ft and 35 minutes in duration 35 Myoglobin which stores oxygen in muscle tissue is much more abundant than in terrestrial animals 83 The blood has a high density of red blood cells which contain oxygen carrying haemoglobin The oxygenated blood can be directed towards only the brain and other essential organs when oxygen levels deplete 84 85 86 The spermaceti organ may also play a role by adjusting buoyancy see below 87 The arterial retia mirabilia are extraordinarily well developed The complex arterial retia mirabilia of the sperm whale are more extensive and larger than those of any other cetacean 78 Senses Spermaceti organ and melon nbsp Anatomy of the sperm whale s head The organs above the jaw are devoted to sound generation Atop the whale s skull is positioned a large complex of organs filled with a liquid mixture of fats and waxes called spermaceti The purpose of this complex is to generate powerful and focused clicking sounds the existence of which was proven by Valentine Worthington and William Schevill when a recording was produced on a research vessel in May 1959 88 The sperm whale uses these sounds for echolocation and communication 89 90 91 92 93 94 95 96 excessive citations The spermaceti organ is like a large barrel of spermaceti Its surrounding wall known as the case is extremely tough and fibrous The case can hold within it up to 1 900 litres of spermaceti 97 It is proportionately larger in males 98 This oil is a mixture of triglycerides and wax esters The proportion of wax esters in the spermaceti organ increases with the age of the whale 38 51 in calves 58 87 in adult females and 71 94 in adult males 99 The spermaceti at the core of the organ has a higher wax content than the outer areas 100 The speed of sound in spermaceti is 2 684 m s at 40 kHz 36 C making it nearly twice as fast as in the oil in a dolphin s melon 101 Below the spermaceti organ lies the junk which consists of compartments of spermaceti separated by cartilage It is analogous to the melon found in other toothed whales 102 The structure of the junk redistributes physical stress across the skull and may have evolved to protect the head during ramming 103 104 102 Running through the head are two air passages The left passage runs alongside the spermaceti organ and goes directly to the blowhole whilst the right passage runs underneath the spermaceti organ and passes air through a pair of phonic lips and into the distal sac at the very front of the nose The distal sac is connected to the blowhole and the terminus of the left passage When the whale is submerged it can close the blowhole and air that passes through the phonic lips can circulate back to the lungs The sperm whale unlike other odontocetes has only one pair of phonic lips whereas all other toothed whales have two 105 and it is located at the front of the nose instead of behind the melon At the posterior end of this spermaceti complex is the frontal sac which covers the concave surface of the cranium The posterior wall of the frontal sac is covered with fluid filled knobs which are about 4 13 mm in diameter and separated by narrow grooves The anterior wall is smooth The knobbly surface reflects sound waves that come through the spermaceti organ from the phonic lips The grooves between the knobs trap a film of air that is consistent whatever the orientation or depth of the whale making it an excellent sound mirror 101 The spermaceti organs may also help adjust the whale s buoyancy It is hypothesized that before the whale dives cold water enters the organ and it is likely that the blood vessels constrict reducing blood flow and hence temperature The wax therefore solidifies and reduces in volume 87 106 The increase in specific density generates a down force of about 392 newtons 88 lbf and allows the whale to dive with less effort citation needed During the hunt oxygen consumption together with blood vessel dilation produces heat and melts the spermaceti increasing its buoyancy and enabling easy surfacing 107 However more recent work 91 has found many problems with this theory including the lack of anatomical structures for the actual heat exchange 108 Another issue is that if the spermaceti does indeed cool and solidify it would affect the whale s echolocation ability just when it needs it to hunt in the depths citation needed Herman Melville s fictional story Moby Dick suggests that the case containing the spermaceti serves as a battering ram for use in fights between males 109 A few famous instances include the well documented sinking of the ships Essex and Ann Alexander by attackers estimated to weigh only one fifth as much as the ships 102 nbsp The phonic lips nbsp The frontal sac exposed Its surface is covered with fluid filled knobs nbsp A piece of the posterior wall of the frontal sac The grooves between the knobs trap a consistent film of air making it an excellent sound mirror 101 Eyes and vision source source source source source source source Like other toothed whales the sperm whale can retract its eyes The sperm whale s eye does not differ greatly from those of other toothed whales except in size It is the largest among the toothed whales weighing about 170 g It is overall ellipsoid in shape compressed along the visual axis measuring about 7 7 3 cm The cornea is elliptical and the lens is spherical The sclera is very hard and thick roughly 1 cm anteriorly and 3 cm posteriorly There are no ciliary muscles The choroid is very thick and contains a fibrous tapetum lucidum Like other toothed whales the sperm whale can retract and protrude its eyes thanks to a 2 cm thick retractor muscle attached around the eye at the equator 110 but are unable to roll the eyes in their sockets 111 According to Fristrup and Harbison 2002 112 sperm whale s eyes afford good vision and sensitivity to light They conjectured that sperm whales use vision to hunt squid either by detecting silhouettes from below or by detecting bioluminescence If sperm whales detect silhouettes Fristrup and Harbison suggested that they hunt upside down allowing them to use the forward parts of the ventral visual fields for binocular vision citation needed Sleeping For some time researchers have been aware that pods of sperm whales may sleep for short periods assuming a vertical position with their heads just below or at the surface or head down 113 A 2008 study published in Current Biology recorded evidence that whales may sleep with both sides of the brain It appears that some whales may fall into a deep sleep for about 7 percent of the time most often between 6 p m and midnight 114 Genetics Sperm whales have 21 pairs of chromosomes 2n 42 