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True squid are molluscs with an elongated soft body, large eyes, eight
arms Arms or ARMS may refer to: *Arm or arms, the upper limbs of the body Arm, Arms, or ARMS may also refer to: People * Ida A. T. Arms (1856–1931), American missionary-educator, temperance leader Coat of arms or weapons *Armaments or weapons **Fi ...
, and two
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work ma ...
s in the superorder
Decapodiformes Decapodiformes is a superorder of Cephalopoda comprising all cephalopod species with ten limbs, specifically eight short arms and two long tentacles. It is hypothesized that the ancestral coleoid had five identical pairs of limbs, and that one ...
, though many other molluscs within the broader
Neocoleoidea Neocoleoidea is a large group of marine cephalopods. This cohort contains two extant groups: Decapodiformes (squid, cuttlefish, and relatives) and Octopodiformes (octopuses and the vampire squid). Species within this group exist in all major hab ...
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 ''Gladius'' () is a Latin word meaning "sword" (of any type), but in its narrow sense it refers to the sword of ancient Roman foot soldiers. Early ancient Roman swords were similar to those of the Greeks, called '' xiphe'' (plural; singular ''xi ...
or pen, made of chitin. Squid diverged from other cephalopods during the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
and occupy a similar role to
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tele ...
fish as open water
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s 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 Jet propulsion is the propulsion of an object in one direction, produced by ejecting a jet of fluid in the opposite direction. By Newton's third law, the moving body is propelled in the opposite direction to the jet. Reaction engines operatin ...
, and largely locate their prey by sight. They are among the most intelligent of
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordat ...
, with groups of
Humboldt squid The Humboldt squid (''Dosidicus gigas''), also known as jumbo squid or jumbo flying squid (EN), and Pota in Peru or Jibia in Chile (ES) is a large, predatory squid living in the eastern Pacific Ocean. It is the only known species of the genus ''D ...
having been observed hunting cooperatively. They are preyed on by
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s, other fish, sea birds,
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
and cetaceans, particularly sperm whales. Squid can change colour for
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
and
signalling In signal processing, a signal is a function that conveys information about a phenomenon. Any quantity that can vary over space or time can be used as a signal to share messages between observers. The ''IEEE Transactions on Signal Processing'' ...
. Some species are bioluminescent, using their light for
counter-illumination Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength. Marine animals of ...
camouflage, while many species can eject a cloud of
ink Ink is a gel, sol, or solution that contains at least one colorant, such as a dye or pigment, and is used to color a surface to produce an image, text, or design. Ink is used for drawing or writing with a pen, brush, reed pen, or quill. Thicker ...
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 is eaten in many cuisines; in English, the culinary name calamari is often used for squid dishes.'' Oxford English Dictionary'', 3rd edition, 2002''s.v.''/ref> There are many ways to prepare and cook squid. Fried squid is common in the ...
". Squid have featured in literature since classical times, especially in tales of giant squid and
sea monster Sea monsters are beings from folklore believed to dwell in the sea and often imagined to be of immense size. Marine monsters can take many forms, including sea dragons, sea serpents, or tentacled beasts. They can be slimy and scaly and are of ...
s.


Taxonomy and phylogeny

Squid are members of the class Cephalopoda, subclass
Coleoidea Subclass Coleoidea, or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less" (i.e., octopuses, squid and cuttlefish). Unlike its extant sister group, Nautiloidea, whos ...
. The squid
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Myopsida Myopsida is one of the four orders of squid. It consists of two families: the monotypic '' Australiteuthis'' and the diverse and commercially important Loliginidae (~50 species). Some taxonomists classify this taxon as a suborder of the order ...
and
Oegopsida Oegopsida is one of the two orders of squid in the superorder Decapodiformes, in the class Cephalopoda. Together with the Myopsina, it was formerly considered to be a suborder of the order Teuthida, in which case it was known as Oegopsina. This ...
are in the superorder
Decapodiformes Decapodiformes is a superorder of Cephalopoda comprising all cephalopod species with ten limbs, specifically eight short arms and two long tentacles. It is hypothesized that the ancestral coleoid had five identical pairs of limbs, and that one ...
(from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
for "ten-legged"). Two other orders of decapodiform cephalopods are also called squid, although they are
taxonomically In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
distinct from squids and differ recognizably in their gross anatomical features. They are the
bobtail squid Bobtail squid (order Sepiolida) are a group of cephalopods closely related to cuttlefish. Bobtail squid tend to have a rounder mantle than cuttlefish and have no cuttlebone. They have eight suckered arms and two tentacles and are generally qui ...
of order
Sepiolida Bobtail squid (order Sepiolida) are a group of cephalopods closely related to cuttlefish. Bobtail squid tend to have a rounder mantle than cuttlefish and have no cuttlebone. They have eight suckered arms and two tentacles and are generally quite ...
and the ram's horn squid of the monotypic order
Spirulida Spirulida is an order of cephalopods comprising one extant species ('' Spirula spirula'') and several extinct taxa. Fossil record * Oldest representative: Carboniferous, though contested: see '' Shimanskya'' * Oldest uncontested representativ ...
. The
vampire squid The vampire squid (''Vampyroteuthis infernalis'', lit. 'vampire squid from hell') is a small cephalopod found throughout temperate and tropical oceans in #Habitat and adaptations, extreme deep sea conditions. The vampire squid uses its Biolumines ...
('' Vampyroteuthis infernalis''), however, is more closely related to the octopus than to any squid. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
, not fully resolved, is based on Sanchez et al., 2018. Their
molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
used mitochondrial and
nuclear DNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. I ...
marker sequences; they comment that a robust
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
"has proven very challenging to obtain". If it is accepted that
Sepiidae Sepiidae is a family of cephalopods in the order Sepiida. Classification *Order Sepiida: cuttlefish **Family Sepiadariidae **Family Sepiidae ***Genus ''Metasepia'' ****''Metasepia pfefferi'', flamboyant cuttlefish ****'' Metasepia tullbergi' ...
cuttlefish are a kind of squid, then the squids, excluding the vampire squid, form a clade as illustrated. 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",


