124-600: Discosorida are an order of cephalopods that lived from the beginning of the Middle Ordovician , through the Silurian , and into the Devonian . Discosorids are unique in the structure and formation of the siphuncle , the tube that runs through and connects the camerae (chambers) in cephalopods, which unlike those in other orders is zoned longitudinally along the segments rather than laterally. Siphuncle structure indicated that
248-440: A closed circulatory system. Like other marine mollusks, cuttlefish have ink stores that are used for chemical deterrence, phagomimicry , sensory distraction, and evasion when attacked. Its composition results in a dark colored ink, rich in ammonium salts and amino acids that may have a role in phagomimicry defenses. The ink can be ejected to create a " smoke screen " to hide the cuttlefish's escape, or it can be released as
372-624: 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. 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. Although cuttlefish (and most other cephalopods) lack color vision, high-resolution polarisation vision may provide an alternative mode of receiving contrast information that
496-802: A pseudomorph of similar size to the cuttlefish, acting as a decoy while the cuttlefish swims away. 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. 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, subsequently used for biomimetic colors and materials. A common gene between cuttlefish and almost all other cephalopods allows them to produce venom, excreting it through their beak to help kill their prey. Additionally,
620-464: A squid , octopus , cuttlefish , or nautilus . These exclusively marine animals are characterized by bilateral body symmetry , a prominent head, and a set of arms or tentacles ( muscular hydrostats ) modified from the primitive molluscan foot. Fishers sometimes call cephalopods " inkfish ", referring to their common ability to squirt ink . The study of cephalopods is a branch of malacology known as teuthology . Cephalopods became dominant during
744-427: A "shell vestige" or "gladius". The Incirrina have either a pair of rod-shaped stylets or no vestige of an internal shell, and some squid also lack a gladius. The shelled coleoids do not form a clade or even a paraphyletic group. The Spirula shell begins as an organic structure, and is then very rapidly mineralized. Shells that are "lost" may be lost by resorption of the calcium carbonate component. Females of
868-399: A cloud of dark ink to confuse predators . This sac is a muscular bag which originated as an extension of the hindgut. It lies beneath the gut and opens into the anus, into which its contents – almost pure melanin – can be squirted; its proximity to the base of the funnel means the ink can be distributed by ejected water as the cephalopod uses its jet propulsion. The ejected cloud of melanin
992-433: 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. 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
1116-606: 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
1240-447: A diversity of backgrounds. Experiments done in Dwarf chameleons testing these hypotheses showed that chameleon taxa with greater capacity for color change had more visually conspicuous social signals but did not come from more visually diverse habitats, suggesting that color change ability likely evolved to facilitate social signaling, while camouflage is a useful byproduct. Because camouflage
1364-798: 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. 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 )
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#17328699602551488-529: A flat fan shape with a mucus film between the individual tentacles, while another, Sepioteuthis sepioidea , has been observed putting the tentacles in a circular arrangement. Cephalopods have advanced vision, can detect gravity with statocysts , and have a variety of chemical sense organs. Octopuses use their arms to explore their environment and can use them for depth perception. Most cephalopods rely on vision to detect predators and prey and to communicate with one another. Consequently, cephalopod vision
1612-613: 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. 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
1736-652: A gunshot-like popping noise, thought to function to frighten away potential predators. Cephalopods employ a similar method of propulsion despite their increasing size (as they grow) changing the dynamics of the water in which they find themselves. Thus their paralarvae do not extensively use their fins (which are less efficient at low Reynolds numbers ) and primarily use their jets to propel themselves upwards, whereas large adult cephalopods tend to swim less efficiently and with more reliance on their fins. Early cephalopods are thought to have produced jets by drawing their body into their shells, as Nautilus does today. Nautilus
1860-453: A jet as a propulsion mechanism. Squids do not have the longitudinal muscles that octopus do. Instead, they have a tunic. This tunic is made of layers of collagen and it surrounds the top and the bottom of the mantle. Because they are made of collagen and not muscle, the tunics are rigid bodies that are much stronger than the muscle counterparts. This provides the squids some advantages for jet propulsion swimming. The stiffness means that there
1984-437: 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 it 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