115 The genome of live whales can be examined by recovering shed skin 116 Vocalization complexFurther information on the sonar operator slang term Carpenter fish source source Sperm whale vocalization After Valentine Worthington and William E Schevill confirmed the existence of sperm whale vocalization 88 further studies found that sperm whales are capable of emitting sounds at a volume of 230 decibels more than an aircraft jet engine at takeoff making the sperm whale the loudest animal in the world 117 Mechanism When echolocating the sperm whale emits a directionally focused beam of broadband clicks Clicks are generated by forcing air through a pair of phonic lips also known as monkey lips or museau de singe at the front end of the nose just below the blowhole The sound then travels backwards along the length of the nose through the spermaceti organ Most of the sound energy is then reflected off the frontal sac at the cranium and into the melon whose lens like structure focuses it 89 Some of the sound will reflect back into the spermaceti organ and back towards the front of the whale s nose where it will be reflected through the spermaceti organ a third time This back and forth reflection which happens on the scale of a few milliseconds creates a multi pulse click structure 118 This multi pulse click structure allows researchers to measure the whale s spermaceti organ using only the sound of its clicks 119 120 Because the interval between pulses of a sperm whale s click is related to the length of the sound producing organ an individual whale s click is unique to that individual However if the whale matures and the size of the spermaceti organ increases the tone of the whale s click will also change 120 The lower jaw is the primary reception path for the echoes A continuous fat filled canal transmits received sounds to the inner ear 121 The source of the air forced through the phonic lips is the right nasal passage While the left nasal passage opens to the blow hole the right nasal passage has evolved to supply air to the phonic lips It is thought that the nostrils of the land based ancestor of the sperm whale migrated through evolution to their current functions the left nostril becoming the blowhole and the right nostril becoming the phonic lips 122 Air that passes through the phonic lips passes into the distal sac then back down through the left nasal passage This recycling of air allows the whale to continuously generate clicks for as long as it is submerged 123 Vocalization types The sperm whale s vocalizations are all based on clicking described in four types the usual echolocation creaks codas and slow clicks 124 The usual echolocation click type is used in searching for prey 124 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 124 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 125 Characteristics of sperm whale clicks 124 Click type Apparent source level dB re 1 µPa m Directionality Centroidfrequency kHz Inter click interval s Duration of click ms Duration of pulse ms Range audible to sperm whale km Inferred function Audio sample Usual 230 High 15 0 5 1 0 15 30 0 1 16 Searching for prey source source Creak 205 High 15 0 005 0 1 0 1 5 0 1 6 Homing in on prey source source Coda 180 Low 5 0 1 0 5 35 0 5 2 Social communication source source Slow 190 Low 0 5 5 8 30 5 60 Communication by males source source Codas The most distinctive vocalizations are codas which are short rhythmic sequences of clicks mostly numbering 3 12 clicks in stereotyped patterns 126 They are classified using variations in the number of clicks rhythm and tempo 127 Codas are the result of vocal learning within a stable social group 128 and are made in the context of the whales social unit 126 The foundation of sperm whale society is the matrilineally based social unit of ten or so females and their offspring The members of the unit travel together suckle each others infants and babysit them while mothers make long deep dives to feed 126 Over 70 of a sperm whale s time is spent independently foraging codas could help whales reunite and reaffirm their social ties in between long foraging dives 127 129 While nonidentity codas are commonly used in multiple different clans 130 some codas express clan identity and denote different patterns of travel foraging and socializing or avoidance among clans 129 131 In particular whales will not group with whales of another clan even though they share the same geographical area 126 Statistically as the clans ranges become more overlapped the distinction in clan identity coda usage becomes more pronounced 130 Distinctive codas identify seven clans described among the approximately 150 000 female sperm whales in the Pacific Ocean and there are another four clans in the Atlantic 126 As arbitrary traits that function as reliable indicators of cultural group membership clan identity codas act as symbolic markers that modulate interactions between individuals 127 Individual identity in sperm whale vocalizations is an ongoing scientific issue however A distinction needs to be made between cues and signals Human acoustic tools can distinguish individual whales by analyzing micro characteristics of their vocalizations 132 and the whales can probably do the same This does not prove that the whales deliberately use some vocalizations to signal individual identity in the manner of the signature whistles that bottlenose dolphins use as individual labels 128 127 EcologyDistribution nbsp Global concentrations of sperm whales Sperm whales are among the most cosmopolitan species They prefer ice free waters over 1 000 metres 3 300 ft deep 3 Although both sexes range through temperate and tropical oceans and seas only adult males populate higher latitudes 11 Among several regions such as along coastal waters of southern Australia sperm whales have been considered to be locally extinct 133 They are relatively abundant from the poles to the equator and are found in all the oceans They inhabit the Mediterranean Sea but not the Black Sea 10 while their presence in the Red Sea is uncertain 3 The shallow entrances to both the Black Sea and the Red Sea may account for their absence 134 The Black Sea s lower layers are also anoxic and contain high concentrations of sulphur compounds such as hydrogen sulphide 135 What does being anoxic have to do with whales that don t breath under water Lack of prey The first ever sighting off the coast of Pakistan was made in 2017 136 137 The first ever record off the west coast of the Korean Peninsula Yellow Sea was made in 2005 138 139 followed by one near Ganghwa Island in 2009 140 Populations are denser close to