A cephalopod is any member of the

mollusca 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 esti ...

class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...

Cephalopoda (

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 ...

plural , ; "head-feet") such as a

squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...

octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttle ...

cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...

, or

nautilus The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in t ...

. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of

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 ...

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 main ...

s (

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 ...

s) modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt

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 ...

. The study of cephalopods is a branch of

malacology Malacology is the branch of invertebrate zoology that deals with the study of the Mollusca (mollusks or molluscs), the second-largest phylum of animals in terms of described species after the arthropods. Mollusks include snails and slugs, clams, ...

known as

teuthology Teuthology (from Greek , "cuttlefish, squid", and , ''-logia'') is the study of cephalopods such as octopus, squid, and cuttlefish.

. Cephalopods became dominant during the

Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...

period, represented by primitive

nautiloid Nautiloids are a group of marine cephalopods ( Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and speciose, with over 2,500 recorded species ...

s. The class now contains two, only distantly related,

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

subclasses:

Coleoidea Subclass (biology), 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, ...

, which includes

es,

, and

; and

Nautiloid Nautiloids are a group of marine cephalopods ( Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and speciose, with over 2,500 recorded species ...

ea, represented by ''

Nautilus The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in t ...

'' and ''

Allonautilus The genus ''Allonautilus'' contains two species of nautiluses, which have a significantly different morphology from those placed in the sister taxon ''Nautilus''. ''Allonautilus'' is now thought to be a descendant of ''Nautilus'', rendering the l ...

''. In the Coleoidea, the molluscan shell has been internalized or is absent, whereas in the Nautiloidea, the external shell remains. About 800 living

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

of cephalopods have been identified. Two important extinct

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

are the

Ammonoidea Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttle ...

(ammonites) and

Belemnoidea Belemnoids are an extinct group of marine cephalopod, very similar in many ways to the modern squid and closely related to the modern cuttlefish. Like them, the belemnoids possessed an ink sac, but, unlike the squid, they possessed ten arms o ...

(belemnites). Extant cephalopods range in size from the 10 mm (0.3 in) ''

Idiosepius thailandicus ''Idiosepius thailandicus'', also known as the thai pygmy squid, is a species of bobtail squid native to the Indo-Pacific waters off Thailand.Reid, A. 2005. Family Idiosepiidae. ''In:'' P. Jereb & C.F.E. Roper, eds. ''Cephalopods of the world. ...

'' to the 14 m (45.1 ft)

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 ...

, the largest extant

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 ...

Distribution

There are over 800

species of cephalopod, although new species continue to be described. An estimated 11,000 extinct

have been described, although the soft-bodied nature of cephalopods means they are not easily fossilised. Cephalopods are found in all the oceans of Earth. None of them can tolerate

fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...

, but the brief squid, ''

Lolliguncula brevis ''Lolliguncula brevis'', or the Atlantic brief squid, is a small species of squid in the Loliginidae family. It is found in shallow parts of the western Atlantic Ocean. Distribution The Atlantic brief squid occurs most frequently in shallow wat ...

'', found in

Chesapeake Bay The Chesapeake Bay ( ) is the largest estuary in the United States. The Bay is located in the Mid-Atlantic (United States), Mid-Atlantic region and is primarily separated from the Atlantic Ocean by the Delmarva Peninsula (including the parts: the ...

, is a notable partial exception in that it tolerates

brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...

. Cephalopods are thought to be unable to live in fresh water due to multiple biochemical constraints, and in their >400 million year existence have never ventured into fully freshwater habitats. Cephalopods occupy most of the depth of the ocean, from the

abyssal plain An abyssal plain is an underwater plain on the deep ocean floor, usually found at depths between and . Lying generally between the foot of a continental rise and a mid-ocean ridge, abyssal plains cover more than 50% of the Earth's surface. T ...

to the sea surface. Their diversity is greatest near the equator (~40 species retrieved in nets at 11°N by a diversity study) and decreases towards the poles (~5 species captured at 60°N).

Biology

Nervous system and behavior

Cephalopods are widely regarded as the most intelligent of the

s, and have well developed senses and large

brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...

s (larger than those of

gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...

s). The

nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...

of cephalopods is the most complex of the invertebrates and their brain-to-body-mass ratio falls between that of

endotherm An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...

ic and

ectotherm An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...

ic vertebrates. Captive cephalopods have also been known to climb out of their aquaria, maneuver a distance of the lab floor, enter another aquarium to feed on the crabs, and return to their own aquarium. The brain is protected in a

cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck and ...

cranium. The giant

nerve A nerve is an enclosed, cable-like bundle of nerve fibers (called axons) in the peripheral nervous system. A nerve transmits electrical impulses. It is the basic unit of the peripheral nervous system. A nerve provides a common pathway for the e ...

fibers of the cephalopod mantle have been widely used for many years as experimental material in

neurophysiology Neurophysiology is a branch of physiology and neuroscience that studies nervous system function rather than nervous system architecture. This area aids in the diagnosis and monitoring of neurological diseases. Historically, it has been dominated b ...

; their large diameter (due to lack of

myelination Myelin is a lipid-rich material that surrounds nerve cell axons (the nervous system's "wires") to insulate them and increase the rate at which electrical impulses (called action potentials) are passed along the axon. The myelinated axon can be l ...

