Cuttlefish or cuttles[3] 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.

Cuttlefish have large, W-shaped pupils, eight arms, and two tentacles furnished
with denticulated suckers, with which they secure their prey. They generally range
in size from 15 to 25 cm (6 to 10 in), with the largest species, Sepia apama,
reaching 50 cm (20 in) in mantle length and over 10.5 kg (23 lb) in mass.[4]

Cuttlefish eat small molluscs, crabs, shrimp, fish, octopus, worms, and other
cuttlefish. Their predators include dolphins, sharks, fish, seals, seabirds, and
other cuttlefish. The average life expectancy of a cuttlefish is about 1–2 years.
Recent studies indicate cuttlefish are among the most intelligent invertebrates.[5]
Cuttlefish also have one of the largest brain-to-body size ratios of all
invertebrates.[5]

The "cuttle" in cuttlefish comes from the Old English name for the species, cudele,
which may be cognate with the Old Norse koddi (cushion) and the Middle Low German
Kudel (rag).[6] The Greco-Roman world valued the cuttlefish as a source of the
unique brown pigment the creature releases from its siphon when it is alarmed. The
word for it in both Greek and Latin, sepia, now refers to the reddish-brown color
sepia in English.

Contents
1 Fossil record
2 Range and habitat
3 Anatomy and physiology
3.1 Visual system
3.2 Circulatory system
3.3 Cuttlebone
3.4 Ink
3.5 Arms and mantle cavity
3.6 Suckers and venom
3.7 Sleep-like behavior
4 Life cycle
4.1 Reproduction
5 Communication
5.1 Chromatic
5.2 Intraspecific communication
5.3 Interspecific communication
6 Diet
7 Taxonomy
8 Human uses
8.1 As food
8.2 Sepia
8.3 Metal casting
8.4 Smart clothing
8.5 Pets
9 See also
10 References
11 External links
Fossil record
The earliest sepia-like fossils of cuttlefish are from the Cretaceous period.[7][8]
Whether the earlier Trachyteuthis is assigned to this class, or to the
Octopodiformes, remains unclear.[9]

Range and habitat

S. mestus swimming (Australia)
The family Sepiidae, which contains all cuttlefish, inhabits tropical and temperate
ocean waters. They are mostly shallow-water animals, although they are known to go
to depths of about 600 m (2,000 ft).[10] They have an unusual biogeographic
pattern: they are present along the coasts of East and South Asia, Western Europe,
and the Mediterranean, as well as all coasts of Africa and Australia, but are
totally absent from the Americas. By the time the family evolved, ostensibly in the
Old World, the North Atlantic possibly had become too cold and deep for these warm-
water species to cross.[11] The common cuttlefish (Sepia officinalis), is found in
the Mediterranean, North and Baltic seas, although populations may occur as far
south as South Africa. They are found in sublittoral depths, between the low tide
line and the edge of the continental shelf, to about 180 m (600 ft).[12] The
cuttlefish is listed under the Red List category of "least concern" by the IUCN Red
List of Threatened Species. This means that while some over-exploitation of the
marine animal has occurred in some regions due to large-scale commercial fishing,
their wide geographic range prevents them from being too threatened. Ocean
acidification, however, caused largely by higher levels of carbon dioxide emitted
into the atmosphere, is cited as a potential threat.[13]

Anatomy and physiology

Visual system

The characteristic W-shape of the cuttlefish eye

File:Sepia eyelid shape.theora.ogv
Pupil expansion in Sepia officinalis
Cuttlefish, like other cephalopods, have sophisticated eyes. The organogenesis and
the final structure of the cephalopod eye fundamentally differ from those of
vertebrates such as humans.[14] Superficial similarities between cephalopod and
vertebrate eyes are thought to be examples of convergent evolution. The cuttlefish
pupil is a smoothly curving W-shape.[15][16] Although cuttlefish cannot see color,
[17] they can perceive the polarization of light, which enhances their perception
of contrast. They have two spots of concentrated sensor cells on their retinas
(known as foveae), one to look more forward, and one to look more backward. The eye
changes focus by shifting the position of the entire lens with respect to the
retina, instead of reshaping the lens as in mammals. Unlike the vertebrate eye, no
blind spot exists, because the optic nerve is positioned behind the retina. They
are capable of using stereopsis, enabling them to discern depth/distance because
their brain calculates the input from both eyes.[18][19]

