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

 1Fossil record
 2Range and habitat
 3Anatomy and physiology
o 3.1Visual system
o 3.2Circulatory system
o 3.3Cuttlebone
o 3.4Specialized components
o 3.5Sleep-like behavior
 4Reproduction
 5Communication
o 5.1Chromatic
o 5.2Intraspecific communication
o 5.3Interspecific communication
 6Diet
 7Taxonomy
 8Human uses
o 8.1As food
o 8.2Sepia
o 8.3Metal casting
o 8.4Smart clothing
o 8.5Pets
 9See also
 10References
 11External links

Fossil record[edit]
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[edit]

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 (100 fathoms)."[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[edit]

Visual system[edit]

The characteristic W-shape of the cuttlefish eye

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 below the retina.
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.[18]

Circulatory system[edit]
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[edit]
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 ventralsiphuncle.[19] 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.[20]

Specialized components[edit]
Ink[edit]
Like other marine mollusks, cuttlefish have ink stores that are used for chemical
deterrence, phagomimicry, sensory distraction, and evasion when attacked.[21] The composition
of the ink leads to a richly colored "black" ink which is rich in ammonium salts and amino
acids that may have a role in phagomimicry defenses.[21]
The use of this color is wide-ranged. A common use is in cooking with squid ink to darken and
flavor rice and pasta.[citation needed] It is purported to add a sweet flavor to the food, and leaves the
food stained a dark black color. 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, [22] subsequently used for biomimetic colors and
materials.[citation needed]
Suckers and venom[edit]
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.[23]
Some cuttlefish are venomous. The genes for venom production are thought to be descended
from a common ancestor.[24] 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.[25]

Sleep-like behavior[edit]
Sleep is a state of immobility characterized by being rapidly reversible, homeostatically
controlled, and increasing an organism's arousal threshold [26][27].
To date one cephalopod species, Octopus vulgaris, has been shown to satisfy these
criteria.[28] Another species, Sepia officinalis, satisfies two of the three criteria but has not yet
been tested on the third (arousal threshold).[27][26] Recent research shows that the sleep-like
state in a common species of cuttlefish, Sepia officinalis, shows predictable periods[27]of rapid
eye movement, arm twitching and rapid chromatophore changes.[26]

Reproduction[edit]
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. The male then guards the female until she lays the eggs a few hours later.[29]
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.[30]
Cuttlefish are indeterminate growers, so smaller cuttlefish always have a chance at finding a
mate the next year when they are bigger.[31] 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.[30][32][33]

Communication[edit]
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. For the Caribbean reef squid (Sepioteuthis sepioidea), they
are thought to have up to 35 distinct signalling states.[34][35]

Visual signals of the common cuttlefish[34]

Chromic - light Chromic - dark Texture Posture Locomotor

White posterior Anterior transverse

Smooth skin Raised arms Sitting
triangle mantle line

Posterior transverse Bottom

White square Coarse skin Waving arms
mantle line suction

White mantle bar Anterior mantle bar Papillate skin Splayed arms Buried

Wrinkled first Drooping

White lateral stripe Posterior mantle bar Hovering
arms arms

White square Extended

White fin spots Paired mantle spots Jetting
papillae fourth arm

Major lateral Flattened

White fin line Median mantle stripe Inking
papillae body
White neck spots Mantle margin stripe Raised head

Iridescent ventral Mantle margin

Flanged fin
mantle scalloping

White zebra bands Dark fin line

White landmark
Black zebra bands
spots

White splotches Mottle

White major lateral Lateroventral

papillae patches

White head bar Anterior head bar

White arm triangle Posterior head bar

Pink iridophore
Pupil
arm stripes

White arms spots

Eye ring
(males only)

Dark arm stripes

Dark arms

Chromatic[edit]
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.[36]
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.[37] 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.[38] The cuttlefish's wide pupil attenuates chromatic aberration,
allowing it to perceive color by focusing specific wavelengths onto the retina.[39]
The three broad categories of color patterns are uniform, mottle, and disruptive.[40] Cuttlefish
can display as many as 12 to 14 patterns,[34] 13 of which have been categorized as seven
"acute" (relatively brief) and six "chronic" (long-lasting) patterns.[41] although other researchers
suggest the patterns occur on a continuum.[40]

Patterns of the common cuttlefish[34]

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.[42][43]
Chromatophores[edit]
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.[35] Furthermore, the chromatophores contain luminescent protein nanostructures in
which tethered pigment granules modify light through absorbance, reflection,
and fluorescence between 650 and 720 nm.[44][45]
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.[35]
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.[44][46][47]
Iridophores[edit]
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.[48] 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.[48][49][47]
Leucophores[edit]

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).[48]
The reflectance spectra of cuttlefish patterns and several natural substrates
(stipple, mottle, disruptive) can be measured using an optic spectrometer.[48]

Intraspecific communication[edit]
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.[50]
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.[51]
Female cuttlefish signal their receptivity to mating using a display called precopulatory
grey.[51] 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.[33]
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.[52]

Interspecific communication[edit]
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,[53] and give no
display at all to chemosensory predators such as crabs and dogfish.[54]
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”[35] – which some have interpreted as a type of "hypnosis".
Camouflage[edit]
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,[54] despite their inability to perceive color, through some
mechanism which is not completely understood.[55] 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.[46]
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.[56] 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.[46]
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.[46]

Diet[edit]

Video of S. mestus in Sydneywaters, hunting and catching prey

While the preferred diet of cuttlefish is crabs and fish, they feed on small shrimp shortly after
hatching.[57]
Cuttlefish use their camouflage to hunt and sneak up on their prey.[58] 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.[57] To achieve a
hypnotic effect and stun prey before catching it, cuttlefish are also known to change color
rapidly.[citation needed]

Taxonomy[edit]
Wikispecies has
information related
to Sepiida
Illustration of Sepia officinalis

Video of a cuttlefish in its natural habitat

Over 120 species of cuttlefish are currently recognised, grouped into six families.

 Order Sepiida: cuttlefish

o Suborder †Vasseuriina
 Family †Vasseuriidae
 Family †Belosepiellidae
o 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[edit]
As food[edit]

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.[59]
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[edit]
Cuttlefish ink was formerly an important dye, called sepia. Today, artificial dyes have mostly
replaced natural sepia.

Metal casting[edit]
Cuttlebone has been used since antiquity to make casts for metal. A model is pushed into the
cuttlebone and removed, leaving an impression. Molten gold, silver or pewter can then be
poured into the cast.[60][61]
Smart clothing[edit]
Research into replicating biological color-changing has led to engineering artificial
chromatophores out of small devices known as dielectric elastomer actuators. Engineers at
the University of Bristol have engineered soft materials that mimic the color-changing skin of
animals like cuttlefish,[62] paving the way for "smart clothing" and camouflage applications.[63]

Pets[edit]
Though cuttlefish are rarely kept as pets, due in part to their fairly short lifetimes, the most
common to be kept are Sepia officinalis or Sepia bandensis.[64]
Cuttlebone is given to parakeets and other cagebirds as a source of dietary calcium