Evolution

Crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
coleoids (the common ancestor of octopuses and squid) diverged in the late
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
( Mississippian), according to fossils of '' Syllipsimopodi'', an early relative of vampire squids and octopuses. True squid diverged during the Jurassic, but many squid families appeared in or after the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
. 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. 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
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
s. The ancestral shell has been lost, with only an internal
gladius ''Gladius'' () is a Latin word meaning "sword" (of any type), but in its narrow sense it refers to the sword of ancient Roman foot soldiers. Early ancient Roman swords were similar to those of the Greeks, called '' xiphe'' (plural; singular ''xi ...
, or pen, remaining. The pen, made of a chitin-like material, is a feather-shaped internal structure that supports the squid's mantle and serves as a site for muscle attachment. The
cuttlebone Cuttlebone, also known as cuttlefish bone, is a hard, brittle internal structure (an internal shell) found in all members of the family Sepiidae, commonly known as cuttlefish, within the cephalopods. In other cephalopod families it is calle ...
or sepion of the Sepiidae is
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an ad ...
and appears to have evolved afresh in the
Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
. File:Rhomboteuthis lehmani (cropped).jpg, Fossil '' Rhomboteuthis'' from the Lower Callovian (, middle Jurassic) of
La Voulte-sur-Rhône La Voulte-sur-Rhône (, literally ''La Voulte on Rhône''; oc, La Vòuta) is a commune in the Ardèche department in southern France. Population Lagerstätte La Voulte-sur-Rhône is a marine Lagerstätte (sedimentary deposit) located in F ...
, France File:Plesioteuthis prisca 01.jpg, Fossil ''
Plesioteuthis ''Plesioteuthis'' is an extinct genus of squids, belonging to the family Plesioteuthididae. This genus was first described in 1859, and its fossils were found in the well-known Solnhofen field in Germany. Species Species within this genus inclu ...
'' from the Tithonian (, upper Jurassic),
Solnhofen Solnhofen is a municipality in the district of Weißenburg-Gunzenhausen in the region of Middle Franconia in the ' of Bavaria in Germany. It is in the Altmühl valley. The local area is famous in geology and palaeontology for Solnhofen limest ...
, Germany


Description

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 A muscular hydrostat is a biological structure found in animals. It is used to manipulate items (including food) or to move its host about and consists mainly of muscles with no skeletal support. It performs its hydraulic movement without fluid in ...
and is flexible and prehensile, usually bearing disc-like suckers. 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. 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. In the mature male, the outer half of one of the left arms is hectocotylised – and ends in a copulatory pad rather than suckers. This is used for depositing a
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
inside the mantle cavity of a female. A ventral part of the foot has been converted into a
funnel A funnel is a tube or pipe that is wide at the top and narrow at the bottom, used for guiding liquid or powder into a small opening. Funnels are usually made of stainless steel, aluminium, glass, or plastic. The material used in its construct ...
through which water exits the mantle cavity. 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. On the functionally ventral part of the body is an opening to the mantle cavity, which contains the
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
s (ctenidia) and openings from the excretory, digestive and
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are als ...
s. 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. 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. Squid are strong swimmers and certain species can "fly" for short distances out of the water.