The cuttlefish's eyes are thought to be fully developed before birth, and they
start observing their surroundings while still in the egg. In consequence, they may
prefer to hunt the prey they saw before hatching.[20]

Circulatory system
The blood of a cuttlefish is an unusual shade of green-blue, because it uses the
copper-containing protein haemocyanin to carry oxygen instead of the red, iron-
containing protein haemoglobin found in vertebrates' blood. The blood is pumped by
three separate hearts: two branchial hearts pump blood to the cuttlefish's pair of
gills (one heart for each), and the third pumps blood around the rest of the body.
Cuttlefish blood must flow more rapidly than that of most other animals because
haemocyanin carries substantially less oxygen than haemoglobin. Unlike most other
mollusks, cephalopods like cuttlefish have a closed circulatory system.

Cuttlebone
Main article: Cuttlebone

Top and bottom view of a cuttlebone, the buoyancy organ and internal shell of a
cuttlefish
Cuttlefish possess an internal structure called the cuttlebone, which is porous and
is made of aragonite. The pores provide it with buoyancy, which the cuttlefish
regulates by changing the gas-to-liquid ratio in the chambered cuttlebone via the
ventral siphuncle.[21] Each species' cuttlebone has a distinct shape, size, and
pattern of ridges or texture. The cuttlebone is unique to cuttlefish, and is one of
the features that distinguish them from their squid relatives.[22]

Ink
Like other marine mollusks, cuttlefish have ink stores that are used for chemical
deterrence, phagomimicry, sensory distraction, and evasion when attacked.[23] Its
composition results in a dark colored ink, rich in ammonium salts and amino acids
that may have a role in phagomimicry defenses.[23] The ink can be ejected to create
a "smoke screen" to hide the cuttlefish's escape, or it can be released as a
pseudomorph of similar size to the cuttlefish, acting as a decoy while the
cuttlefish swims away.[24]

Human use of this substance is wide-ranged. A common use is in cooking with squid
ink to darken and flavor rice and pasta. It adds a black tint and a sweet flavor to
the food. In addition to food, cuttlefish ink can be used with plastics and
staining of materials.[citation needed] The diverse composition of cuttlefish ink,
and its deep complexity of colors, allows for dilution and modification of its
color. Cuttlefish ink can be used to make noniridescent reds, blues, and greens,
[25] subsequently used for biomimetic colors and materials.[citation needed]

Arms and mantle cavity

Cuttlefish have eight arms and two additional elongated tentacles that are used to
grasp prey. The elongated tentacles and mantle cavity serve as defense mechanisms;
when approached by a predator, the cuttlefish can suck water into its mantle cavity
and spread its arms in order to appear larger than normal.[26] Though the mantle
cavity is used for jet propulsion, the main parts of the body that are used for
basic mobility are the fins, which can maneuver the cuttlefish in all directions.

Suckers and venom

The suckers of cuttlefish extend most of the length of their arms and along the
distal portion of their tentacles. Like other cephalopods, cuttlefish have "taste-
by-touch" sensitivity in their suckers, allowing them to discriminate among objects
and water currents that they contact.[27]

Some cuttlefish are venomous. The genes for venom production are thought to be
descended from a common ancestor.[28] The muscles of the flamboyant cuttlefish
(Metasepia pfefferi) contain a highly toxic, unidentified compound[5] as lethal as
that of a fellow cephalopod, the blue-ringed octopus.[29]

Sleep-like behavior
Sleep is a state of immobility characterized by being rapidly reversible,
homeostatically controlled, and increasing an organism's arousal threshold.[30][31]