continental shelves and canyons 11 Sperm whales are usually found in deep off shore waters but may be seen closer to shore in areas where the continental shelf is small and drops quickly to depths of 310 to 920 metres 1 020 to 3 020 ft 10 Coastal areas with significant sperm whale populations include the Azores and Dominica 141 In east Asian waters whales are also observed regularly in coastal waters in places such as the Commander and Kuril Islands Shiretoko Peninsula which is one of few locations where sperm whales can be observed from shores 142 off Kinkasan vicinity to Tokyo Bay 143 and the Bōsō Peninsula to the Izu 144 145 and the Izu Islands the Volcano Islands Yakushima and the Tokara Islands to the Ryukyu Islands 146 147 Taiwan the Northern Mariana Islands 148 and so forth Historical catch records suggest there could have been smaller aggression grounds in the Sea of Japan as well 149 Along the Korean Peninsula the first confirmed observation within the Sea of Japan eight animals off Guryongpo was made in 2004 since after the last catches of five whales off Ulsan in 1911 139 150 while nine whales were observed in the East China Sea side of the peninsula in 1999 151 Grown males are known to enter surprisingly shallow bays to rest whales will be in a state of rest during these occasions Unique coastal groups have been reported from various areas around the globe such as near Scotland s coastal waters 152 and the Shiretoko Peninsula off Kaikōura in Davao Gulf Such coastal groups were more abundant in pre whaling days 153 Genetic analysis indicates that the world population of sperm whales originated in the Pacific Ocean from a population of about 10 000 animals around 100 000 years ago when expanding ice caps blocked off their access to other seas In particular colonization of the Atlantic was revealed to have occurred multiple times during this expansion of their range 154 Diet nbsp A piece of sperm whale skin with giant squid sucker scars Sperm whales usually dive between 300 and 800 metres 980 and 2 620 ft and sometimes 1 to 2 kilometres 3 300 to 6 600 ft in search of food 155 Such dives can last more than an hour 155 They feed on several species notably the giant squid but also the colossal squid octopuses and fish such as demersal rays and sharks citation needed but their diet is mainly medium sized squid 156 Some prey may be taken accidentally while eating other items 156 Most of what is known about deep sea squid has been learned from specimens in captured sperm whale stomachs although more recent studies analysed faeces One study carried out around the Galapagos found that squid from the genera Histioteuthis 62 Ancistrocheirus 16 and Octopoteuthis 7 weighing between 12 and 650 grams 0 026 and 1 433 lb were the most commonly taken 157 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 One study published in 2010 collected evidence that suggests that female sperm whales may collaborate when hunting Humboldt squid 158 Tagging studies have shown that sperm whales hunt upside down at the bottom of their deep dives It is suggested that the whales can see the squid silhouetted above them against the dim surface light 159 An older study examining whales captured by the New Zealand whaling fleet in the Cook Strait region found a 1 69 1 ratio of squid to fish by weight 160 Sperm whales sometimes take sablefish and toothfish from long lines Long line fishing operations in the Gulf of Alaska complain that sperm whales take advantage of their fishing operations to eat desirable species straight off the line sparing the whales the need to hunt 161 However the amount of fish taken is very little compared to what the sperm whale needs per day Video footage has been captured of a large male sperm whale bouncing a long line to gain the fish 162 Sperm whales are believed to prey on the megamouth shark a rare and large deep sea species discovered in the 1970s 163 In one case three sperm whales were observed attacking or playing with a megamouth 164 Sperm whales have also been noted to feed on bioluminescent pyrosomes such as Pyrosoma atlanticum 165 166 167 It is thought that the foraging strategy of sperm whales for bioluminescent squids may also explain the presence of these light emitting pyrosomes in the diet of the sperm whale 167 nbsp Ambergris The sharp beak of a consumed squid lodged in the whale s intestine may lead to the production of ambergris analogous to the production of pearls in oysters 168 The irritation of the intestines caused by squid beaks stimulates the secretion of this lubricant like substance Sperm whales are prodigious feeders and eat around 3 of their body weight per day The total annual consumption of prey by sperm whales worldwide is estimated to be about 272 million tonnes 300 million short tons 169 170 In comparison human consumption of seafood is estimated to be 157 million tonnes 173 million short tons 171 Sperm whales hunt through echolocation Their clicks are among the most powerful sounds in the animal kingdom see above It has been hypothesised that it can stun prey with its clicks Experimental studies attempting to duplicate this effect have been unable to replicate the supposed injuries casting doubt on this idea 172 One study showing that sound pressure levels on the squid are more than an order of magnitude below levels required for debilitation and therefore precluding acoustic stunning to facilitate prey capture 173 Sperm whales as well as other large cetaceans help fertilise the surface of the ocean by consuming nutrients in the depths and transporting those nutrients to the oceans surface when they defecate an effect known as the whale pump 174 This fertilises phytoplankton and other plants on the surface of the ocean and contributes to ocean productivity and the drawdown of atmospheric carbon 175 Life cycleSee also Whale reproduction Sperm whales can live 70 years or more 10 11 12 They are a prime example of a species that has been K selected meaning their reproductive strategy is associated with stable environmental conditions and comprises a low birth rate significant parental aid to offspring slow maturation and high longevity 35 How they choose mates has not been definitively determined Bulls will fight with each other over females and males will mate with multiple females making them polygynous but they do not dominate the group as in a harem 176 177 Bulls do not provide paternal care to their offspring but rather play a fatherly role to younger bulls to show dominance 178 Females become fertile at around 9 years of age 179 The oldest pregnant female ever recorded was 41 years old 180 Gestation requires 