2108-406: A length of 8 metres. They may terminate in a broadened, sucker-coated club. The shorter four pairs are termed arms , and are involved in holding and manipulating the captured organism. They too have suckers, on the side closest to the mouth; these help to hold onto the prey. Octopods only have four pairs of sucker-coated arms, as the name suggests, though developmental abnormalities can modify
2232-474: A muscle, which is why they can change their skin hue as rapidly as they do. Coloration is typically stronger in near-shore species than those living in the open ocean, whose functions tend to be restricted to disruptive camouflage . These chromatophores are found throughout the body of the octopus, however, they are controlled by the same part of the brain that controls elongation during jet propulsion to reduce drag. As such, jetting octopuses can turn pale because
2356-512: A novel mechanism for spectral discrimination in cephalopods was described. This relies on the exploitation of chromatic aberration (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. In 2015, molecular evidence
2480-536: 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. 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
2604-500: A rapidly expanding siphuncle with segments that extend into the adjacent chambers, and parietal deposits within the siphuncle that overlap to form endocones. The Discosorida include these families, more or less in phylogenetic sequence beginning with the oldest: These form three basin evolutionary lineages. The first, formed by the Reudemannoceratidae, Cyrtogomphoceratidae, and Phragmoceratidae, are fundamentally endogastric with
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#17328699602552728-465: A rare form of physiological color change which utilizes neural control of muscles to change the morphology of their chromatophores. This neural control of chromatophores has evolved convergently in both cephalopods and teleosts fishes. With the exception of the Nautilidae and the species of octopus belonging to the suborder Cirrina , all known cephalopods have an ink sac, which can be used to expel
2852-530: A shell with a phragmocone divided into chambers separated by septa. 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 . 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. Cuttlefish, like other cephalopods, have sophisticated eyes. The organogenesis and
2976-429: A shell-less subclass of cephalopods (squid, cuttlefish, and octopuses), have complex pigment containing cells called chromatophores which are capable of producing rapidly changing color patterns. These cells store pigment within an elastic sac which produces the color seen from these cells. Coleoids can change the shape of this sac, called the cytoelastic sacculus, which then causes changes in the translucency and opacity of
3100-675: 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). The reflectance spectra of cuttlefish patterns and several natural substrates ( stipple , mottle , disruptive ) can be measured using an optic spectrometer . Cuttlefish sometimes use their color patterns to signal future intent to other cuttlefish. For example, during agonistic encounters, male cuttlefish adopt
3224-620: A startling array of fashions. As well as providing camouflage with their background, some cephalopods bioluminesce, shining light downwards to disguise their shadows from any predators that may lurk below. The bioluminescence is produced by bacterial symbionts; the host cephalopod is able to detect the light produced by these organisms. Bioluminescence may also be used to entice prey, and some species use colorful displays to impress mates, startle predators, or even communicate with one another. Cephalopods can change their colors and patterns in milliseconds, whether for signalling (both within
3348-433: Is about 1–2 years. Studies are said to indicate cuttlefish to be among the most intelligent invertebrates . Cuttlefish also have one of the largest brain-to-body size ratios of all invertebrates. 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 the cuttlefish in both Greek and Latin , sepia , now refers to
3472-516: Is acute: training experiments have shown that the common octopus 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, nautiluses do not have good vision; their eye structure
3596-458: Is also capable of creating a jet by undulations of its funnel; this slower flow of water is more suited to the extraction of oxygen from the water. When motionless, Nautilus can only extract 20% of oxygen from the water. The jet velocity in Nautilus is much slower than in coleoids , but less musculature and energy is involved in its production. Jet thrust in cephalopods is controlled primarily by
3720-570: Is aragonite. As for other mollusc shells or coral skeletons, the smallest visible units are irregular rounded granules. Cephalopods, as the name implies, have muscular appendages extending from their heads and surrounding their mouths. These are used in feeding, mobility, and even reproduction. In coleoids they number eight or ten. Decapods such as cuttlefish and squid have five pairs. The longer two, termed tentacles , are actively involved in capturing prey; they can lengthen rapidly (in as little as 15 milliseconds ). In giant squid they may reach
3844-427: Is highly developed, but lacks a solid lens . They have a simple " pinhole " eye through which water can pass. Instead of vision, the animal is thought to use olfaction as the primary sense for foraging , as well as locating or identifying potential mates. All octopuses and most cephalopods are considered to be color blind . Coleoid cephalopods (octopus, squid, cuttlefish) have a single photoreceptor type and lack