Camouflage

Squid make use of different kinds of camouflage, namely
active camouflage Active camouflage or adaptive camouflage is camouflage that adapts, often rapidly, to the surroundings of an object such as an animal or military vehicle. In theory, active camouflage could provide perfect concealment from visual detection. Activ ...
for background matching (in shallow water) and counter-illumination. This helps to protect them from their predators and allows them to approach their prey. The skin is covered in controllable
chromatophore Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contrast, ...
s of different colours, enabling the squid to match its coloration to its surroundings. The play of colours may in addition distract prey from the squid's approaching tentacles. The skin also contains light reflectors called iridophores and
leucophore Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contras ...
s that, when activated, in milliseconds create changeable skin patterns of polarized light. Such skin camouflage may serve various functions, such as communication with nearby squid, prey detection, navigation, and orientation during hunting or seeking shelter. Neural control of the iridophores enabling rapid changes in skin iridescence appears to be regulated by a
cholinergic Cholinergic agents are compounds which mimic the action of acetylcholine and/or butyrylcholine. In general, the word " choline" describes the various quaternary ammonium salts containing the ''N'',''N'',''N''-trimethylethanolammonium cati ...
process affecting reflectin
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s. Some
mesopelagic The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins at ...
squid such as the
firefly squid The firefly squid (''Watasenia scintillans''), also commonly known as the sparkling enope squid or hotaru-ika in Japan, is a species of squid in the family Enoploteuthidae. It is the sole species in the monotypic genus ''Watasenia''. These tiny ...
(''Watasenia scintillans'') and the midwater squid (''Abralia veranyi'') use counter-illumination camouflage, generating light to match the downwelling light from the ocean surface. This creates the effect of
countershading Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, a ...
, making the underside lighter than the upperside. 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. 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 Iris most often refers to: *Iris (anatomy), part of the eye *Iris (mythology), a Greek goddess * ''Iris'' (plant), a genus of flowering plants * Iris (color), an ambiguous color term Iris or IRIS may also refer to: Arts and media Fictional ent ...
, which has branches (diverticula) of its
ink sac An ink sac is an anatomical feature that is found in many cephalopod mollusks used to produce the defensive cephalopod ink. With the exception of nocturnal and very deep water cephalopods, all Coleoidea (squid, octopus and cuttlefish) which dwell ...
, 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. 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. File:Chromatophores.jpg, upright=1.25, Controllable
chromatophore Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contrast, ...
s of different colours in the skin of a squid allow it to change its coloration and patterns rapidly, whether for camouflage or signalling. File:Squid Counterillumination.png, Principle of
counter-illumination Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength. Marine animals of ...
camouflage of the firefly squid, '' Watasenia scintillans''. When seen from below by a
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
, the animal's light helps to match its brightness and colour to the sea surface above.


Predator distraction with ink

Squid distract attacking predators by ejecting a cloud of
ink Ink is a gel, sol, or solution that contains at least one colorant, such as a dye or pigment, and is used to color a surface to produce an image, text, or design. Ink is used for drawing or writing with a pen, brush, reed pen, or quill. Thicker ...
, giving themselves an opportunity to escape. 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. The ink is a suspension of
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
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
chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
s.


Nervous system and sense organs

Cephalopods have the most highly developed nervous systems among
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s. Squids have a complex brain in the form of a nerve ring encircling the
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to th ...
, enclosed in a cartilaginous cranium. Paired cerebral ganglia above the oesophagus receive sensory information from the eyes and
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in '' Xenoturbella''. The statocyst c ...
s, and further ganglia below control the muscles of the mouth, foot, mantle and viscera. Giant axons up to 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. The paired eyes, on either side of the head, are housed in capsules fused to the cranium. Their structure is very similar to that of a fish eye, with a globular
lens A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements ...
that has a
depth of focus Depth of focus is a lens optics concept that measures the tolerance of placement of the image plane (the film plane in a camera) in relation to the lens. In a camera, depth of focus indicates the tolerance of the film's displacement within the ca ...
from to infinity. The image is focused by changing the position of the lens, as in a camera or
telescope A telescope is a device used to observe distant objects by their emission, absorption, or reflection of electromagnetic radiation. Originally meaning only an optical instrument using lenses, curved mirrors, or a combination of both to observ ...
, rather than changing the shape of the lens, as in the human eye. Squid adjust to changes in light intensity by expanding and contracting the slit-shaped
pupil The pupil is a black hole located in the center of the Iris (anatomy), iris of the Human eye, eye that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing ...
. Deep sea squids in the family Histioteuthidae have eyes of two different types and orientation. The large left eye is tubular in shape and looks upwards, presumably searching for the silhouettes of animals higher in the
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
. The normally-shaped right eye points forwards and downwards to detect prey. The statocysts are involved in maintaining balance and are analogous to the inner ear of fish. They are housed in cartilaginous capsules on either side of the cranium. They provide the squid with information on its body position in relation to gravity, its orientation, acceleration and rotation, and are able to perceive incoming vibrations. Without the statocysts, the squid cannot maintain equilibrium. Squid appear to have limited hearing, but the head and arms bear lines of hair-cells that are weakly sensitive to water movements and changes in pressure, and are analogous in function to the lateral line system of fish.


