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NOTE: This article uses the English pluralisation, "octopuses". This is correct and intentional, please maintain it. Other variants are discussed in the == Etymology and pluralisation == section.

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An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a beaked mouth at the centre point of the eight limbs. An octopus can radically deform its shape, enabling it to squeeze through small gaps. They trail their appendages behind them as they swim backwards. The siphon is used for respiration and locomotion (by water jet propulsion). Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse invertebrates.

Octopuses inhabit various ocean habitats, including coral reefs, pelagic waters, and the seabed; some live in the intertidal zone and others at abyssal depths. Most species grow quickly, mature early, and are short-lived. In most species, the male uses a specially-adapted arm to deliver sperm directly into the female's mantle cavity, after which he becomes senescent and dies, while the female deposits fertilised eggs in a den and cares for them until they hatch, after which she also dies. They are predators and hunt crustaceans, bivalves, gastropods and fish. Strategies to defend themselves against their own predators include expelling ink, camouflage, and threat displays, the ability to jet quickly through the water and hide, and deceit. All octopuses are venomous, but only the blue-ringed octopuses are known to be deadly to humans.

Octopuses appear in mythology as sea monsters such as the kraken of Norway and the Akkorokamui of the Ainu, and possibly the Gorgons of ancient Greece. A battle with an octopus appears in Victor Hugo's book Toilers of the Sea. Octopuses appear in Japanese shunga erotic art. They are eaten and considered a delicacy by humans in many parts of the world, especially the Mediterranean and Asia.

==Etymology and pluralisation==<!--Avoid changing. Used in links to subsection.-->

The scientific Latin term was derived from Ancient Greek (), a compound form of (, 'eight') and (, 'foot'), itself a variant form of , a word used for example by Alexander of Tralles ( – ).

The standard pluralised form of octopus in English is octopuses; the Ancient Greek plural , (), has also been used historically. The alternative plural octopi is usually considered etymologically incorrect, because it wrongly assumes that octopus is a Latin second-declension noun or adjective when, in either Greek or Latin, it is a third-declension noun. Historically, the first plural to commonly appear in English language sources, in the early 19th century, is the Latinate form octopi, followed by the English form octopuses in the latter half of the same century. The Hellenic plural is roughly contemporary in usage, although it is also the rarest.

A Dictionary of Modern English Usage by H. W. Fowler states that the only acceptable plural in English is octopuses, that octopi is misconceived, and octopodes pedantic; the last is nonetheless used frequently enough to be acknowledged by the descriptivist Merriam-Webster 11th Collegiate Dictionary and Webster's New World College Dictionary. The Oxford English Dictionary lists octopuses, octopi, and octopodes, in that order, reflecting frequency of use, calling octopodes rare and noting that octopi is based on a misunderstanding. The New Oxford American Dictionary (3rd Edition, 2010) lists octopuses as the only acceptable pluralisation, and indicates that octopodes is occasionally used, but that octopi is incorrect.

Anatomy and physiology

Size

thumb|A [[giant Pacific octopus at Echizen Matsushima Aquarium, Japan|alt=Captured specimen of a giant octopus]]

The giant Pacific octopus (Enteroctopus dofleini) is often cited as the largest octopus species. Adults usually weigh , with an arm span of up to . The largest specimen of this species to be scientifically documented reached a live mass of . Much larger sizes have been claimed: one specimen was recorded as with an arm span of . A carcass of the seven-arm octopus, Haliphron atlanticus, weighed and was estimated to have had a live mass of . The smallest species is Octopus wolfi, which is around and weighs less than .

External characteristics

The octopus has an elongated body that is bilaterally symmetrical along its dorso-ventral (back to belly) axis; the head and foot are on the ventral side, but act as the anterior (front). The head contains both the mouth and the brain. The two rear appendages are generally used to walk on the sea floor, while the other six are used to forage for food. The bulbous and hollow mantle is fused to the back of the head and contains most of the vital organs.

thumb|400px|left|alt=Schematic of external anatomy|Diagram of octopus from side, with gills, funnel, eye, [[ocellus (eyespot), web, arms, suckers, hectocotylus and ligula labelled.]]