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3968-436: Is inevitable. Cuttlefish are indeterminate growers , so smaller cuttlefish always have a chance of finding a mate the next year when they are bigger. 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
4092-455: Is just as defined. The cuttlefish's wide pupil may accentuate chromatic aberration, allowing it to perceive color by focusing specific wavelengths onto the retina. The three broad categories of color patterns are uniform, mottle, and disruptive. Cuttlefish can display as many as 12 to 14 patterns, 13 of which have been categorized as seven "acute" (relatively brief) and six "chronic" (long-lasting) patterns. although other researchers suggest
4216-418: Is more efficient, but in environments with little oxygen and in low temperatures, hemocyanin has the upper hand. The hemocyanin molecule is much larger than the hemoglobin molecule, allowing it to bond with 96 O 2 or CO 2 molecules, instead of the hemoglobin's just four. But unlike hemoglobin, which are attached in millions on the surface of a single red blood cell, hemocyanin molecules float freely in
4340-403: Is needed, compensating for their small size. However, organisms which spend most of their time moving slowly along the bottom do not naturally pass much water through their cavity for locomotion; thus they have larger gills, along with complex systems to ensure that water is constantly washing through their gills, even when the organism is stationary. The water flow is controlled by contractions of
4464-439: Is no necessary muscle flexing to keep the mantle the same size. In addition, tunics take up only 1% of the squid mantle's wall thickness, whereas the longitudinal muscle fibers take up to 20% of the mantle wall thickness in octopuses. Also because of the rigidity of the tunic, the radial muscles in squid can contract more forcefully. The mantle is not the only place where squids have collagen. Collagen fibers are located throughout
4588-457: Is referred to as a pseudomorph ). This strategy often results in the predator attacking the pseudomorph, rather than its rapidly departing prey. For more information, see Inking behaviors . The ink sac of cephalopods has led to a common name of "inkfish", formerly the pen-and-ink fish. Cephalopods are the only molluscs with a closed circulatory system. Coleoids have two gill hearts (also known as branchial hearts ) that move blood through
4712-413: Is supplemented with fin motion; in the squid, the fins flap each time that a jet is released, amplifying the thrust; they are then extended between jets (presumably to avoid sinking). Oxygenated water is taken into the mantle cavity to the gills and through muscular contraction of this cavity, the spent water is expelled through the hyponome , created by a fold in the mantle. The size difference between
4836-553: Is the first evidence that cephalopod dermal tissues may possess the required combination of molecules to respond to light. Some squids have been shown to detect sound using their statocysts , but, in general, cephalopods are deaf. Most cephalopods possess an assemblage of skin components that interact with light. These may include iridophores, leucophores , chromatophores and (in some species) photophores . Chromatophores are colored pigment cells that expand and contract in accordance to produce color and pattern which they can use in
4960-423: Is the most complex of the invertebrates and their brain-to-body-mass ratio falls between that of endothermic and ectothermic 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 captive crabs, and return to their own aquarium. The brain is protected in a cartilaginous cranium. The giant nerve fibers of
5084-441: Is thought to communicate to potential prey – "stop and watch me" – which some have interpreted as a type of "hypnosis". Cuttlefish are able to rapidly change the color of their skin to match their surroundings and create chromatically complex patterns, despite their inability to perceive color, through some mechanism which is not completely understood. They have been seen to have the ability to assess their surroundings and match
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5208-418: Is thought to have up to 35 distinct signalling states. 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
5332-426: Is unknown, but chromatophores are under the control of neural pathways, allowing the cephalopod to coordinate elaborate displays. Together, chromatophores and iridophores are able to produce a large range of colors and pattern displays. Cephalopods utilize chromatophores' color changing ability in order to camouflage themselves. Chromatophores allow Coleoids to blend into many different environments, from coral reefs to
5456-400: Is used for multiple adaptive purposes in cephalopods, color change could have evolved for one use and the other developed later, or it evolved to regulate trade offs within both. Color change is widespread in ectotherms including anoles, frogs, mollusks, many fish, insects, and spiders. The mechanism behind this color change can be either morphological or physiological. Morphological change is
5580-417: Is usually mixed, upon expulsion, with mucus , produced elsewhere in the mantle, and therefore forms a thick cloud, resulting in visual (and possibly chemosensory) impairment of the predator, like a smokescreen . However, a more sophisticated behavior has been observed, in which the cephalopod releases a cloud, with a greater mucus content, that approximately resembles the cephalopod that released it (this decoy