To date one cephalopod species, Octopus vulgaris, has been shown to satisfy these
criteria.[32] Another species, Sepia officinalis, satisfies two of the three
criteria but has not yet been tested on the third (arousal threshold).[31][30]
Recent research shows that the sleep-like state in a common species of cuttlefish,
Sepia officinalis, shows predictable periods[31] of rapid eye movement, arm
twitching and rapid chromatophore changes.[30]

Life cycle
The lifespan of cuttlefish is only around one to two years, depending on the
species. They hatch from eggs fully developed, around a fourth of an inch long,
reaching one inch around the first two months. Before death cuttlefish go through
senescence when the cephalopod essentially deteriorates, or rots in place. Their
eyesight begins to fail which affects their ability to see, move, and hunt
efficiently. Once this process begins, cuttlefish tend to not live long due to
predation by other organisms. Captive breeders may euthanize dying cuttlefish by
freezing them or using life-ending chemicals that are made by aquarium companies.
[26]

Reproduction
Cuttlefish start to actively mate at around five months of age. Male cuttlefish
challenge one another for dominance and the best den during mating season. During
this challenge, no direct contact is usually made. The animals threaten each other
until one of them backs down and swims away. Eventually, the larger male cuttlefish
mate with the females by grabbing them with their tentacles, turning the female so
that the two animals are face-to-face, then using a specialized tentacle to insert
sperm sacs into an opening near the female's mouth. As males can also use their
funnels to flush others' sperm out of the female's pouch, the male then guards the
female until she lays the eggs a few hours later.[33] After laying her cluster of
eggs, the female cuttlefish secretes ink on them making them look very similar to
grapes. The egg case is produced through a complex capsule of the female accessory
genital glands and the ink bag.[34]

On occasion, a large competitor arrives to threaten the male cuttlefish. In these

instances, the male first attempts to intimidate the other male. If the competitor
does not flee, the male eventually attacks it to force it away. The cuttlefish that
can paralyze the other first, by forcing near its mouth, wins the fight and the
female. Since typically four or five (and sometimes as many as 10) males are
available for every female, this behavior is inevitable.[35]

Cuttlefish are indeterminate growers, so smaller cuttlefish always have a chance of

finding a mate the next year when they are bigger.[36] Additionally, cuttlefish
unable to win in a direct confrontation with a guard male have been observed
employing several other tactics to acquire a mate. The most successful of these
methods is camouflage; smaller cuttlefish use their camouflage abilities to
disguise themselves as a female cuttlefish. Changing their body color, and even
pretending to be holding an egg sack, disguised males are able to swim past the
larger guard male and mate with the female.[35][37][38]

Communication
Cephalopods are able to communicate visually using a diverse range of signals. To
produce these signals, cephalopods can vary four types of communication element:
chromatic (skin coloration), skin texture (e.g. rough or smooth), posture, and
locomotion. Changes in body appearance such as these are sometimes called
polyphenism. The common cuttlefish can display 34 chromatic, six textural, eight
postural and six locomotor elements, whereas flamboyant cuttlefish use between 42
and 75 chromatic, 14 postural, and seven textural and locomotor elements. The
Caribbean reef squid (Sepioteuthis sepioidea) is thought to have up to 35 distinct
signalling states.[39][40]

Visual signals of the common cuttlefish[39]

Chromic – light Chromic – dark Texture Posture Locomotor
White posterior triangle Anterior transverse mantle line Smooth skin Raised
arms Sitting
White square Posterior transverse mantle line Coarse skin Waving arms Bottom
suction
White mantle bar Anterior mantle bar Papillate skin Splayed arms Buried
White lateral stripe Posterior mantle bar Wrinkled first arms Drooping
arms Hovering
White fin spots Paired mantle spots White square papillae Extended fourth
arm Jetting
White fin line Median mantle stripe Major lateral papillae Flattened body
 Inking
White neck spots Mantle margin stripe Raised head
Iridescent ventral mantle Mantle margin scalloping Flanged fin
White zebra bands Dark fin line
White landmark spots Black zebra bands
White splotches Mottle
White major lateral papillae Lateroventral patches
White head bar Anterior head bar
White arm triangle Posterior head bar
Pink iridophore arm stripes Pupil
White arms spots (males only) Eye ring
Dark arm stripes
Dark arms
Chromatic
Two cuttlefish with dramatically different coloration
This broadclub cuttlefish (Sepia latimanus) can change from camouflage tans and
browns (top) to yellow with dark highlights (bottom) in less than one second.
Cuttlefish are sometimes referred to as the "chameleons of the sea" because of
their ability to rapidly alter their skin color – this can occur within one second.
Cuttlefish change color and pattern (including the polarization of the reflected
light waves), and the shape of the skin to communicate to other cuttlefish, to
camouflage themselves, and as a deimatic display to warn off potential predators.
Under some circumstances, cuttlefish can be trained to change color in response to
stimuli, thereby indicating their color changing is not completely innate.[41]