The skin consists of a thin epidermis with mucous cells and sensory cells and a fibrous dermis made of collagen and containing various cells that allow colour change.

The interior surfaces of the arms are covered with circular, adhesive suckers. The suckers allow the octopus to secure itself in place or to handle objects. Each sucker is typically circular and bowl-like and has two distinct parts: an outer disc-shaped infundibulum and an inner cup-like acetabulum, both of which are thick muscles covered in connective tissue. A chitinous cuticle lines the outer surface. When a sucker attaches to a surface, the orifice between the two structures is sealed and the infundibulum flattens. Muscle contractions allow for attachment and detachment.

thumb |A finned [[Grimpoteuthis species with its atypical octopus body plan|alt=A stubby round sea-creature with short ear-like fins]]

The cranium has two cartilaginous capsules each containing one large eye, which resembles those of fish. The cornea is formed from a translucent epidermal layer; the slit-shaped pupil forms a hole in the iris just behind the cornea. The lens hangs behind the pupil; photoreceptive retinal cells line the back. The pupil can expand and contract; a retinal pigment screens incident light in bright conditions. In cold conditions with low oxygen levels, haemocyanin transports oxygen more efficiently than haemoglobin. Extensive connective tissue lattices support the respiratory muscles and allow them to inflate the respiratory chamber. Respiration can also play a role in locomotion, as an octopus can propel its body shooting water out of the siphon.

Digestion and excretion

The digestive system begins with the buccal mass which consists of the mouth with the beak, the pharynx, radula and salivary glands. One part is localised in the brain, contained in a cartilaginous capsule. Two-thirds of the neurons are in the nerve cords of its arms. This allows their arms to perform actions with a degree of independence. Learning mainly occurs in the brain, while arms make decisions independently when supplied with information. A severed arm can still move and respond to stimuli. Unlike in many other animals, including other mollusks, the movement of octopuses and their relatives are not organised in their brains via internal somatotopic maps of their bodies. Octopuses have the same jumping genes that are active in the human brain, implying an evolutionary convergence at molecular level.

thumb|left|Eye of [[common octopus|alt=Close up of an octopus showing its eye and an arm with suckers]]

Like other cephalopods, octopuses have camera-like eyes. Opsins in the skin respond to different wavelengths of light and help the animals choose a colouration that matches the surroundings and camouflages them; chromatophores in the skin can respond to light independently of the eyes. An alternative hypothesis<!--Stubbs et al--> is that cephalopod eyes in species that only have a single photoreceptor protein may use chromatic aberration to turn monochromatic vision into colour vision, though this lowers image quality. This would explain pupils shaped like the letter "U", the letter "W", or a dumbbell, as well as the need for colourful mating displays.

Attached to the optic capsules are two organs called statocysts (sac-like structures containing a mineralised mass and sensitive hairs), that allow the octopus to sense the orientation of its body, relative to both gravity and time (angular acceleration). An autonomic response keeps the octopus's eyes oriented so that the pupil is always horizontal.

Octopuses have an excellent somatosensory system. Their suction cups are equipped with chemoreceptors so they can taste what they touch. Octopus arms move easily because the sensors recognise octopus skin and prevent self-attachment. Octopuses appear to have poor proprioceptive sense and must see their arms to keep track of their position.

Ink sac

The ink sac is located under the digestive gland. A gland attached to the sac produces the ink, and the sac holds it. The sac is close enough to the funnel for the octopus to shoot out the ink with a water jet. As the animal begins to shoot, the ink passes through glands which mix it with mucus and it leaves the funnel as a thick, dark blob, which helps the animal to escape from a predator. Cirrate octopuses usually lack the ink sac. The male has a specialised arm called a hectocotylus which it uses to transfer spermatophores (packets of sperm) into the female's mantle cavity. A complex hydraulic mechanism releases the sperm from the spermatophore. Fertilised octopus eggs are laid as strings within a shelter.