5704-468: The Ammonoidea (ammonites) and Belemnoidea (belemnites). Extant cephalopods range in size from the 10 mm (0.3 in) Idiosepius thailandicus to the 700 kilograms (1,500 lb) heavy Colossal squid , the largest extant invertebrate . There are over 800 extant species of cephalopod, although new species continue to be described. An estimated 11,000 extinct taxa have been described, although
5828-658: The Ordovician period, represented by primitive nautiloids . The class now contains two, only distantly related, extant subclasses: Coleoidea , which includes octopuses , squid , and cuttlefish ; and Nautiloidea , represented by Nautilus and Allonautilus . In the Coleoidea, the molluscan shell has been internalized or is absent, whereas in the Nautiloidea, the external shell remains. About 800 living species of cephalopods have been identified. Two important extinct taxa are
5952-824: 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. Cuttlefish thick soup is a signature dish in Taiwan . 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), and in various recipes (either grilled or stewed) often served together with polenta . Catalan cuisine , especially that of
6076-462: The diffraction of light within the stacked plates. Orientation of the chemochromes 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
6200-567: The sparkling enope squid ( Watasenia scintillans ). It achieves color vision with three photoreceptors , which are based on the same opsin , but use distinct retinal 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,
6324-521: The Discosorida evolved directly from the Plectronoceratida rather than through the more developed Ellesmerocerida , as did the other orders. Finally and most diagnostic, discosorids developed a reinforcing, grommet -like structure in the septal opening of the siphuncle known as the bullette, formed by a thickening of the connecting ring as it draped around the folded back septal neck. The origin of
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#17328699602556448-674: The Discosorida is unknown, thought at one time to be directly from the Plectronocerida . Evolution within the order begins with the lower Middle Ordovician Reudemannoceratidae and from there diverges into three main lineages. Questionable discosorids have been reported as early as the Middle Tremadocian - near the start of the Ordovician, however the first bona fide examples date to the Middle Ordovician. The diversification of
6572-481: The Discosorida resembled the Oncocerida , which lived about the same time, but evolved from a completely different stock. The two convergent groups differ in their internal detail. Cephalopod A cephalopod / ˈ s ɛ f ə l ə p ɒ d / is any member of the molluscan class Cephalopoda / s ɛ f ə ˈ l ɒ p ə d ə / ( Greek plural κεφαλόποδες , kephalópodes ; "head-feet") such as
6696-572: The Discosorida, in terms of genera, peaked at the beginning in the Middle Ordovician (modern Darriwilian stage) followed by a decline in the Upper Ordovician (modern Sandbian and Katian stages) only to peak again in the Middle Silurian. Afterwards their diversity declined drastically and remained low until their end in the late Devonian. Some were endogastrically curved, with the lower, siphuncle side concave, others were exogastrically curved with
6820-487: The European common squid ( Alloteuthis subulata ) has yellow and red, and the common octopus has yellow, orange, red, brown, and black. In cuttlefish, activation of a chromatophore can expand its surface area by 500%. Up to 200 chromatophores per mm 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. Retracting
6944-556: The ability to determine color by comparing detected photon intensity across multiple spectral channels. When camouflaging 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 leucophores 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 has been found in
7068-449: The acidity of the organic shell matrix (see Mollusc shell ); shell-forming cephalopods have an acidic matrix, whereas the gladius of squid has a basic matrix. The basic arrangement of the cephalopod outer wall is: an outer (spherulitic) prismatic layer, a laminar (nacreous) layer and an inner prismatic layer. The thickness of every layer depends on the taxa. In modern cephalopods, the Ca carbonate
7192-434: The air for distances of up to 50 metres (160 ft). 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 , has been observed spreading tentacles in
7316-454: The appearance of their surroundings is notable given that cephalopods' vision is monochromatic. Cephalopods also use their fine control of body coloration and patterning to perform complex signaling displays for both conspecific and intraspecific communication. Coloration is used in concert with locomotion and texture to send signals to other organisms. Intraspecifically this can serve as a warning display to potential predators. For example, when
7440-417: The atmosphere, is cited as a potential threat. Some studies suggest that ocean acidification does not impair normal embryonic development, survival rates or body size. Unlike other cephalopods, cuttlefish possess a unique internal structure called the cuttlebone , a highly modified internal shell, which is porous and is made of aragonite . Except for spirula , they are the only coleoid cephalopods with
7564-408: 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