14 to 16 months producing a single calf 10 Sexually mature females give birth once every 4 to 20 years pregnancy rates were higher during the whaling era 179 Birth is a social event as the mother and calf need others to protect them from predators The other adults may jostle and bite the newborn in its first hours 181 Lactation proceeds for 19 to 42 months but calves rarely may suckle up to 13 years 10 Like that of other whales the sperm whale s milk has a higher fat content than that of terrestrial mammals about 36 182 compared to 4 in cow milk This gives it a consistency similar to cottage cheese 183 which prevents it from dissolving in the water before the calf can drink it 184 It has an energy content of roughly 3 840 kcal kg 182 compared to just 640 kcal kg in cow milk 185 Calves may be allowed to suckle from females other than their mothers 10 Males become sexually mature at 18 years Upon reaching sexual maturity males move to higher latitudes where the water is colder and feeding is more productive Females remain at lower latitudes 10 Males reach their full size at about age 50 35 Social behaviourRelations within the species nbsp Sperm whales adopt the marguerite formation to defend a vulnerable pod member Like elephants females and their young live in matriarchal groups called pods while bulls live apart Bulls sometimes form loose bachelor groups with other males of similar age and size As they grow older they typically live solitary lives only returning to the pod to socialize or to breed 35 Bulls have beached themselves together suggesting a degree of cooperation which is not yet fully understood 35 The whales rarely if ever leave their group 186 A social unit is a group of sperm whales who live and travel together over a period of years Individuals rarely if ever join or leave a social unit There is a huge variance in the size of social units They are most commonly between six and nine individuals in size but can have more than twenty 187 Unlike orcas sperm whales within a social unit show no significant tendency to associate with their genetic relatives 188 Females and calves spend about three quarters of their time foraging and a quarter of their time socializing Socializing usually takes place in the afternoon 189 When sperm whales socialize they emit complex patterns of clicks called codas They will spend much of the time rubbing against each other Tracking of diving whales suggests that groups engage in herding of prey similar to bait balls created by other species though the research needs to be confirmed by tracking the prey 190 191 Relations with other speciesThe most common natural predator of sperm whales is the orca killer whale but pilot whales and false killer whales sometimes harass them 192 193 Orcas prey on target groups of females with young usually making an effort to extract and kill a calf The females will protect their calves or an injured adult by encircling them They may face inwards with their tails out the marguerite formation named after the flower The heavy and powerful tail of an adult whale is potentially capable of delivering lethal blows 194 Alternatively they may face outwards the heads out formation Other than sperm whales southern right whales had been observed to perform similar formations 195 However formations in non dangerous situations have been recorded as well 195 Early whalers exploited this behaviour attracting a whole unit by injuring one of its members 196 Such a tactic is described in Moby Dick Say you strike a Forty barrel bull poor devil all his comrades quit him But strike a member of the harem school and her companions swim around her with every token of concern sometimes lingering so near her and so long as themselves to fall a prey 197 If the killer whale pod is large its members may sometimes be able to kill adult female sperm whales and can at least injure an entire pod of sperm whales 198 199 Bulls have no predators and are believed to be too large powerful and aggressive to be threatened by killer whales 200 Solitary bulls are known to interfere and come to the aid of vulnerable groups nearby 201 However the bull sperm whale when accompanying pods of female sperm whales and their calves as such may be reportedly unable to effectively dissuade killer whales from their attacks on the group although the killer whales may end the attack sooner when a bull is present 202 203 However male sperm whales have been observed to attack and intimidate killer whale pods in competitive feeding instances An incident was filmed from a long line trawler a killer whale pod was systematically taking fish caught on the trawler s long lines as the lines were being pulled into the ship when a male sperm whale appeared to repeatedly charge the killer whale pod in an attempt to drive them away it was speculated by the film crew that the sperm whale was attempting to access the same fish The killer whales employed a tail outward and tail slapping defensive position against the bull sperm whale similar to that used by female sperm whales against attacking killer whales 204 However at some potential feeding sites the killer whales may prevail over sperm whales even when outnumbered by the sperm whales Some authors consider the killer whales usually behaviorally dominant over sperm whales but express that the two species are fairly evenly matched with the killer whales greater aggression more considerable biting force for their size and predatory prowess more than compensating for their smaller size 202 205 Sperm whales are not known for forging bonds with other species but it was observed that a bottlenose dolphin with a spinal deformity had been accepted into a pod of sperm whales 206 They are known to swim alongside other cetaceans such as humpback 207 fin minke pilot 208 and killer whales on occasion 209 Parasites Sperm whales can suffer from parasites Out of 35 sperm whales caught during the 1976 1977 Antarctic whaling season all of them were infected by Anisakis physeteris in their stomachs and Phyllobothrium delphini in their blubber Both whales with a placenta were infected with Placentonema gigantissima 210 potentially the largest nematode worm ever described citation needed Evolutionary historySee also Sperm whale family Cetacea Toothed whales Physeteroidea Other Physeteroidea Kogiidae Pygmy sperm whale Dwarf sperm whale Physeteridae Other Physeteridae Sperm whale South Asian river dolphin Other river dolphins Oceanic dolphins Porpoises Arctic whales Beaked whales Baleen whales Evolutionary family tree of sperm whales 211 including simplified summary of extinct groups 212 Fossil record Although