Lifespan

Octopuses have short lifespans, living up to four years. Senescence is triggered by the optic glands and experimental removal of them after spawning was found to extend their lifecycle and activity.

Distribution and habitat

thumb|Octopus cyanea in Kona, Hawaii|alt=An octopus nearly hidden in a crack in some coral

Octopuses inhabit every ocean, with species adapted to many habitats. As juveniles, common octopuses inhabit shallow tide pools. The Hawaiian day octopus (Octopus cyanea) lives on coral reefs, while argonauts float in pelagic waters. Abdopus aculeatus is a near-shore species and can be found in seagrass beds. Some species can survive in deeper environments. The spoon-armed octopus (Bathypolypus arcticus) can live deep, and Vulcanoctopus hydrothermalis lives in depths of around hydrothermal vents. No species are known to live in fresh water.

The cirrate species are often free-swimming and live in deep-water habitats. Although several species live at bathyal and abyssal depths, only a single indisputable record documents their presence in the hadal zone; a species of Grimpoteuthis (dumbo octopus) photographed at .

Behaviour and ecology

Octopuses are mostly solitary The Larger Pacific striped octopus has been described as particularly social, living in groups of up to 40. Octopuses hide in dens, which are typically crevices in rocky or other hard structures, including man-made ones. Small species may use abandoned shells and bottles. They are not migratory. On rare occasions, octopuses hunt cooperatively with other species, with fish as their partners. They regulate the species composition of the hunting group and the behavior of their partners by punching them.

Feeding

thumb|left|[[Amphioctopus marginatus|Veined octopus eating a crab|alt=An octopus in an open seashell on a sandy surface, surrounding a small crab with the suckers on its arms]]

Octopuses are generally predatory and feed on prey such as crustaceans, bivalves, gastropods, fish, and other cephalopods, including members of the same species. Major items in the diet of the giant Pacific octopus include bivalves such as the cockle Clinocardium nuttallii, clams and scallops and crustaceans such as crabs. It typically rejects moon snails because they are too large; limpets, rock scallops, chitons and abalone, because they are too securely fixed to the rock.

Octopuses typically locate prey by feeling through their environment;

Some species have other modes of feeding. Grimpoteuthis either lacks or has a small radula and swallows prey whole.

Locomotion

thumb|right|Octopuses swim with their arms trailing behind.|alt=An octopus swimming with its round body to the front, its arms forming a streamlined tube behind

Octopuses mainly move about by relatively slow crawling with some swimming in a head-first position. Jet propulsion or backward swimming, is their fastest means of locomotion, while crawling is slowest. While crawling, the suckers adhere and detach from the substrate as the animal hauls itself forward with its powerful arm muscles. This form of locomotion allows these octopuses to move quickly away from a potential predator without being recognised. Maze and problem-solving experiments have shown evidence of a memory system that can store both short- and long-term memory. In laboratory experiments, octopuses can readily be trained to distinguish between different shapes and patterns. They have been reported to practise observational learning, although the validity of these findings is contested. Octopuses have also been observed in what has been described as play: including moving around a bottle by jetting water at it. Octopuses often break out of aquariums and sometimes into others in search of food. Interpretation of evidence has been used to suggest that octopuses have sentience and can feel pain.

Camouflage and colour change

thumb|Video of Octopus cyanea moving and changing its colour, shape, and texture|alt=A video of an octopus changing its appearance

Octopuses use camouflage to hunt and to avoid predators. To do this, they use specialised skin cells that change colour. Chromatophores contain yellow, orange, red, brown, or black pigments; most species have three of these colours, while some have two or four. Other colour-changing cells are reflective iridophores and white leucophores. This colour-changing ability is also used to communicate with or warn other octopuses.