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#17328699602557688-403: The bloodstream. Cephalopods exchange gases with the seawater by forcing water through their gills, which are attached to the roof of the organism. Water enters the mantle cavity on the outside of the gills, and the entrance of the mantle cavity closes. When the mantle contracts, water is forced through the gills, which lie between the mantle cavity and the funnel. The water's expulsion through
7812-517: The body cavity; others, like some fish, accumulate oils in the liver; and some octopuses have a gelatinous body with lighter chloride ions replacing sulfate in the body chemistry. Squids are the primary sufferers of negative buoyancy in cephalopods. The negative buoyancy means that some squids, especially those whose habitat depths are rather shallow, have to actively regulate their vertical positions. This means that they must expend energy, often through jetting or undulations, in order to maintain
7936-527: The brain is unable to achieve both controlling elongation and controlling the chromatophores. Most octopuses mimic select structures in their field of view rather than becoming a composite color of their full background. Evidence of original coloration has been detected in cephalopod fossils dating as far back as the Silurian ; these orthoconic individuals bore concentric stripes, which are thought to have served as camouflage. Devonian cephalopods bear more complex color patterns, of unknown function. Coleoids,
8060-401: The capillaries of the gills . A single systemic heart then pumps the oxygenated blood through the rest of the body. Like most molluscs, cephalopods use hemocyanin , a copper-containing protein, rather than hemoglobin , to transport oxygen. As a result, their blood is colorless when deoxygenated and turns blue when bonded to oxygen. In oxygen-rich environments and in acidic water, hemoglobin
8184-436: The cavity by entering not only through the orifices, but also through the funnel. Squid can expel up to 94% of the fluid within their cavity in a single jet thrust. To accommodate the rapid changes in water intake and expulsion, the orifices are highly flexible and can change their size by a factor of twenty; the funnel radius, conversely, changes only by a factor of around 1.5. Some octopus species are also able to walk along
8308-561: The cell. By rapidly changing multiple chromatophores of different colors, cephalopods are able to change the color of their skin at astonishing speeds, an adaptation that is especially notable in an organism that sees in black and white. Chromatophores are known to only contain three pigments, red, yellow, and brown, which cannot create the full color spectrum. However, cephalopods also have cells called iridophores, thin, layered protein cells that reflect light in ways that can produce colors chromatophores cannot. The mechanism of iridophore control
8432-399: The cephalopod mantle have been widely used for many years as experimental material in neurophysiology ; their large diameter (due to lack of myelination ) 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 with fish. Some cephalopods are able to fly through
8556-474: 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. Furthermore,
8680-461: The chromatophores contain luminescent protein nanostructures in which tethered pigment granules modify light through absorbance, reflection, and fluorescence between 650 and 720 nm. 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 ( Doryteuthis opalescens ) have yellow, red, and brown,
8804-414: 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
8928-464: 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
9052-746: 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. 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
9176-403: The color, contrast and texture of the substrate even in nearly total darkness. 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. Sepia officinalis changes color to match
9300-465: The cuttlefish becomes texturally as well as chromatically similar to objects in its environment such as kelp or rocks. While the preferred diet of cuttlefish is crabs and fish, they feed on small shrimp shortly after hatching. 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
9424-411: The cuttlefish, but use the flamboyant display towards larger, more dangerous fish, and give no display at all to chemosensory predators such as crabs and dogfish. 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
9548-491: The depth of the ocean, from the abyssal plains to the sea surface, and have also been found in the hadal zone . 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). Cephalopods are widely regarded as the most intelligent of the invertebrates and have well developed senses and large brains (larger than those of gastropods ). The nervous system of cephalopods
9672-403: The egg. In consequence, they may prefer to hunt the prey they saw before hatching. 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. Though
9796-869: The end of the Cretaceous period, represented by Ceratisepia from the Late Maastrichtian Maastricht Formation of the Netherlands. Although the Jurassic Trachyteuthis was historically considered possibly related to cuttlefish, later studies considered it to be more closely related to octopuses and vampire squids. 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). They have an unusual biogeographic pattern; they are present along