the fossil record is poor 213 several extinct genera have been assigned to the clade Physeteroidea which includes the last common ancestor of the modern sperm whale pygmy sperm whales dwarf sperm whales and extinct physeteroids These fossils include Ferecetotherium Idiorophus Diaphorocetus Aulophyseter Orycterocetus Scaldicetus Placoziphius Zygophyseter and Acrophyseter 23 212 214 Ferecetotherium found in Azerbaijan and dated to the late Oligocene about 28 to 23 million years ago is the most primitive fossil that has been found which possesses sperm whale specific features such as an asymmetric rostrum beak or snout 215 Most sperm whale fossils date from the Miocene period 23 to 5 million years ago Diaphorocetus from Argentina has been dated to the early Miocene Fossil sperm whales from the Middle Miocene include Aulophyseter Idiorophus and Orycterocetus all of which were found on the West Coast of the United States and Scaldicetus found in Europe and Japan 215 216 Orycterocetus fossils have also been found in the North Atlantic Ocean and the Mediterranean Sea in addition to the west coast of the United States 217 Placoziphius found in Europe and Acrophyseter from Peru are dated to the late Miocene 23 215 Fossil sperm whales differ from modern sperm whales in tooth count and the shape of the face and jaws 215 For example Scaldicetus had a tapered rostrum 216 Genera from the Oligocene and early and middle Miocene with the possible exception of Aulophyseter had teeth in their upper jaws 215 Acrophyseter from the late Miocene also had teeth in both the upper and lower jaws as well as a short rostrum and an upward curving mandible lower jaw 23 These anatomical differences suggest that fossil species may not have necessarily been deep sea squid eaters such as the modern sperm whale but that some genera mainly ate fish 215 Zygophyseter dated from the middle to late Miocene and found in southern Italy had teeth in both jaws and appears to have been adapted to feed on large prey rather like the modern killer whale orca Other fossil sperm whales with adaptations similar to this are collectively known as killer sperm whales 212 Two poorly known fossil species belonging to the modern genus Physeter have been recognized so far P antiquus Neogene of France 218 and P vetus Neogene of eastern North America 219 Physeter vetus is very likely an invalid species as the few teeth that were used to identify this species appear to be identical to those of another toothed whale Orycterocetus quadratidens 220 Phylogeny The traditional view has been that Mysticeti baleen whales and Odontoceti toothed whales arose from more primitive whales early in the Oligocene period and that the super family Physeteroidea which contains the sperm whale dwarf sperm whale and pygmy sperm whale diverged from other toothed whales soon after that over 23 million years ago 213 215 From 1993 to 1996 molecular phylogenetics analyses by Milinkovitch and colleagues based on comparing the genes of various modern whales suggested that the sperm whales are more closely related to the baleen whales than they are to other toothed whales which would have meant that Odontoceti were not monophyletic in other words it did not consist of a single ancestral toothed whale species and all its descendants 211 However more recent studies based on various combinations of comparative anatomy and molecular phylogenetics criticised Milinkovitch s analysis on technical grounds and reaffirmed that the Odontoceti are monophyletic 211 221 222 These analyses also confirm that there was a rapid evolutionary radiation diversification of the Physeteroidea in the Miocene period 212 The Kogiidae dwarf and pygmy sperm whales diverged from the Physeteridae true sperm whales at least 8 million years ago 221 Relationship with humansSperm whaling Main articles Whaling and Sperm whaling nbsp In the 19th century sperm whales were hunted using rowboats and hand thrown harpoons a rather dangerous method as the whales sometimes fought back Spermaceti obtained primarily from the spermaceti organ and sperm oil obtained primarily from the blubber in the body were much sought after by 18th 19th and 20th century whalers These substances found a variety of commercial applications such as candles soap cosmetics machine oil other specialised lubricants lamp oil pencils crayons leather waterproofing rust proofing materials and many pharmaceutical compounds 223 224 225 226 Ambergris a highly expensive solid waxy flammable substance produced in the digestive system of sperm whales was also sought as a fixative in perfumery citation needed Prior to the early eighteenth century hunting was mostly by indigenous Indonesians 227 Legend has it that sometime in the early 18th century around 1712 Captain Christopher Hussey while cruising for right whales near shore was blown offshore by a northerly wind where he encountered a sperm whale pod and killed one 228 Although the story may not be true sperm whales were indeed soon exploited by American whalers Judge Paul Dudley in his Essay upon the Natural History of Whales 1725 states that a certain Atkins 10 or 12 years in the trade was among the first to catch sperm whales sometime around 1720 off the New England coast 229 There were only a few recorded instances during the first few decades 1709 1730s of offshore sperm whaling Instead sloops concentrated on the Nantucket Shoals where they would have taken right whales or went to the Davis Strait region to catch bowhead whales By the early 1740s with the advent of spermaceti candles before 1743 American vessels began to focus on sperm whales The diary of Benjamin Bangs 1721 1769 shows that along with the bumpkin sloop he sailed he found three other sloops flensing sperm whales off the coast of North Carolina in late May 1743 230 On returning to Nantucket in the summer 1744 on a subsequent voyage he noted that 45 spermacetes are brought in here this day another indication that American sperm whaling was in full swing 230 American sperm whaling soon spread from the east coast of the American colonies to the Gulf Stream the Grand Banks West Africa 1763 the Azores 1765 and the South Atlantic 1770s From 1770 to 1775 Massachusetts New York Connecticut and Rhode Island ports produced 45 000 barrels of sperm oil annually compared to 8 500 of whale oil 231 In the same decade the British began sperm whaling employing American ships and personnel 232 By the following decade the French had entered the trade also employing American expertise 232 Sperm whaling increased until the mid nineteenth century Spermaceti oil was important in public lighting for example in