Reproductive system

The sexes are separate in squid, there being a single gonad in the posterior part of the body with fertilisation being external, and usually taking place in the mantle cavity of the female. The male has a
testis A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testostero ...
from which sperm pass into a single gonoduct where they are rolled together into a long bundle, or spermatophore. The gonoduct is elongated into a "penis" that extends into the mantle cavity and through which spermatophores are ejected. In shallow water species, the penis is short, and the spermatophore is removed from the mantle cavity by a tentacle of the male, which is specially adapted for the purpose and known as a
hectocotylus A hectocotylus (plural: ''hectocotyli'') is one of the arms of male cephalopods that is specialized to store and transfer spermatophores to the female. Structurally, hectocotyli are muscular hydrostats. Depending on the species, the male may use i ...
, and placed inside the mantle cavity of the female during mating. The female has a large translucent ovary, situated towards the posterior of the visceral mass. From here, eggs travel along the gonocoel, where there are a pair of white nidamental glands, which lie anterior to the gills. Also present are red-spotted accessory nidamental glands containing symbiotic bacteria; both organs are associated with nutrient manufacture and forming shells for the eggs. The gonocoel enters the mantle cavity at the
gonopore A gonopore, sometimes called a gonadopore, is a genital pore in many invertebrates. Hexapods, including insects have a single common gonopore, except mayflies, which have a pair of gonopores. More specifically, in the unmodified female it is t ...
, and in some species, receptacles for storing spermatophores are located nearby, in the mantle wall. In shallow-water species of the continental shelf and
epipelagic The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
or mesopelagic zones, it is frequently one or both of arm pair IV of males that are modified into hectocotyli. However, most deep-sea squid lack hectocotyl arms and have longer penises; Ancistrocheiridae and Cranchiinae are exceptions.
Giant squid The giant squid (''Architeuthis dux'') is a species of deep-ocean dwelling squid in the family Architeuthidae. It can grow to a tremendous size, offering an example of abyssal gigantism: recent estimates put the maximum size at around Trac ...
of the genus ''Architeuthis'' are unusual in that they possess both a large penis and modified arm tips, although whether the latter are used for spermatophore transfer is uncertain. Penis elongation has been observed in the deep-water species '' Onykia ingens''; when erect, the penis may be as long as the mantle, head, and arms combined. As such, deep-water squid have the greatest known penis length relative to body size of all mobile animals, second in the entire animal kingdom only to certain sessile barnacles.


Digestive system

Like all cephalopods, squids are predators and have complex digestive systems. The mouth is equipped with a sharp, horny
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for foo ...
mainly made of chitin and
cross-link In chemistry and biology a cross-link is a bond or a short sequence of bonds that links one polymer chain to another. These links may take the form of covalent bonds or ionic bonds and the polymers can be either synthetic polymers or natural ...
ed proteins, which is used to kill and tear prey into manageable pieces. The beak is very robust, but does not contain minerals, unlike the teeth and jaws of many other organisms; the cross-linked proteins are histidine- and glycine-rich and give the beak a stiffness and hardness greater than most equivalent synthetic organic materials. The stomachs of captured whales often have indigestible squid beaks inside. The mouth contains the radula, the rough tongue common to all
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is estim ...
except
bivalvia Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biva ...
, which is equipped with multiple rows of teeth. In some species, toxic saliva helps to control large prey; when subdued, the food can be torn in pieces by the beak, moved to the oesophagus by the radula, and swallowed. The food bolus is moved along the gut by waves of muscular contractions (
peristalsis Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, whic ...
). The long oesophagus leads to a muscular
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
roughly in the middle of the visceral mass. The
digestive gland The hepatopancreas, digestive gland or midgut gland is an organ of the digestive tract of arthropods and molluscs. It provides the functions which in mammals are provided separately by the liver and pancreas, including the production of digestive e ...
, which is equivalent to a vertebrate liver, diverticulates here, as does the
pancreas The pancreas is an organ of the digestive system and endocrine system of vertebrates. In humans, it is located in the abdomen behind the stomach and functions as a gland. The pancreas is a mixed or heterocrine gland, i.e. it has both an en ...
, and both of these empty into the
caecum The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, to which it is joined). The wo ...
, a pouch-shaped sac where most of the absorption of nutrients takes place. Indigestible food can be passed directly from the stomach to the rectum where it joins the flow from the caecum and is voided through the anus into the mantle cavity. Cephalopods are short-lived, and in mature squid, priority is given to reproduction; the female ''
Onychoteuthis banksii ''Onychoteuthis banksii'', the common clubhook squid, is a species of squid in the family Onychoteuthidae. It is the type species of the genus '' Onychoteuthis''. This species was thought to have a worldwide distribution but with the revision o ...
'' for example, sheds its feeding tentacles on reaching maturity, and becomes flaccid and weak after spawning.


Cardiovascular and excretory systems

The squid mantle cavity is a seawater-filled sac containing three hearts and other organs supporting circulation, respiration, and
excretion Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after ...
. Squid have a main systemic heart that pumps blood around the body as part of the general
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
, and two branchial hearts. The systemic heart consists of three chambers, a lower ventricle and two upper atria, all of which can contract to propel the blood. The branchial hearts pump blood specifically to the gills for oxygenation, before returning it to the systemic heart. The blood contains the
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
-rich protein
hemocyanin Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2) ...
, which is used for oxygen transport at low ocean temperatures and low oxygen concentrations, and makes the oxygenated blood a deep, blue color. As systemic blood returns via two
vena cavae In anatomy, the venae cavae (; singular: vena cava ; ) are two large veins (great vessels) that return deoxygenated blood from the body into the heart. In humans they are the superior vena cava and the inferior vena cava, and both empty into the ...
to the branchial hearts, excretion of
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excreted from the body through the urethra. Cellular ...
,
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
, and waste
solute In chemistry, a solution is a special type of homogeneous mixture composed of two or more substances. In such a mixture, a solute is a substance dissolved in another substance, known as a solvent. If the attractive forces between the solvent ...
s occurs through outpockets (called nephridial appendages) in the vena cavae walls that enable gas exchange and excretion via the mantle cavity seawater.