Cuttlefish can also affect the light's polarization, which can be used to signal to
other marine animals, many of which can also sense polarization, as well as being
able to influence the color of light as it reflects off their skin.[42] Although
cuttlefish (and most other cephalopods) lack color vision, high-resolution
polarisation vision may provide an alternative mode of receiving contrast
information that is just as defined.[43] The cuttlefish's wide pupil attenuates
chromatic aberration, allowing it to perceive color by focusing specific
wavelengths onto the retina.[44]

The three broad categories of color patterns are uniform, mottle, and disruptive.
[45] Cuttlefish can display as many as 12 to 14 patterns,[39] 13 of which have been
categorized as seven "acute" (relatively brief) and six "chronic" (long-lasting)
patterns.[46] although other researchers suggest the patterns occur on a continuum.
[45]

Patterns of the common cuttlefish[39]

Chronic Acute
Uniform light Uniform blanching
Stipple Uniform darkening
Light mottle Acute disruptive
Disruptive Deimatic
Dark mottle Flamboyant
Weak zebra Intense zebra
Passing cloud
The color-changing ability of cuttlefish is due to multiple types of cells. These
are arranged (from the skin's surface going deeper) as pigmented chromatophores
above a layer of reflective iridophores and below them, leucophores.[47][48]

Chromatophores
The chromatophores are sacs containing hundreds of thousands of pigment granules
and a large membrane that is folded when retracted. Hundreds of muscles radiate
from the chromatophore. These are under neural control and when they expand, they
reveal the hue of the pigment contained in the sac. Cuttlefish have three types of
chromatophore: yellow/orange (the uppermost layer), red, and brown/black (the
deepest layer). The cuttlefish can control the contraction and relaxation of the
muscles around individual chromatophores, thereby opening or closing the elastic
sacs and allowing different levels of pigment to be exposed.[40] Furthermore, the
chromatophores contain luminescent protein nanostructures in which tethered pigment
granules modify light through absorbance, reflection, and fluorescence between 650
and 720 nm.[49][50]

For cephalopods in general, the hues of the pigment granules are relatively
constant within a species, but can vary slightly between species. For example, the
common cuttlefish and the opalescent inshore squid (Loligo opalescens) have yellow,
red, and brown, the European common squid (Alloteuthis subulata) has yellow and
red, and the common octopus has yellow, orange, red, brown, and black.[40]

In cuttlefish, activation of a chromatophore can expand its surface area by 500%.

Up to 200 chromatophores per mm2 of skin may occur. In Loligo plei, an expanded
chromatophore may be up to 1.5 mm in diameter, but when retracted, it can measure
as little as 0.1 mm.[49][51][52]

Iridophores
Retracting the chromatophores reveals the iridophores and leucophores beneath them,
thereby allowing cuttlefish to use another modality of visual signalling brought
about by structural coloration.

Iridophores are structures that produce iridescent colors with a metallic sheen.
They reflect light using plates of crystalline chemochromes made from guanine. When
illuminated, they reflect iridescent colors because of the diffraction of light
within the stacked plates. Orientation of the schemochrome determines the nature of
the color observed. By using biochromes as colored filters, iridophores create an
optical effect known as Tyndall or Rayleigh scattering, producing bright blue or
blue-green colors. Iridophores vary in size, but are generally smaller than 1 mm.
Squid at least are able to change their iridescence. This takes several seconds or
minutes, and the mechanism is not understood.[53] However, iridescence can also be
altered by expanding and retracting the chromatophores above the iridophores.
Because chromatophores are under direct neural control from the brain, this effect
can be immediate.