9920-469: The expansion of the mantle at the end of the jet. In some tests, the collagen has been shown to be able to begin raising mantle pressure up to 50ms before muscle activity is initiated. These anatomical differences between squid and octopuses can help explain why squid can be found swimming comparably to fish while octopuses usually rely on other forms of locomotion on the sea floor such as bipedal walking, crawling, and non-jetting swimming. Nautiluses are
10044-422: 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. 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. On occasion,
10168-595: The final structure of the cephalopod eye fundamentally differ from those of vertebrates , such as humans. Superficial similarities between cephalopod and vertebrate eyes are thought to be examples of convergent evolution . The cuttlefish pupil is a smoothly curving W-shape. Although cuttlefish cannot see color, 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
10292-447: 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. Cuttlefish start to actively mate at around five months of age. Male cuttlefish challenge one another for dominance and
10416-413: The form of jetting. The composition of these mantles differs between the two families, however. In octopuses, the mantle is made up of three muscle types: longitudinal, radial, and circular. The longitudinal muscles run parallel to the length of the octopus and they are used in order to keep the mantle the same length throughout the jetting process. Given that they are muscles, it can be noted that this means
10540-456: The funnel can be used to power jet propulsion. If respiration is used concurrently with jet propulsion, large losses in speed or oxygen generation can be expected. The gills, which are much more efficient than those of other mollusks, are attached to the ventral surface of the mantle cavity. There is a trade-off with gill size regarding lifestyle. To achieve fast speeds, gills need to be small – water will be passed through them quickly when energy
10664-401: The largest species , the giant cuttlefish ( Sepia apama ), reaching 50 cm (20 in) in mantle length and over 10.5 kg (23 lb) in mass. Cuttlefish eat small molluscs, crabs, shrimp, fish, octopuses, worms, and other cuttlefish. Their predators include dolphins, larger fish (including sharks), seals, seabirds, and other cuttlefish. The typical life expectancy of a cuttlefish
10788-611: The low tide line and the edge of the continental shelf, to about 180 m (600 ft). 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
10912-477: 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. 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. The blood of
11036-472: The maximum diameter of the funnel orifice (or, perhaps, the average diameter of the funnel) and the diameter of the mantle cavity. Changes in the size of the orifice are used most at intermediate velocities. The absolute velocity achieved is limited by the cephalopod's requirement to inhale water for expulsion; this intake limits the maximum velocity to eight body-lengths per second, a speed which most cephalopods can attain after two funnel-blows. Water refills
11160-521: The mechanism is not understood. 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
11284-617: The muscles of the flamboyant cuttlefish ( Metasepia pfefferi ) contain a highly toxic, unidentified compound as lethal as the venom of fellow cephalopod, the blue-ringed octopus . However, this toxin is found only in the muscle and is not injected in any form, classifying it as poisonous, not venomous. Sleep is a state of immobility characterized by being rapidly reversible, homeostatically controlled, and increasing an organism's arousal threshold. To date one cephalopod species, Octopus vulgaris , has been shown to satisfy these criteria. Another species, Sepia officinalis , satisfies two of
11408-404: The non threatening herbivorous parrotfish to approach unaware prey. The octopus Thaumoctopus mimicus is known to mimic a number of different venomous organisms it cohabitates with to deter predators. While background matching, a cephalopod changes its appearance to resemble its surroundings, hiding from its predators or concealing itself from prey. The ability to both mimic other organisms and match
11532-563: The number of arms expressed. Cuttlefish Cuttlefish , or cuttles , are marine molluscs of the suborder Sepiina . 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 buoyancy . 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
11656-413: The octopus Callistoctopus macropus is threatened, it will turn a bright red brown color speckled with white dots as a high contrast display to startle predators. Conspecifically, color change is used for both mating displays and social communication. Cuttlefish have intricate mating displays from males to females. There is also male to male signaling that occurs during competition over mates, all of which are
11780-443: The octopus genus Argonauta secrete a specialized paper-thin egg case in which they reside, and this is popularly regarded as a "shell", although it is not attached to the body of the animal and has a separate evolutionary origin. The largest group of shelled cephalopods, the ammonites , are extinct, but their shells are very common as fossils . The deposition of carbonate, leading to a mineralized shell, appears to be related to
11904-444: The octopus must actively flex the longitudinal muscles during jetting in order to keep the mantle at a constant length. The radial muscles run perpendicular to the longitudinal muscles and are used to thicken and thin the wall of the mantle. Finally, the circular muscles are used as the main activators in jetting. They are muscle bands that surround the mantle and expand/contract the cavity. All three muscle types work in unison to produce