lighthouses where it was used in the United States until 1862 when it was replaced by lard oil in turn replaced by petroleum and for lubricating the machines such as those used in cotton mills of the Industrial Revolution Sperm whaling declined in the second half of the nineteenth century as petroleum came into broader use In that sense petroleum use may be said to have protected whale populations from even greater exploitation 233 234 Sperm whaling in the 18th century began with small sloops carrying only one or two whaleboats The fleet s scope and size increased over time and larger ships entered the fishery In the late 18th century and early 19th century sperm whaling ships sailed to the equatorial Pacific the Indian Ocean Japan the coast of Arabia Australia and New Zealand 232 235 236 Hunting could be dangerous to the crew since sperm whales especially bulls will readily fight to defend themselves against attack unlike most baleen whales When dealing with a threat sperm whales will use their huge head effectively as a battering ram 102 Arguably the most famous sperm whale counter attack occurred on 20 November 1820 when a whale claimed to be about 25 9 metres 85 ft long rammed and sank the Nantucket whaleship Essex Only 8 out of 21 sailors survived to be rescued by other ships 237 nbsp Scrimshaw was the art of engraving on the teeth of sperm whales It was a way for whalers to pass the time between hunts The sperm whale s ivory like teeth were often sought by 18th and 19th century whalers who used them to produce inked carvings known as scrimshaw 30 teeth of the sperm whale can be used for ivory Each of these teeth up to 20 cm 8 in and 8 cm 3 in across are hollow for the first half of their length Like walrus ivory sperm whale ivory has two distinct layers However sperm whale ivory contains a much thicker inner layer Though a widely practised art in the 19th century scrimshaw using genuine sperm whale ivory declined substantially after the retirement of the whaling fleets in the 1880s citation needed Modern whaling was more efficient than open boat whaling employing steam powered ships and exploding harpoons Initially modern whaling activity focused on large baleen whales but as these populations were taken sperm whaling increased Spermaceti the fine waxy oil produced by sperm whales was in high demand In both the 1941 1942 and 1942 1943 seasons Norwegian expeditions took over 3 000 sperm whales off the coast of Peru alone After World War II whaling continued unabated to obtain oil for cosmetics and high performance machinery such as automobile transmissions citation needed The hunting led to the near extinction of large whales including sperm whales until bans on whale oil use were instituted in 1972 The International Whaling Commission gave the species full protection in 1985 but hunting by Japan in the northern Pacific Ocean continued until 1988 234 It is estimated that the historic worldwide population numbered 1 100 000 before commercial sperm whaling began in the early 18th century 3 By 1880 it had declined by an estimated 29 percent 3 From that date until 1946 the population appears to have partially recovered as whaling activity decreased but after the Second World War the population declined even further to 33 per cent of the pre whaling population citation needed Between 184 000 and 236 000 sperm whales were killed by the various whaling nations in the 19th century 238 while in the 20th century at least 770 000 were taken the majority between 1946 and 1980 239 nbsp Sperm whaling peaked in the 1830s and 1960s Sperm whales increase levels of primary production and carbon export by depositing iron rich faeces into surface waters of the Southern Ocean The iron rich faeces cause phytoplankton to grow and take up more carbon from the atmosphere When the phytoplankton dies it sinks to the deep ocean and takes the atmospheric carbon with it By reducing the abundance of sperm whales in the Southern Ocean whaling has resulted in an extra 2 million tonnes of carbon remaining in the atmosphere each year 240 Remaining sperm whale populations are large enough that the species conservation status is rated as vulnerable rather than endangered 3 However the recovery from centuries of commercial whaling is a slow process particularly in the South Pacific where the toll on breeding age males was severe 241 Current conservation status The total number of sperm whales in the world is unknown but is thought to be in the hundreds of thousands 3 The conservation outlook is brighter than for many other whales Commercial whaling has ceased 3 and the species is protected almost worldwide though records indicate that in the 11 year period starting from 2000 Japanese vessels have caught 51 sperm whales needs update Fishermen do not target sperm whales to eat 3 but long line fishing operations in the Gulf of Alaska have complained about sperm whales stealing fish from their lines 161 Currently entanglement in fishing nets and collisions with ships represent the greatest threats to the sperm whale population 11 Other threats include ingestion of marine debris ocean noise and chemical pollution 242 The International Union for Conservation of Nature IUCN regards the sperm whale as being vulnerable 3 The species is listed as endangered on the United States Endangered Species Act 243 Sperm whales are listed on Appendix I 244 and Appendix II 244 of the Convention on the Conservation of Migratory Species of Wild Animals CMS It is listed on Appendix I 244 as this species has been categorized as being in danger of extinction throughout all or a significant proportion of their range and CMS Parties strive towards strictly protecting these animals conserving or restoring the places where they live mitigating obstacles to migration and controlling other factors that might endanger them It is listed on Appendix II 244 as it has an unfavourable conservation status or would benefit significantly from international co operation organised by tailored agreements It is also covered by the Agreement on the Conservation of Cetaceans in the Black Sea Mediterranean Sea and Contiguous Atlantic Area ACCOBAMS and the Memorandum of Understanding for the Conservation of Cetaceans and Their Habitats in the Pacific Islands Region Pacific Cetaceans MOU citation needed The species is protected under Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora CITES This makes commercial international trade including in parts and derivatives prohibited with all other international trade strictly regulated through a system of permits and certificates 4 Cultural importance nbsp Sperm whale teeth necklace