Buoyancy

Unlike nautiloids which have gas-filled chambers inside their shells which provide buoyancy, and octopuses which live near and rest on the seabed and do not require to be buoyant, many squid have a fluid-filled receptacle, equivalent to the swim bladder of a fish, in the coelom or connective tissue. This reservoir acts as a chemical buoyancy chamber, with the heavy metallic cations typical of seawater replaced by low
molecular-weight The molecular mass (''m'') is the mass of a given molecule: it is measured in daltons (Da or u). Different molecules of the same compound may have different molecular masses because they contain different isotopes of an element. The related quanti ...
ammonium ions, a product of excretion. The small difference in density provides a small contribution to buoyancy per unit volume, so the mechanism requires a large buoyancy chamber to be effective. Since the chamber is filled with liquid, it has the advantage over a swim bladder of not changing significantly in volume with pressure. Glass squids in the family Cranchiidae for example, have an enormous transparent coelom containing ammonium ions and occupying about two-thirds the volume of the animal, allowing it to float at the required depth. About half of the 28 families of squid use this mechanism to solve their buoyancy issues.


Largest and smallest

The majority of squid are no more than long, although the giant squid may reach . The smallest species are probably the benthic pygmy squids '' Idiosepius'', which grow to a mantle length of , and have short bodies and stubby arms. In 1978, sharp, curved claws on the suction cups of squid tentacles cut up the rubber coating on the hull of the USS ''Stein''. The size suggested the largest squid known at the time. In 2003, a large specimen of an abundant but poorly understood species, ''Mesonychoteuthis hamiltoni'' (the
colossal squid The colossal squid (''Mesonychoteuthis hamiltoni'') is part of the family Cranchiidae. It is sometimes called the Antarctic squid or giant cranch squid and is believed to be the largest squid species in terms of mass. It is the only recognized ...
), was discovered. This species may grow to in length, making it the largest invertebrate. In February 2007, a New Zealand fishing vessel caught the largest squid ever documented, weighing and measuring around off the coast of Antarctica. Dissection showed that the eyes, used to detect prey in the deep Southern Ocean, exceeded the size of footballs; these may be among the largest eyes ever to exist in the animal kingdom.


Development

The eggs of squid are large for a mollusc, containing a large amount of yolk to nourish the embryo as it develops directly, without an intervening
veliger A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells. Description The veliger is the characteristic larva of the gastropod, bivalve and scaphopod ...
larval stage. The embryo grows as a disc of cells on top of the yolk. During the gastrulation stage, the margins of the disc grow to surround the yolk, forming a yolk sac, which eventually forms part of the animal's gut. The dorsal side of the disc grows upwards and forms the embryo, with a shell gland on its dorsal surface, gills, mantle and eyes. The arms and funnel develop as part of the foot on the ventral side of the disc. The arms later migrate upwards, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo grows. Some juvenile squid live higher in the water column than do adults. Squids tend to be short-lived; ''
Loligo ''Loligo'' is a genus of squid and one of the most representative and widely distributed groups of myopsid squid. The genus was first described by Jean Baptiste Lamarck in 1798. However, the name had been used earlier than Lamarck (Schneider, ...
'' for example lives from one to three years according to species, typically dying soon after spawning. In a well-studied bioluminescent species, the Hawaiian bobtail squid, a special light organ in the squid's mantle is rapidly colonized with ''
Aliivibrio fischeri ''Aliivibrio fischeri'' (also called ''Vibrio fischeri'') is a Gram-negative, rod-shaped bacterium found globally in marine environments. This species has bioluminescent properties, and is found predominantly in symbiosis with various marine anim ...
'' bacteria within hours of hatching. This light-organ colonization requires this particular bacterial species for a symbiotic relationship; no colonization occurs in the absence of ''A. fischeri''. Colonization occurs in a horizontal manner, such that the hosts acquires its bacterial partners from the environment. The symbiosis is obligate for the squid, but facultative for the bacteria. Once the bacteria enter the squid, they colonize interior
epithelial cells Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercell ...
in the light organ, living in
crypts A crypt (from Latin '' crypta'' "vault") is a stone chamber beneath the floor of a church or other building. It typically contains coffins, sarcophagi, or religious relics. Originally, crypts were typically found below the main apse of a chu ...
with complex microvilli protrusions. The bacteria also interact with hemocytes, macrophage-like blood cells that migrate between epithelial cells, but the mechanism and function of this process is not well understood. Bioluminescence reaches its highest levels during the early evening hours and bottoms out before dawn; this occurs because at the end of each day, the contents of the squid's crypts are expelled into the surrounding environment. Approximately 95% of the bacteria are voided each morning before the bacterial population builds up again by nightfall.


Behaviour


Locomotion

Squid can move about in several different ways. Slow movement is achieved by a gentle undulation of the muscular lateral fins on either side of the trunk which drives the animal forward. A more common means of locomotion providing sustained movement is achieved using jetting, during which contraction of the muscular wall of the mantle cavity provides jet propulsion. Slow jetting is used for ordinary locomotion, and ventilation of the gills is achieved at the same time. The circular muscles in the mantle wall contract; this causes the inhalant valve to close, the exhalant valve to open and the mantle edge to lock tightly around the head. Water is forced out through the funnel which is pointed in the opposite direction to the required direction of travel. The inhalant phase is initiated by the relaxation of the circular muscles causes them to stretch, the connective tissue in the mantle wall recoils elastically, the mantle cavity expands causing the inhalant valve to open, the exhalant valve to close and water to flow into the cavity. This cycle of exhalation and inhalation is repeated to provide continuous locomotion. Fast jetting is an escape response. In this form of locomotion, radial muscles in the mantle wall are involved as well as circular ones, making it possible to hyper-inflate the mantle cavity with a larger volume of water than during slow jetting. On contraction, water flows out with great force, the funnel always being pointed anteriorly, and travel is backwards. During this means of locomotion, some squid exit the water in a similar way to
flying fish The Exocoetidae are a family of marine fish in the order Beloniformes class Actinopterygii, known colloquially as flying fish or flying cod. About 64 species are grouped in seven to nine genera. While they cannot fly in the same way a bird d ...
, gliding through the air for up to , and occasionally ending up on the decks of ships.


Feeding

Squid are
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
s, and, with their strong arms and suckers, can overwhelm relatively large animals efficiently. Prey is identified by sight or by touch, grabbed by the tentacles which can be shot out with great rapidity, brought back to within reach of the arms, and held by the hooks and suckers on their surface. In some species, the squid's saliva contains
toxin A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849 ...
s which act to subdue the prey. These are injected into its bloodstream when the prey is bitten, along with vasodilators and chemicals to stimulate the heart, and quickly circulate to all parts of its body. The deep sea squid '' Taningia danae'' has been filmed releasing blinding flashes of light from large photophores on its arms to illuminate and disorientate potential prey. Although squid can catch large prey, the mouth is relatively small, and the food must be cut into pieces by the chitinous beak with its powerful muscles before being swallowed. The radula is located in the buccal cavity and has multiple rows of tiny teeth that draw the food backwards and grind it in pieces. The deep sea squid '' Mastigoteuthis'' has the whole length of its whip-like tentacles covered with tiny suckers; it probably catches small organisms in the same way that flypaper traps flies. The tentacles of some bathypelagic squids bear
photophore A photophore is a glandular organ that appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors, ...
s which may bring food within its reach by attracting prey. Squid are among the most intelligent invertebrates. For example, groups of
Humboldt squid The Humboldt squid (''Dosidicus gigas''), also known as jumbo squid or jumbo flying squid (EN), and Pota in Peru or Jibia in Chile (ES) is a large, predatory squid living in the eastern Pacific Ocean. It is the only known species of the genus ''D ...
hunt cooperatively, spiralling up through the water at night and coordinating their vertical and horizontal movements while foraging.


Reproduction

Courtship in squid takes place in the open water and involves the male selecting a female, the female responding, and the transfer by the male of spermatophores to the female. In many instances, the male may display to identify himself to the female and drive off any potential competitors. Elaborate changes in body patterning take place in some species in both agonistic and courtship behaviour. The
Caribbean reef squid The Caribbean reef squid (''Sepioteuthis sepioidea''), commonly called the reef squid, is a species of small, torpedo-shaped squid with undulating fins that extend nearly the entire length of the body, approximately in length. In 2001, marine b ...
(''Sepioteuthis sepioidea''), for example, employs a complex array of colour changes during courtship and social interactions and has a range of about 16 body patterns in its repertoire. The pair adopt a head-to-head position, and "jaw locking" may take place, in a similar manner to that adopted by some cichlid fish. The heterodactylus of the male is used to transfer the spermatophore and deposit it in the female's mantle cavity in the position appropriate for the species; this may be adjacent to the gonopore or in a seminal receptacle. The sperm may be used immediately or may be stored. As the eggs pass down the oviduct, they are wrapped in a gelatinous coating, before continuing to the mantle cavity, where they are fertilised. In ''
Loligo ''Loligo'' is a genus of squid and one of the most representative and widely distributed groups of myopsid squid. The genus was first described by Jean Baptiste Lamarck in 1798. However, the name had been used earlier than Lamarck (Schneider, ...
'', further coatings are added by the nidimental glands in the walls of the cavity and the eggs leave through a funnel formed by the arms. The female attaches them to the substrate in strings or groups, the coating layers swelling and hardening after contact with sea water. ''Loligo'' sometimes forms breeding aggregations which may create a "community pile" of egg strings. Some pelagic and deep sea squid do not attach their egg masses, which float freely.