Octopuses can create distracting patterns with waves of dark colouration across the body, a display known as the "passing cloud". Muscles in the skin change the texture of the mantle to achieve greater camouflage. In some species, the mantle can take on the bumpy appearance of algae-covered rocks. Diurnal, shallow water octopuses have more complex skin than their nocturnal and deep-sea counterparts. In the latter species, skin anatomy is limited to one colour or pattern.

Defence

thumb|left|upright|[[Aposematism|Warning display of greater blue-ringed octopus (Hapalochlaena lunulata)|alt=An octopus among coral displaying conspicuous rings of turquoise outlined in black against a sandy background]]

Aside from humans, octopuses are prey for fishes, seabirds, sea otters, pinnipeds, cetaceans, and other cephalopods. The blue rings of the venomous blue-ringed octopus are hidden in muscular skin folds which contract when the animal is threatened, revealing the iridescent warning. The Atlantic white-spotted octopus (Callistoctopus macropus) becomes redder with bright white spots in a deimatic display. Displays are often reinforced by stretching out the animal's arms, fins or web to make it look as big and threatening as possible. When severed by a predator, some octopuses can detach their arm,

Pathogens and parasites

Cephalopods are known to be the intermediate or final hosts of various parasitic cestodes, nematodes and copepods; 150 species of protistan and metazoan parasites are recognised. The Dicyemidae are a family of tiny worms found in the renal appendages of many species; it is unclear whether they are parasitic or endosymbionts. Coccidians in the genus Aggregata living in the gut cause serious illness in the host. Octopuses have an innate immune system; their haemocytes locate the foreign invader and attack it via phagocytosis, encapsulation, infiltration, or cytotoxicity. The haemocytes also contribute to healing injures. A gram-negative bacterium, Vibrio lentus, can cause skin lesions, exposure of muscle and sometimes death.

Evolution

The scientific name Octopoda was first given as the order of octopuses in 1818 by English biologist William Elford Leach, who classified them as Octopoida the previous year. More recent evidence suggests cirrates are the most basal species, not a unique clade.

Fossil history and phylogeny

thumb|The octopuses evolved from the [[Muensterelloidea (fossil pictured) in the Jurassic period. The octopuses arose from the Muensterelloidea within the Vampyropoda in the Jurassic. The earliest octopus likely lived near the sea floor (benthic to demersal) in shallow marine environments. Octopuses consist mostly of soft tissue, and so fossils are relatively rare. As soft-bodied cephalopods, they lack the external shell of most molluscs, including other cephalopods like the nautiloids and the (extinct) Ammonoidea. They have eight limbs like other Coleoidea, but lack the extra specialised feeding appendages known as tentacles which are longer and thinner with suckers only at their club-like ends. The vampire squid (Vampyroteuthis) also lacks tentacles but has sensory filaments.

The cladograms are based on Sanchez et al., 2018, who created a molecular phylogeny based on mitochondrial and nuclear DNA marker sequences. The position of the Eledonidae is from Ibáñez et al., 2020, with a similar methodology. Divergence dates are from Kröger et al., 2011 and Fuchs et al., 2019.

The genome of octopuses has also gone through several chromosomal fusions and rearrangements, unlike that of their closest relative the vampire squid, whose chromosomal structure is more basal and squid-like. The octopus genome is unremarkably bilaterian except for large developments of two gene families: protocadherins, which regulate the development of neurons; and the C2H2 zinc-finger transcription factors. Many novel genes in both cephalopods generally and octopus specifically manifest in the animals' skin, suckers, and nervous system. The Akkorokamui is a gigantic octopus-like monster from Ainu folklore, worshipped in Shinto.<!---->