from Fiji Rope mounted teeth are important cultural objects throughout the Pacific In New Zealand the Maori know them as rei puta such whale tooth pendants were rare objects because sperm whales were not actively hunted in traditional Maori society 245 Whale ivory and bone were taken from beached whales In Fiji the teeth are known as tabua traditionally given as gifts for atonement or esteem called sevusevu and were important in negotiations between rival chiefs 246 Friedrich Ratzel in The History of Mankind reported in 1896 that in Fiji whales or cachalots teeth were the most demanded article of ornament or value They occurred often in necklaces 247 Today the tabua remains an important item in Fijian life The teeth were originally rare in Fiji and Tonga which exported teeth but with the Europeans arrival teeth flooded the market and this currency collapsed The oversupply led in turn to the development of the European art of scrimshaw 248 Herman Melville s novel Moby Dick is based on a true story about a sperm whale that attacked and sank the whaleship Essex 249 250 Melville associated the sperm whale with the Bible s Leviathan 250 251 The fearsome reputation perpetuated by Melville was based on bull whales ability to fiercely defend themselves from attacks by early whalers smashing whaling boats and occasionally attacking and destroying whaling ships citation needed In Jules Verne s Twenty Thousand Leagues Under the Sea the Nautilus fights a group of cachalots sperm whales to protect a pod of southern right whales from their attacks Verne portrays them as being savage hunters nothing but mouth and teeth citation needed The sperm whale was designated as the Connecticut state animal by the General Assembly in 1975 252 It was selected because of its specific contribution to the state s history and because of its present day plight as an endangered species 253 Watching sperm whales See also Whale watching Sperm whales are not the easiest of whales to watch due to their long dive times and ability to travel long distances underwater However due to the distinctive look and large size of the whale watching is increasingly popular citation needed Sperm whale watchers often use hydrophones to listen to the clicks of the whales and locate them before they surface 254 Popular locations for sperm whale watching include the town of Kaikōura on New Zealand s South Island Andenes and Tromso in Arctic Norway as well as the Azores where the continental shelf is so narrow that whales can be observed from the shore 141 255 and Dominica 256 where a long term scientific research program The Dominica Sperm Whale Project has been in operation since 2005 257 Plastic waste The introduction of plastic waste to the ocean environment by humans is relatively new From the 1970s sperm whales have occasionally been found with pieces of plastic in their stomachs 167 258 259 260 See also nbsp Cetaceans portal nbsp Mammals portal nbsp Marine life portal List of sperm whale strandings List of cetaceans Marine biology LivyatanNotes ˈ k ae ʃ e l ɒ t ˈ k ae ʃ e l oʊ cachalot Oxford English Dictionary Online ed Oxford University Press Subscription or participating institution membership required References Mead J G Brownell R L Jr 2005 Order Cetacea In Wilson D E Reeder D M eds Mammal Species of the World A Taxonomic and Geographic Reference 3rd ed Johns Hopkins University Press p 737 ISBN 978 0 8018 8221 0 OCLC 62265494 Physeter macrocephalus Linnaeus 1758 sperm whale Fossilworks Gateway to the Paleobiology Database Retrieved 17 December 2021 a b c d e f g h i j Taylor B L Baird R Barlow J Dawson S M Ford J Mead J G Notarbartolo di Sciara G Wade P Pitman R L 2019 Physeter macrocephalus IUCN Red List of Threatened Species 2019 e T41755A160983555 doi 10 2305 IUCN UK 2008 RLTS T41755A160983555 en Retrieved 19 November 2021 date doi mismatch a b Appendices CITES cites org Retrieved 14 January 2022 Sperm Whale acsonline org Archived from the original on 22 April 2017 Retrieved 13 May 2017 a b Gregory S Schorr Erin A Falcone David J Moretti Russel D Andrews 2014 First long 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Huggenberger Michel Andre amp Helmut H A Oelschlager 2014 The nose of the sperm whale overviews of functional design structural homologies and evolution Journal of the Marine Biological Association of the United Kingdom 96 4 1 24 doi 10 1017 S0025315414001118 hdl 2117 97052 S2CID 27312770 Taxonomy Natural History Museum Retrieved 2013 03 19 Whitehead p 321 Perrin p 1164 Morris Robert J 1975 Further studies into the lipid structure of the spermaceti organ of the sperm whale Physeter catodon Deep Sea Research 22 7 483 489 Bibcode 1975DSRA 22 483M doi 10 1016 0011 7471 75 90021 2 a b c Norris Kenneth S amp Harvey George W 1972 A Theory for the Function of the Spermaceti Organ of the Sperm Whale Animal orientation and navigation NASA a b c d Carrier David R Deban Stephen M Otterstrom Jason 15 June 2002 The face that sank the Essex potential function of the spermaceti organ in aggression Journal of Experimental Biology 205 12 1755 1763 doi 10 1242 jeb 205 12 1755 PMID 12042334 Science Says 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January 2018 Fais A Aguilar Soto N Johnson M Perez Gonzalez C Miller P J O Madsen P T April 2015 Sperm whale echolocation behaviour reveals a directed prior based search strategy informed by prey distribution Behavioral Ecology and Sociobiology 69 4 663 674 doi 10 1007 s00265 015 1877 1 hdl 10023 8168 PMID 12077166 S2CID 13711121 a b c d Whitehead p 135 Whitehead p 144 a b c d e Hal Whitehead 2024 Sperm whale clans and human societies Royal Society Open Science 11 1 Bibcode 2024RSOS 1131353W doi 10 1098 rsos 231353 PMC 10776220 PMID 38204796 a b c d Taylor A Hersh et al 2022 Evidence from sperm whale clans of symbolic marking in non human cultures PNAS 119 37 National Academy of Sciences e2201692119 Bibcode 2022PNAS 11901692H doi 10 1073 pnas 2201692119 hdl 10023 27122 PMC 9478646 PMID 36074817 a b Gero Shane Whitehead Hal Rendell Luke 2016 Individual unit and vocal clan level identity cues in sperm whale codas Royal Society Open Science 3 1 Bibcode 2016RSOS 350372G doi 10 1098 rsos 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ウォッチングセンター 静岡県伊東市城ヶ崎 富戸港 光海丸で行く 本当の大自然との ふれあい ドルフィンウォッチング エコツーリスト エコツーリズム KOHKAIMARU 石井泉 光海丸 Archived from the original on 3 March 2016 a href Template Cite web html title Template Cite web cite web a CS1 maint numeric names authors list link Archived at Ghostarchive and the Wayback Machine 相模湾でマッコウクジラに遭遇 Sperm Whale Encounter in Japan via YouTube 琉球諸島 くじらガイドがお届けするクジラ シャチ イルカ 自然 エコツアー情報 ޥah졪 ϡȥɤYۥ Archived at Ghostarchive and the Wayback Machine Guam Whales via YouTube Smith Tim D Reeves Randall R Josephson Elizabeth A Lund Judith N 27 April 2012 Spatial and Seasonal Distribution of American Whaling and Whales in the Age of Sail PLOS ONE 7 4 e34905 Bibcode 2012PLoSO 734905S doi 10 1371 journal pone 0034905 PMC 3338773 PMID 22558102 JoongAng Ilbo 2004 マッコウクジラ ９０年ぶりに東海出現 Retrieved 17 August 2017 Chang K Zhang C Park C Kang D Ju S Lee S Wimbush M eds 2015 Oceanography of the East Sea Japan Sea Springer International Publishing p 380 ISBN 9783319227207 Retrieved 8 September 2015 Sperm 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France and Britain for Control of the Southern Whale Fishery 1785 1825 The University of Massachusetts Press ISBN 978 0 87023 104 9 Baldwin R Gallagher M amp van Waerebeek K A Review of Cetaceans from Waters off the Arabian Peninsula PDF p 6 Retrieved 15 October 2008 The Wreck of the Whaleship Essex BBC Retrieved 11 October 2008 Davis L Gallman R amp Gleiter K 1997 In Pursuit of Leviathan Technology Institutions Productivity and Profits in American Whaling 1816 1906 National Bureau of Economic Research Series on Long Term Factors in Economic Dev University of Chicago Press p 135 ISBN 978 0 226 13789 6 Over 680 000 officially reported at Whaling Statistics Archived from the original on 15 October 2019 Retrieved 15 October 2008 In addition studies have found that official reports understated USSR catches by at least 89 000 Sperm Whale Physeter macrocephalus California Oregon Washington Stock PDF Retrieved 16 October 2008 Furthermore other countries such as Japan have been found to have understated catches The RMS A Question of Confidence Manipulations and Falsifications in Whaling PDF Archived from the original PDF on 7 October 2008 Retrieved 16 October 2008 Lavery Trish L Ben Roudnew Peter Gill Justin Seymour Laurent Seuront Genevieve Johnson James G Mitchell amp Victor Smetacek 2010 Iron defecation by sperm whales stimulates carbon export in the Southern Ocean Proceedings of the Royal Society B 277 1699 3527 3531 doi 10 1098 rspb 2010 0863 PMC 2982231 PMID 20554546 Whitehead pp 360 362 Whitehead pp 362 368 Sperm whale Physeter catodon species profile Environmental Conservation Online System United States Fish and Wildlife Service 16 November 2010 a b c d Appendix I and Appendix II of the Convention on the Conservation of Migratory Species of Wild Animals CMS As amended by the Conference of the Parties in 1985 1988 1991 1994 1997 1999 2002 2005 and 2008 Effective 5 March 2009 Museum of New Zealand Te Papa Tongarewa Collections Online Search Rei puta Archived from the original on 4 November 2012 Arno A 2005 Cobo and tabua in Fiji Two forms of cultural currency in an economy of sentiment American Ethnologist 32 1 46 62 doi 10 1525 ae 2005 32 1 46 INIST 16581746 Ratzel Friedrich 1896 Dress and Weapons of the Melanesians Ornament The History of Mankind London MacMillan Retrieved 21 October 2009 Constantine R Folklore and Legends p 449 in Perrin Van Doren Carl 1921 Chapter 3 Romances of Adventure Section 2 Herman Melville The American Novel Bartleby com Retrieved 19 October 2008 a b Zwart H 2000 What is a Whale Moby Dick marine science and the sublime PDF Erzahlen und Moral Narrativitat Im Spannungsfeld von Ethik und Asthetik Tubingen Attempo 185 214 Archived from the original PDF on 20 March 2009 Edwards B The Playful Learnings PDF Australasian Journal of American Studies 25 1 1 13 9 Archived from the original PDF on 20 July 2008 Sperm whale designated Connecticut state animal Cetacean Times 1 3 May 1975 p 6 The State Animal State of Connecticut Sites Seals and Symbols State of Connecticut Archived from the original on 1 September 2011 Retrieved 26 December 2010 Reproduced from the Connecticut State Register amp Manual Sperm whales Whaletrips 26 May 2016 Retrieved 11 December 2022 Whale and dolphin watching in the Azores Wildlife Extra Retrieved 26 September 2008 Whale Watching Dominica Archived from the original on 27 January 2010 Retrieved 26 September 2008 The Dominica Sperm Whale Project Retrieved 25 January 2016 Whales are starving their stomachs full of our plastic waste Philip Hoare The Guardian 30 March 2016 The Times News Google News Archive Search news google com Jacobsen Jeff K Massey Liam Gulland Frances May 2010 Fatal ingestion of floating net debris by two sperm whales Physeter macrocephalus Marine Pollution Bulletin 60 5 765 767 Bibcode 2010MarPB 60 765J doi 10 1016 j marpolbul 2010 03 008 PMID 20381092 Further readingWhitehead H 2003 Sperm Whales Social Evolution in the Ocean Chicago University of Chicago Press p 4 ISBN 978 0 226 89518 5 Perrin William F Wursig Bernd Thewissen J G M eds 2002 Encyclopedia of Marine Mammals San Diego Calif Academic Press ISBN 978 0 12 551340 1 Carwardine Hoyt Fordyce amp Gill 1998 Whales amp Dolphins The Ultimate Guide to Marine Mammals London HarperCollins ISBN 978 0 00 220105 6 Heptner V G Nasimovich A A Bannikov Andrei Grigorevich Hoffmann Robert S Mammals of the Soviet Union Volume II part 3 1996 Washington D C Smithsonian Institution Libraries and National Science FoundationExternal links nbsp Wikimedia Commons has media related to Physeter macrocephalus nbsp Wikispecies has information related to Physeter macrocephalus nbsp Wikisource has the text of the 1911 Encyclopaedia Britannica article Sperm Whale The Dominica Sperm Whale Project a long term scientific research program focusing on the behaviour of sperm whale units Spermaceti in candles 22 July 2007 Society for Marine Mammalogy Sperm Whale Fact Sheet US National Marine Fisheries Service Sperm Whale web page 70South information on the sperm whale Physty stranded sperm whale nursed back to health and released in 1981 ARKive Photographs video Whale Trackers An online documentary film exploring the sperm whales in the Mediterranean Sea Convention on Migratory Species page on the sperm whale Website of the Memorandum of Understanding for the Conservation of Cetaceans and Their Habitats in the Pacific Islands Region Official website of the Agreement on the Conservation of Cetaceans in the Black Sea Mediterranean Sea and Contiguous Atlantic Area Retroposon analysis of major cetacean lineages The monophyly of toothed whales and the paraphyly of river dolphins 19 June 2001 Voices in the Sea sounds of the sperm whale Sperm whales quickly learned to avoid humans who were hunting them in the 19th century scientists say ABC News 16 March 2021 Retrieved from https en wikipedia org w index php title Sperm whale amp oldid 1219675412, wikipedia, wiki, book, books, library,

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