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[Vanya Lamunyon]

Asquid (pl.: squid) is a mollusc with an elongated soft body, large eyes, eight arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida. Though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting these criteria. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.

Squid diverged from other cephalopods during the Jurassic and occupy a similar role to teleost fish as open water predators of similar size and behaviour. They play an important role in the open water food web. The two long tentacles are used to grab prey and the eight arms to hold and control it. The beak then cuts the food into suitable size chunks for swallowing. Squid are rapid swimmers, moving by jet propulsion, and largely locate their prey by sight. They are among the most intelligent of invertebrates, with groups of Humboldt squid having been observed hunting cooperatively. They are preyed on by sharks, other fish, sea birds, seals and cetaceans, particularly sperm whales.

Squid can change colour for camouflage and signalling. Some species are bioluminescent, using their light for counter-illumination camouflage, while many species can eject a cloud of ink to distract predators.

Squid are used for human consumption with commercial fisheries in Japan, the Mediterranean, the southwestern Atlantic, the eastern Pacific and elsewhere. They are used in cuisines around the world, often known as "calamari". Squid have featured in literature since classical times, especially in tales of giant squid and sea monsters.

Squid are members of the class Cephalopoda, subclass Coleoidea. The squid orders Myopsida and Oegopsida are in the superorder Decapodiformes (from the Greek for "ten-legged"). Two other orders of decapodiform cephalopods are also called squid, although they are taxonomically distinct from squids and differ recognizably in their gross anatomical features. They are the bobtail squid of order Sepiolida and the ram's horn squid of the monotypic order Spirulida. The vampire squid (Vampyroteuthis infernalis), however, is more closely related to the octopus than to any squid.[2]

The cladogram, not fully resolved, is based on Sanchez et al., 2018.[2] Their molecular phylogeny used mitochondrial and nuclear DNA marker sequences; they comment that a robust phylogeny "has proven very challenging to obtain". If it is accepted that Sepiidae cuttlefish are a kind of squid, then the squids, excluding the vampire squid, form a clade as illustrated.[2] Orders are shown in boldface; all the families not included in those orders are in the paraphyletic order "Oegopsida", except Sepiadariidae and Sepiidae that are in the paraphyletic order "Sepiida",

Crown coleoids (the common ancestor of octopuses and squid) diverged in the late Paleozoic (Mississippian), according to fossils of Syllipsimopodi, an early relative of vampire squids and octopuses.[3] True squid diverged during the Jurassic, but many squid families appeared in or after the Cretaceous.[4] Both the coleoids and the teleost fish were involved in much adaptive radiation at this time, and the two modern groups resemble each other in size, ecology, habitat, morphology and behaviour, however some fish moved into fresh water while the coleoids remained in marine environments.[5]

The ancestral coleoid was probably nautiloid-like with a strait septate shell that became immersed in the mantle and was used for buoyancy control. Four lines diverged from this, Spirulida (with one living member), the cuttlefishes, the squids and the octopuses. Squid have differentiated from the ancestral mollusc such that the body plan has been condensed antero-posteriorly and extended dorso-ventrally. What may have been the foot of the ancestor is modified into a complex set of appendages around the mouth. The sense organs are highly developed and include advanced eyes similar to those of vertebrates.[5]

The ancestral shell has been lost, with only an internal gladius, or pen, remaining. The pen, made of a chitin-like material,[5][6] is a feather-shaped internal structure that supports the squid's mantle and serves as a site for muscle attachment. The cuttlebone or sepion of the Sepiidae is calcareous and appears to have evolved afresh in the Tertiary.[7]

Squid are soft-bodied molluscs whose forms evolved to adopt an active predatory lifestyle. The head and foot of the squid are at one end of a long body, and this end is functionally anterior, leading the animal as it moves through the water. A set of eight arms and two distinctive tentacles surround the mouth; each appendage takes the form of a muscular hydrostat and is flexible and prehensile, usually bearing disc-like suckers.[5]

The suckers may lie directly on the arm or be stalked. Their rims are stiffened with chitin and may contain minute toothlike denticles. These features, as well as strong musculature, and a small ganglion beneath each sucker to allow individual control, provide a very powerful adhesion to grip prey. Hooks are present on the arms and tentacles in some species, but their function is unclear.[8] The two tentacles are much longer than the arms and are retractile. Suckers are limited to the spatulate tip of the tentacle, known as the manus.[5]