Cephalopod iridophores polarize light. Cephalopods have a rhabdomeric visual system

which means they are visually sensitive to polarized light. Cuttlefish use their
polarization vision when hunting for silvery fish (their scales polarize light).
Female cuttlefish exhibit a greater number of polarized light displays than males
and also alter their behavior when responding to polarized patterns. The use of
polarized reflective patterns has led some to suggest that cephalopods may
communicate intraspecifically in a mode that is "hidden" or "private" because many
of their predators are insensitive to polarized light.[53][54][52]

Leucophores

The white spots and bands on this cuttlefish are produced by leucophores.
Leucophores, usually located deeper in the skin than iridophores, are also
structural reflectors using crystalline purines, often guanine, to reflect light.
Unlike iridophores, however, leucophores have more organized crystals that reduce
diffraction. Given a source of white light, they produce a white shine, in red they
produce red, and in blue they produce blue. Leucophores assist in camouflage by
providing light areas during background matching (e.g. by resembling light-colored
objects in the environment) and disruptive coloration (by making the body appear to
be composed of high-contrasting patches).[53]

The reflectance spectra of cuttlefish patterns and several natural substrates

(stipple, mottle, disruptive) can be measured using an optic spectrometer.[53]
Intraspecific communication
Cuttlefish sometimes use their color patterns to signal future intent to other
cuttlefish. For example, during agonistic encounters, male cuttlefish adopt a
pattern called the intense zebra pattern, considered to be an honest signal. If a
male is intending to attack, it adopts a "dark face" change, otherwise, it remains
pale.[55]

In at least one species, female cuttlefish react to their own reflection in a

mirror and to other females by displaying a body pattern called "splotch". However,
they do not use this display in response to males, inanimate objects, or prey. This
indicates they are able to discriminate same-sex conspecifics, even when human
observers are unable to discern the sex of a cuttlefish in the absence of sexual
dimorphism.[56]

Female cuttlefish signal their receptivity to mating using a display called

precopulatory grey.[56] Male cuttlefish sometimes use deception toward guarding
males to mate with females. Small males hide their sexually dimorphic fourth arms,
change their skin pattern to the mottled appearance of females, and change the
shape of their arms to mimic those of nonreceptive, egg-laying females.[38]

Displays on one side of a cuttlefish can be independent of the other side of the
body; males can display courtship signals to females on one side while
simultaneously showing female-like displays with the other side to stop rival males
interfering with their courtship.[57]

Interspecific communication
The deimatic display (a rapid change to black and white with dark ‘eyespots’ and
contour, and spreading of the body and fins) is used to startle small fish that are
unlikely to prey on the cuttlefish, but use the flamboyant display towards larger,
more dangerous fish,[58] and give no display at all to chemosensory predators such
as crabs and dogfish.[59]

One dynamic pattern shown by cuttlefish is dark mottled waves apparently repeatedly
moving down the body of the animals. This has been called the passing cloud
pattern. In the common cuttlefish, this is primarily observed during hunting, and
is thought to communicate to potential prey – “stop and watch me”[40] – which some
have interpreted as a type of "hypnosis".

Camouflage
Further information: Camouflage, Crypsis, and Animal coloration

Juvenile cuttlefish camouflaged against the seafloor

External video
Kings of Camouflage– Nova documentary
Cuttlefish are able to rapidly change the color of their skin to match their
surroundings and create chromatically complex patterns,[59] despite their inability
to perceive color, through some mechanism which is not completely understood.[60]
They have been seen to have the ability to assess their surroundings and match the
color, contrast and texture of the substrate even in nearly total darkness.[51]