Ecology

Squid mostly have an annual life cycle, growing fast and dying soon after spawning. The diet changes as they grow but mostly consists of large
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
and small
nekton Nekton or necton (from the ) refers to the actively swimming aquatic organisms in a body of water. The term was proposed by German biologist Ernst Haeckel to differentiate between the active swimmers in a body of water, and the passive organisms t ...
. In Antarctica for example,
krill Krill are small crustaceans of the order Euphausiacea, and are found in all the world's oceans. The name "krill" comes from the Norwegian word ', meaning "small fry of fish", which is also often attributed to species of fish. Krill are consi ...
is the main constituent of the diet, with other food items being
amphipods Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far describ ...
, other small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s, and large
arrow worms The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and can ...
. Fish are also eaten, and some squid are
cannibalistic Cannibalism is the act of consuming another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded in more than 1,500 species. Human cannibalism is well documented, bo ...
. As well as occupying a key role in the food chain, squid are an important prey for predators including sharks, sea birds, seals and whales. Juvenile squid provide part of the diet for
worms Worms may refer to: *Worm, an invertebrate animal with a tube-like body and no limbs Places *Worms, Germany Worms () is a city in Rhineland-Palatinate, Germany, situated on the Upper Rhine about south-southwest of Frankfurt am Main. It had ...
and small fish. When researchers studied the contents of the stomachs of
elephant seal Elephant seals are very large, oceangoing earless seals in the genus ''Mirounga''. Both species, the northern elephant seal (''M. angustirostris'') and the southern elephant seal (''M. leonina''), were hunted to the brink of extinction for oi ...
s in South Georgia, they found 96% squid by weight. In a single day, a sperm whale can eat 700 to 800 squid, and a
Risso's dolphin Risso's dolphin (''Grampus griseus'') is a dolphin, the only species of the genus ''Grampus''. Some of the closest related species to these dolphins include: pilot whales (''Globicephala'' spp.), pygmy killer whales (''Feresa attenuata''), melon ...
entangled in a net in the Mediterranean was found to have eaten angel clubhook squid, umbrella squid, reverse jewel squid and European flying squid, all identifiable from their indigestible beaks. '' Ornithoteuthis volatilis'', a common squid from the tropical Indo-Pacific, is predated by
yellowfin tuna The yellowfin tuna (''Thunnus albacares'') is a species of tuna found in pelagic waters of tropical and subtropical oceans worldwide. Yellowfin is often marketed as ahi, from the Hawaiian , a name also used there for the closely related bigeye ...
, longnose lancetfish, common dolphinfish and swordfish, the
tiger shark The tiger shark (''Galeocerdo cuvier'') is a species of requiem shark and the last extant member of the family Galeocerdonidae. It is a large macropredator, capable of attaining a length over . Populations are found in many tropical and tempera ...
, the
scalloped hammerhead shark The scalloped hammerhead (''Sphyrna lewini'') is a species of hammerhead shark in the family Sphyrnidae. It was originally known as ''Zygaena lewini''. The Greek word ''sphyrna'' translates into "hammer" in English, referring to the shape of thi ...
and the
smooth hammerhead shark The smooth hammerhead (''Sphyrna zygaena'') is a species of hammerhead shark, and part of the family Sphyrnidae. This species is named "smooth hammerhead" because of the distinctive shape of the head, which is flattened and laterally extended int ...
. Sperm whales also hunt this species extensively as does the
brown fur seal The brown fur seal (''Arctocephalus pusillus''), also known as the Cape fur seal, South African fur seal and Australian fur seal, is a species of fur seal. Description The brown fur seal is the largest and most robust member of the fur seal ...
. In the
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
, penguins and
wandering albatross The wandering albatross, snowy albatross, white-winged albatross or goonie (''Diomedea exulans'') is a large seabird from the family Diomedeidae, which has a circumpolar range in the Southern Ocean. It was the last species of albatross to be desc ...
es are major predators of '' Gonatus antarcticus''.


Human uses


In literature and art

Giant squid have featured as monsters of the deep since classical times. Giant squid were described by Aristotle (4th century BC) in his ''
History of Animals ''History of Animals'' ( grc-gre, Τῶν περὶ τὰ ζῷα ἱστοριῶν, ''Ton peri ta zoia historion'', "Inquiries on Animals"; la, Historia Animalium, "History of Animals") is one of the major texts on biology by the ancient Gr ...
'' and
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
(1st century AD) in his '' Natural History''. The
Gorgon A Gorgon ( /ˈɡɔːrɡən/; plural: Gorgons, Ancient Greek: Γοργών/Γοργώ ''Gorgṓn/Gorgṓ'') is a creature in Greek mythology. Gorgons occur in the earliest examples of Greek literature. While descriptions of Gorgons vary, the te ...
of
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities ...
may have been inspired by squid or octopus, the animal itself representing the severed head of
Medusa In Greek mythology, Medusa (; Ancient Greek: Μέδουσα "guardian, protectress"), also called Gorgo, was one of the three monstrous Gorgons, generally described as winged human females with living venomous snakes in place of hair. Those ...
, the beak as the protruding tongue and fangs, and its tentacles as the snakes. The six-headed sea monster of the ''
Odyssey The ''Odyssey'' (; grc, Ὀδύσσεια, Odýsseia, ) is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the ''Iliad'', th ...
'', Scylla, may have had a similar origin. The Nordic legend of the kraken may also have derived from sightings of large cephalopods. In literature,
H. G. Wells Herbert George Wells"Wells, H. G."
Revised 18 May 2015. ''
The Sea Raiders" featured a man-eating squid species ''Haploteuthis ferox''. The
science fiction Science fiction (sometimes shortened to Sci-Fi or SF) is a genre of speculative fiction which typically deals with imaginative and futuristic concepts such as advanced science and technology, space exploration, time travel, parallel uni ...
writer Jules Verne told a tale of a kraken-like monster in his 1870 novel ''
Twenty Thousand Leagues Under the Sea ''Twenty Thousand Leagues Under the Seas'' (french: Vingt mille lieues sous les mers) is a classic science fiction adventure novel by French writer Jules Verne. The novel was originally serialized from March 1869 through June 1870 in Pierre- ...
''.