) makes them relatively easy to study compared with other animals. Many cephalopods are social creatures; when isolated from their own kind, some species have been observed

shoaling In biology, any group of fish that stay together for social reasons are shoaling, and if the group is swimming in the same direction in a coordinated manner, they are schooling. In common usage, the terms are sometimes used rather loosely. Ab ...

with fish. Some cephalopods are able to fly through the air for distances of up to 50 m. While cephalopods are not particularly aerodynamic, they achieve these impressive ranges by jet-propulsion; water continues to be expelled from the funnel while the organism is in the air. The animals spread their fins and tentacles to form wings and actively control lift force with body posture. One species, ''

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 ...

'', has been observed spreading tentacles in a flat fan shape with a mucus film between the individual tentacles while another, ''

Sepioteuthis sepioidea 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 ...

'', has been observed putting the tentacles in a circular arrangement.

Senses

Cephalopods have advanced vision, can detect gravity with

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 cons ...

s, and have a variety of chemical sense organs. Octopuses use their arms to explore their environment and can use them for depth perception.

Vision

Most cephalopods rely on vision to detect predators and prey, and to communicate with one another. Consequently, cephalopod vision is acute: training experiments have shown that the

common octopus The common octopus (''Octopus vulgaris'') is a mollusc belonging to the class Cephalopoda. ''Octopus vulgaris'' is one of the most studied of all octopus species, and also one of the most intelligent. It ranges from the eastern Atlantic, exte ...

can distinguish the brightness, size, shape, and horizontal or vertical orientation of objects. The morphological construction gives cephalopod eyes the same performance as shark eyes, however their construction differs, as cephalopods lack a cornea and have an everted retina. Cephalopods' eyes are also sensitive to the plane of polarization of light. Unlike many other cephalopods,

es do not have good vision; their eye structure is highly developed, but lacks a solid

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''), ...

. They have a simple "

pinhole A hole is an opening in or through a particular medium, usually a solid body. Holes occur through natural and artificial processes, and may be useful for various purposes, or may represent a problem needing to be addressed in many fields of en ...

" eye through which water can pass. Instead of vision, the animal is thought to use

olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...

as the primary sense for

foraging Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...

, as well as locating or identifying potential mates. Cuttlefish eye

Surprisingly, given their ability to change color, all octopuses and most cephalopods are considered to be

color blind Color blindness or color vision deficiency (CVD) is the decreased ability to see color or differences in color. It can impair tasks such as selecting ripe fruit, choosing clothing, and reading traffic lights. Color blindness may make some aca ...

. Coleoid cephalopods (octopus, squid, cuttlefish) have a single photoreceptor type and lack the ability to determine color by comparing detected photon intensity across multiple spectral channels. When

camouflaging 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 ...

themselves, they use their chromatophores to change brightness and pattern according to the background they see, but their ability to match the specific color of a background may come from cells such as 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 reflect light from the environment. They also produce visual pigments throughout their body, and may sense light levels directly from their body. Evidence of

color vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different wavelengths (i.e., different spectral power distributions) independently of light intensity. Color perception is a part of ...

has been found in the

sparkling enope 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''). It achieves color vision with three photoreceptors, which are based on the same

opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most pro ...

, but use distinct

retinal Retinal (also known as retinaldehyde) is a polyene chromophore. Retinal, bound to proteins called opsins, is the chemical basis of visual phototransduction, the light-detection stage of visual perception (vision). Some microorganisms use retin ...

molecules as chromophores: A1 (retinal), A3 (3-dehydroretinal), and A4 (4-hydroxyretinal). The A1-photoreceptor is most sensitive to green-blue (484 nm), the A2-photoreceptor to blue-green (500 nm), and the A4-photoreceptor to blue (470 nm) light. In 2015, a novel mechanism for spectral discrimination in cephalopods was described. This relies on the exploitation of

chromatic aberration In optics, chromatic aberration (CA), also called chromatic distortion and spherochromatism, is a failure of a lens to focus all colors to the same point. It is caused by dispersion: the refractive index of the lens elements varies with the wave ...

(wavelength-dependence of focal length). Numerical modeling shows that chromatic aberration can yield useful chromatic information through the dependence of image acuity on accommodation. The unusual off-axis slit and annular pupil shapes in cephalopods enhance this ability by acting as prisms which are scattering white light in all directions.

Photoreception

In 2015, molecular evidence was published indicating that cephalopod chromatophores are photosensitive; reverse transcription polymerase chain reactions (RT-PCR) revealed transcripts encoding

rhodopsin Rhodopsin, also known as visual purple, is a protein encoded by the RHO gene and a G-protein-coupled receptor (GPCR). It is the opsin of the rod cells in the retina and a light-sensitive receptor protein that triggers visual phototransduction ...

and retinochrome within the retinas and skin of the

longfin inshore squid The longfin inshore squid (''Doryteuthis pealeii'') is a species of squid of the family Loliginidae. Description This species of squid is often seen with a reddish hue, but like many types of squid can manipulate its color, varying from a deep ...

(''Doryteuthis pealeii''), and the

common cuttlefish #REDIRECT Common cuttlefish #REDIRECT Common cuttlefish {{redirect category shell, {{R from other capitalisation{{R from move ...

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