12028-497: The only extant cephalopods with a true external shell. However, all molluscan shells are formed from the ectoderm (outer layer of the embryo); in cuttlefish ( Sepia spp.), for example, an invagination of the ectoderm forms during the embryonic period, resulting in a shell ( cuttlebone ) that is internal in the adult. The same is true of the chitinous gladius of squid and octopuses. Cirrate octopods have arch-shaped cartilaginous fin supports , which are sometimes referred to as
12152-440: The other muscle fibers in the mantle. These collagen fibers act as elastics and are sometimes named "collagen springs". As the name implies, these fibers act as springs. When the radial and circular muscles in the mantle contract, they reach a point where the contraction is no longer efficient to the forward motion of the creature. In such cases, the excess contraction is stored in the collagen which then efficiently begins or aids in
12276-449: The patterns occur on a continuum. 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 . 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
12400-484: 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. The cuttlefish's eyes are thought to be fully developed before birth, and they start observing their surroundings while still in
12524-442: The posterior and anterior ends of this organ control the speed of the jet the organism can produce. The velocity of the organism can be accurately predicted for a given mass and morphology of animal. Motion of the cephalopods is usually backward as water is forced out anteriorly through the hyponome, but direction can be controlled somewhat by pointing it in different directions. Some cephalopods accompany this expulsion of water with
12648-407: The product of chromatophore coloration displays. There are two hypotheses about the evolution of color change in cephalopods. One hypothesis is that the ability to change color may have evolved for social, sexual, and signaling functions. Another explanation is that it first evolved because of selective pressures encouraging predator avoidance and stealth hunting. For color change to have evolved as
12772-431: The radial and circular mantle cavity muscles. The gills of cephalopods are supported by a skeleton of robust fibrous proteins; the lack of mucopolysaccharides distinguishes this matrix from cartilage. The gills are also thought to be involved in excretion, with NH 4 being swapped with K from the seawater. While most cephalopods can move by jet propulsion, this is a very energy-consuming way to travel compared to
12896-712: The reddish-brown color sepia in English. "Cuttle" in "cuttlefish", sometimes called "cuttles", is derived from the Old English name for the species, cudele . The word may be cognate with the Old Norse koddi (cushion) and the Middle Low German Kudel (rag). Over 120 species of cuttlefish are currently recognized, grouped into six families divided between two suborders. One superfamily and three families are extinct. The earliest fossils of cuttlefish are from
13020-441: The result of a change in the density of pigment containing cells and tends to change over longer periods of time. Physiological change, the kind observed in cephalopod lineages, is typically the result of the movement of pigment within the chromatophore, changing where different pigments are localized within the cell. This physiological change typically occurs on much shorter timescales compared to morphological change. Cephalopods have
13144-439: The result of natural selection different parameters would have to be met. For one, you would need some phenotypic diversity in body patterning among the population. The species would also need to cohabitate with predators which rely on vision for prey identification. These predators should have a high range of visual sensitivity, detecting not just motion or contrast but also colors. The habitats they occupy would also need to display
13268-424: The result of social selection the environment of cephalopods' ancestors would have to fit a number of criteria. One, there would need to be some kind of mating ritual that involved signaling. Two, they would have to experience demonstrably high levels of sexual selection. And three, the ancestor would need to communicate using sexual signals that are visible to a conspecific receiver. For color change to have evolved as
13392-549: The same class. Octopuses are generally not seen as active swimmers; they are often found scavenging the sea floor instead of swimming long distances through the water. Squids, on the other hand, can be found to travel vast distances, with some moving as much as 2000 km in 2.5 months at an average pace of 0.9 body lengths per second. There is a major reason for the difference in movement type and efficiency: anatomy. Both octopuses and squids have mantles (referenced above) which function towards respiration and locomotion in
13516-399: The same depth. As such, the cost of transport of many squids are quite high. That being said, squid and other cephalopod that dwell in deep waters tend to be more neutrally buoyant which removes the need to regulate depth and increases their locomotory efficiency. The Macrotritopus defilippi , or the sand-dwelling octopus, was seen mimicking both the coloration and the swimming movements of