As food

Squid form a major food resource and are used in cuisines around the world, notably in Japan where it is eaten as
ika sōmen refer to a type of sashimi that is made from raw squid cut into fine strips, vaguely resembling '' sōmen'' type noodles. They are typically served with grated ginger and soy sauce or a soy sauce-based mentsuyu sauce. They are slurped up, muc ...
, sliced into vermicelli-like strips; as
sashimi is a Japanese delicacy consisting of fresh raw fish or meat sliced into thin pieces and often eaten with soy sauce. Origin The word ''sashimi'' means "pierced body", i.e. " 刺身" = ''sashimi'', where 刺 し = ''sashi'' (pierced, stu ...
; and as
tempura is a typical Japanese dish usually consisting of seafood, meat and vegetables that have been battered and deep fried. The dish was introduced by the Portuguese in Nagasaki through fritter-cooking techniques in the 16th century. The word ...
. Three species of ''Loligo'' are used in large quantities, '' L. vulgaris'' in the Mediterranean (known as ''Calamar'' in Spanish, ''Calamaro'' in Italian); '' L. forbesii'' in the Northeast Atlantic; and '' L. pealei'' on the American East Coast. Among the Ommastrephidae, ''
Todarodes pacificus The Japanese flying squid, Japanese common squid or Pacific flying squid, scientific name ''Todarodes pacificus'', is a squid of the family Ommastrephidae. This animal lives in the northern Pacific Ocean, in the area surrounding Japan, along th ...
'' is the main commercial species, harvested in large quantities across the North Pacific in Canada, Japan and China. In English-speaking countries, squid as food is often called ''
calamari Squid is eaten in many cuisines; in English, the culinary name calamari is often used for squid dishes.'' Oxford English Dictionary'', 3rd edition, 2002''s.v.''/ref> There are many ways to prepare and cook squid. Fried squid is common in the ...
'', adopted from Italian into English in the 17th century. Squid are found abundantly in certain areas, and provide large catches for fisheries. The body can be stuffed whole, cut into flat pieces, or sliced into rings. The arms, tentacles, and ink are also edible; the only parts not eaten are the beak and gladius (pen). Squid is a good food source for
zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
and
manganese Manganese is a chemical element with the symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese is a transition metal with a multifaceted array of industrial alloy use ...
, and high in copper,
selenium Selenium is a chemical element with the symbol Se and atomic number 34. It is a nonmetal (more rarely considered a metalloid) with properties that are intermediate between the elements above and below in the periodic table, sulfur and tellurium, ...
, vitamin B12, and riboflavin.


Commercial fishing

According to the
FAO The Food and Agriculture Organization of the United Nations (FAO)french: link=no, Organisation des Nations unies pour l'alimentation et l'agriculture; it, Organizzazione delle Nazioni Unite per l'Alimentazione e l'Agricoltura is an intern ...
, the cephalopod catch for 2002 was . Of this, 2,189,206 tonnes, or 75.8 percent, was squid. The following table lists squid species fishery catches that exceeded in 2002.


In biomimicry

Prototype chromatophores that
mimic MIMIC, known in capitalized form only, is a former simulation computer language developed 1964 by H. E. Petersen, F. J. Sansom and L. M. Warshawsky of Systems Engineering Group within the Air Force Materiel Command at the Wright-Patterson AFB in ...
the squid's adaptive camouflage have been made by Bristol University researchers using an electroactive
dielectric In electromagnetism, a dielectric (or dielectric medium) is an electrical insulator that can be polarised by an applied electric field. When a dielectric material is placed in an electric field, electric charges do not flow through the mate ...
elastomer, a flexible "smart" material that changes its colour and texture in response to electrical signals. The researchers state that their goal is to create an artificial skin that provides rapid active camouflage. The squid giant axon inspired
Otto Schmitt Otto Herbert Schmitt (April 6, 1913 – January 6, 1998) was an American inventor, engineer, and biophysicist known for his scientific contributions to biophysics and for establishing the field of biomedical engineering. Schmitt also coined ...
to develop a
comparator In electronics, a comparator is a device that compares two voltages or currents and outputs a digital signal indicating which is larger. It has two analog input terminals V_+ and V_- and one binary digital output V_\text. The output is ideally : ...
circuit with hysteresis now called the
Schmitt trigger In electronics, a Schmitt trigger is a comparator circuit with hysteresis implemented by applying positive feedback to the noninverting input of a comparator or differential amplifier. It is an active circuit which converts an analog input ...
, replicating the axon's propagation of
nerve impulse An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s.


See also

* Paralarva


Notes


References


Sources

*


External links


CephBase: Teuthida

Colossal Squid at the Museum of New Zealand Te Papa Tongarewa

Market squid mating, laying eggs (video)

Scientific American – Giant Squid

The Cephalopod Page

The Octopus News Magazine Online
{{Authority control Commercial molluscs Cenozoic cephalopods Cretaceous cephalopods Extant Devonian first appearances Mollusc common names