In the Asuka-era Japanese legend Taishokan, a female diver battles an octopus to recover a stolen jewel, which became the inspiration for woodblock printings. Similarly, in the 1973 novel Gravity's Rainbow an octopus named Grigori attacks a woman on the beach. A battle with an octopus plays a significant role in Victor Hugo's 1866 book Travailleurs de la mer (Toilers of the Sea). The octopus continues to be depicted as antagonistic in films such as Wake of the Red Witch (1948). All species are venomous, but only blue-ringed octopuses have venom that is lethal to humans. Blue-ringed octopuses rank amongst the most dangerous marine animals; their bites are reported each year across the animals' range from Australia to the eastern Indo-Pacific Ocean. They bite only when provoked or accidentally touched; bites are small and usually painless. The venom appears to be able to penetrate the skin without a puncture, given prolonged contact. It contains tetrodotoxin, which causes paralysis by blocking the transmission of nerve impulses to the muscles. This causes death by respiratory failure leading to cerebral anoxia. No antidote is known, but if breathing can be kept going artificially, patients recover within 24 hours. Bites have been recorded from captive octopuses of other species; they leave temporary swellings. The world catch peaked in 2007 at 380,000 tons, and had fallen by a tenth by 2012. Methods to capture octopuses include pots, trapping, trawling, snaring, drift fishing, spearing, hooking and catching by hands. Attempts to farm octopuses commercially are controversial.

Octopus is eaten in many cultures, such as those on the Mediterranean and Asian coasts. The arms and other body parts are prepared in ways that vary by species and geography. Live octopuses or their wriggling pieces are consumed as san-nakji in Korean cuisine. If not prepared properly, however, the severed arms can choke the diner with their suction cups, causing at least one death in 2010. Animal welfare groups have objected to the live consumption of octopuses on the basis that they can experience pain.

Science and technology

In classical Greece, Aristotle (384–322 BC) commented on their colour-changing abilities, both for camouflage and for signalling, in his Historia animalium: "The octopus&nbsp;... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed." Aristotle noted that the octopus had a hectocotyl arm and suggested it might be used in reproduction. This claim was widely ignored until the 19th century. It was described in 1829 by the French zoologist Georges Cuvier, who supposed it to be a parasitic worm, naming it as a new species, Hectocotylus octopodis. Other zoologists thought it a spermatophore; the German zoologist Heinrich Müller believed it was "designed" to detach during copulation. In 1856, the Danish zoologist Japetus Steenstrup demonstrated that it is used to transfer sperm, and only rarely detaches.

thumb|Flexible [[biomimetic 'Octopus' robotics arm. The BioRobotics Institute, Scuola Superiore Sant'Anna, Pisa, 2011]]

Octopuses offer many possibilities in biological research; the California two-spot octopus had its genome sequenced, allowing exploration of its molecular adaptations. Having independently evolved mammal-like intelligence, octopuses were compared by the philosopher Peter Godfrey-Smith, who studied the nature of intelligence, to hypothetical intelligent extraterrestrials. Their intelligence and flexible bodies enable them to escape from supposedly secure tanks in public aquariums.

Due to their intelligence, many argue that octopuses should be given protections when used for experiments. In the UK from 1993 to 2012, the common octopus (Octopus vulgaris) was the only invertebrate protected under the Animals (Scientific Procedures) Act 1986. In 2012, this legislation was extended to include all cephalopods in accordance with a general EU directive.

Some robotics research is exploring biomimicry of octopus features. Octopus arms can move and sense largely autonomously without intervention from the animal's central nervous system. In 2015 a team in Italy built soft-bodied robots able to crawl and swim, requiring only minimal computation. In 2017, a German company made an arm with a soft pneumatically controlled silicone gripper fitted with two rows of suckers. It was able to grasp objects such as a metal tube, a magazine, or a ball, and to fill a glass by pouring water from a bottle.

See also

Notes

References

  • Octopuses – Overview at the Encyclopedia of Life
  • Octopoda at the Tree of Life Web Project
  • "Can We Really Be Friends with an Octopus?" at Hakai Magazine, January 11, 2022