The main body mass is enclosed in the mantle, which has a swimming fin along each side. These fins are not the main source of locomotion in most species. The mantle wall is heavily muscled and internal. The visceral mass, which is covered by a thin, membranous epidermis, forms a cone-shaped posterior region known as the "visceral hump". The mollusc shell is reduced to an internal, longitudinal chitinous "pen" in the functionally dorsal part of the animal; the pen acts to stiffen the squid and provides attachments for muscles.[5]

On the functionally ventral part of the body is an opening to the mantle cavity, which contains the gills (ctenidia) and openings from the excretory, digestive and reproductive systems. An inhalant siphon behind the funnel draws water into the mantle cavity via a valve. The squid uses the funnel for locomotion via precise jet propulsion.[9] In this form of locomotion, water is sucked into the mantle cavity and expelled out of the funnel in a fast, strong jet. The direction of travel is varied by the orientation of the funnel.[5] Squid are strong swimmers and certain species can "fly" for short distances out of the water.[10]

Squid make use of different kinds of camouflage, namely active camouflage for background matching (in shallow water) and counter-illumination. This helps to protect them from their predators and allows them to approach their prey.[11][12]

The skin is covered in controllable chromatophores of different colours, enabling the squid to match its coloration to its surroundings.[11][13] The play of colours may in addition distract prey from the squid's approaching tentacles.[14] The skin also contains light reflectors called iridophores and leucophores that, when activated, in milliseconds create changeable skin patterns of polarized light.[15][16] Such skin camouflage may serve various functions, such as communication with nearby squid, prey detection, navigation, and orientation during hunting or seeking shelter.[15] Neural control of the iridophores enabling rapid changes in skin iridescence appears to be regulated by a cholinergic process affecting reflectin proteins.[16]

Some mesopelagic squid such as the firefly squid (Watasenia scintillans) and the midwater squid (Abralia veranyi) use counter-illumination camouflage, generating light to match the downwelling light from the ocean surface.[12][17][18] This creates the effect of countershading, making the underside lighter than the upperside.[12]

Counter-illumination is also used by the Hawaiian bobtail squid (Euprymna scolopes), which has symbiotic bacteria (Aliivibrio fischeri) that produce light to help the squid avoid nocturnal predators.[19] This light shines through the squid's skin on its underside and is generated by a large and complex two-lobed light organ inside the squid's mantle cavity. From there, it escapes downwards, some of it travelling directly, some coming off a reflector at the top of the organ (dorsal side). Below there is a kind of iris, which has branches (diverticula) of its ink sac, with a lens below that; both the reflector and lens are derived from mesoderm. The squid controls light production by changing the shape of its iris or adjusting the strength of yellow filters on its underside, which presumably change the balance of wavelengths emitted.[17] Light production shows a correlation with intensity of down-welling light, but it is about one third as bright; the squid can track repeated changes in brightness. Because the Hawaiian bobtail squid hides in sand during the day to avoid predators, it does not use counter-illumination during daylight hours.[17]

Squid distract attacking predators by ejecting a cloud of ink, giving themselves an opportunity to escape.[20][21] The ink gland and its associated ink sac empties into the rectum close to the anus, allowing the squid to rapidly discharge black ink into the mantle cavity and surrounding water.[8] The ink is a suspension of melanin particles and quickly disperses to form a dark cloud that obscures the escape manoeuvres of the squid. Predatory fish may also be deterred by the alkaloid nature of the discharge which may interfere with their chemoreceptors.[5]

Cephalopods have the most highly developed nervous systems among invertebrates. Squids have a complex brain in the form of a nerve ring encircling the oesophagus, enclosed in a cartilaginous cranium. Paired cerebral ganglia above the oesophagus receive sensory information from the eyes and statocysts, and further ganglia below control the muscles of the mouth, foot, mantle and viscera. Giant axons up to 1 mm (0.04 in) in diameter convey nerve messages with great rapidity to the circular muscles of the mantle wall, allowing a synchronous, powerful contraction and maximum speed in the jet propulsion system.[5]

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