13640-528: The same side convex. In some, the aperture was a simple opening. In others, it became contracted into a pattern of slits. In earlier, Ordovician forms, the bullette became quite large and readily noticeable. In later forms, the bullette became reduced, in some to the point of being vestigial. The Discosoridae, one of the last families to evolve, found in Silurian and questionably in Devonian rocks, are characterized by
13764-414: 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. 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
13888-452: The sand-dwelling flounder Bothus lunatus to avoid predators. The octopuses were able to flatten their bodies and put their arms back to appear the same as the flounders as well as move with the same speed and movements. Females of two species, Ocythoe tuberculata and Haliphron atlanticus , have evolved a true swim bladder . Two of the categories of cephalopods, octopus and squid, are vastly different in their movements despite being of
14012-446: The sandy sea floor. The color change of chromatophores works in concert with papillae, epithelial tissue which grows and deforms through hydrostatic motion to change skin texture. Chromatophores are able to perform two types of camouflage, mimicry and color matching. Mimicry is when an organism changes its appearance to appear like a different organism. The squid Sepioteuthis sepioide has been documented changing its appearance to appear as
14136-489: The seabed. Squids and cuttlefish can move short distances in any direction by rippling of a flap of muscle around the mantle. While most cephalopods float (i.e. are neutrally buoyant or nearly so; in fact most cephalopods are about 2–3% denser than seawater ), they achieve this in different ways. Some, such as Nautilus , allow gas to diffuse into the gap between the mantle and the shell; others allow purer water to ooze from their kidneys, forcing out denser salt water from
14260-481: The sex of a cuttlefish in the absence of sexual dimorphism . Female cuttlefish signal their receptivity to mating using a display called precopulatory grey. 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. Displays on one side of
14384-597: The siphuncle near the inside or longitudinally concave curvature. The second, formed by the Westonoceratidae, Lowoceratidae, and Discosoridae, are fundamentally exogastric with the siphuncle near the outside or longitudinally convex curvature, although the Discosoridae are somewhat different. The third, consisting of the Mandaloceratidae and Mesoceratidae are basically straight (orthoconic). Families differ primarily in
14508-601: 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 , but the brief squid, Lolliguncula brevis , found in Chesapeake Bay , is a notable partial exception in that it tolerates brackish water . 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
14632-440: The species and for warning ) or active camouflage , as their chromatophores are expanded or contracted. Although color changes appear to rely primarily on vision input, there is evidence that skin cells, specifically chromatophores , can detect light and adjust to light conditions independently of the eyes. The octopus changes skin color and texture during quiet and active sleep cycles. Cephalopods can use chromatophores like
14756-536: The structural details of the siphuncle and in the nature of the aperture. Discosorids were probably benthic forms that crawled over the bottom in search of food or safety, or hovered close to the bottom. The general orientation during life was most likely head down, with the aperture of the shell facing the general direction of the sea floor and shell carried above. Nothing is known of what the animal itself may have looked like; how many tentacles they had and relative length or how well they may have seen. In general form
14880-409: 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
15004-454: The tail propulsion used by fish. The efficiency of a propeller -driven waterjet (i.e. Froude efficiency ) is greater than a rocket . The relative efficiency of jet propulsion decreases further as animal size increases; paralarvae are far more efficient than juvenile and adult individuals. Since the Paleozoic era , as competition with fish produced an environment where efficient motion
15128-505: The three criteria but has not yet been tested on the third (arousal threshold). Recent research shows that the sleep-like state in a common species of cuttlefish, Sepia officinalis , shows predictable periods of rapid eye movement, arm twitching and rapid chromatophore changes. The lifespan of a cuttlefish is typically around one to two years, depending on the species. They hatch from eggs fully developed, around 6 mm ( 1 ⁄ 4 in) long, reaching 25 mm (1 in) around
15252-463: Was crucial to survival, jet propulsion has taken a back role, with fins and tentacles used to maintain a steady velocity. Whilst jet propulsion is never the sole mode of locomotion, the stop-start motion provided by the jets continues to be useful for providing bursts of high speed – not least when capturing prey or avoiding predators . Indeed, it makes cephalopods the fastest marine invertebrates, and they can out-accelerate most fish. The jet
15376-401: Was published indicating that cephalopod chromatophores are photosensitive; reverse transcription polymerase chain reactions (RT-PCR) revealed transcripts encoding rhodopsin and retinochrome within the retinas and skin of the longfin inshore squid ( Doryteuthis pealeii ), and the common cuttlefish ( Sepia officinalis ) and broadclub cuttlefish ( Sepia latimanus ). The authors claim this
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