The color variations in the mimicked substrate and animal skin are similar.
Depending on the species, the skin of cuttlefish responds to substrate changes in
distinctive ways. By changing naturalistic backgrounds, the camouflage responses of
different species can be measured.[61] Sepia officinalis changes color to match the
substrate by disruptive patterning (contrast to break up the outline), whereas S.
pharaonis matches the substrate by blending in. Although camouflage is achieved in
different ways, and in an absence of color vision, both species change their skin
colors to match the substrate. Cuttlefish adapt their own camouflage pattern in
ways that are specific for a particular habitat. An animal could settle in the sand
and appear one way, with another animal a few feet away in a slightly different
microhabitat, settled in algae for example, will be camouflaged quite differently.
[51]

Cuttlefish are also able to change the texture of their skin. The skin contains
bands of circular muscle which as they contract, push fluid up. These can be seen
as little spikes, bumps, or flat blades. This can help with camouflage when the
cuttlefish becomes texturally as well as chromatically similar to objects in its
environment such as kelp or rocks.[51]

Diet
File:Red cuttle hunting.webm
Video of S. mestus in Sydney waters, hunting and catching prey
While the preferred diet of cuttlefish is crabs and fish, they feed on small shrimp
shortly after hatching.[62]

Cuttlefish use their camouflage to hunt and sneak up on their prey.[63] They swim
at the bottom, where shrimp and crabs are found, and shoot out a jet of water to
uncover the prey buried in the sand. Then when the prey tries to escape, the
cuttlefish open their eight arms and shoot out two long feeding tentacles to grab
them. Each arm has a pad covered in suckers, which grabs and pulls prey toward its
beak, paralyzing it with venom before eating it.[62] To achieve a hypnotic effect
and stun prey before catching it, cuttlefish are also known to change color
rapidly.

Taxonomy
Wikispecies has information related to Sepiida

Illustration of Sepia officinalis

File:Cuttlefish.ogv
Video of a cuttlefish in its natural habitat
Over 120 species of cuttlefish are currently recognised, grouped into six families.

Order Sepiida: cuttlefish

Suborder †Vasseuriina
Family †Vasseuriidae
Family †Belosepiellidae
Suborder Sepiina
Family †Belosaepiidae
Family Sepiadariidae
Family Sepiidae
Family Sepiolidae

The common cuttlefish (Sepia officinalis) is the best-known cuttlefish species

Hooded cuttlefish (Sepia prashadi)

Engravings by the Dutch zoologist Albertus Seba, 1665–1736

Human uses
As food
Three-sided white plate containing linguini
Linguine with cuttlefish and ink sauce served at a Venetian osteria
Cuttlefish are caught for food in the Mediterranean, East Asia, the English
Channel, and elsewhere.

In East Asia, dried, shredded cuttlefish is a popular snack food. In the Qing
Dynasty manual of Chinese gastronomy, the Suiyuan shidan, the roe of the cuttlefish
is considered a difficult-to-prepare, but sought-after delicacy.[64]

Cuttlefish are quite popular in Europe. For example, in northeast Italy, they are
used in risotto al nero di seppia (risotto with cuttlefish ink), also found in
Croatia and Montenegro as crni rižot (black risotto). Catalan cuisine, especially
that of the coastal regions, uses cuttlefish and squid ink in a variety of tapas
and dishes such as arròs negre. Breaded and deep-fried cuttlefish is a popular dish
in Andalusia. In Portugal, cuttlefish is present in many popular dishes. Chocos com
tinta (cuttlefish in black ink), for example, is grilled cuttlefish in a sauce of
its own ink. Cuttlefish is also popular in the region of Setúbal, where it is
served as deep-fried strips or in a variant of feijoada, with white beans. Black
pasta is often made using cuttlefish ink.

Sepia
Cuttlefish ink was formerly an important dye, called sepia. To extract the sepia
pigment from a cuttlefish (or squid), the inc sac is removed and dried then
dissolved in a dilute alkali. The resulting solution is filtered to isolate the
pigment, which is then precipitated with dilute hydrochloric acid. The isolated
precipitate is the sepia pigment.[citation needed] It is relatively chemically
inert, which contributes to its longevity. Today, artificial dyes have mostly
replaced n