cephalopod

/sef"euh leuh pod'/, n.

1. any mollusk of the class Cephalopoda, having tentacles attached to the head, including the cuttlefish, squid, and octopus.

adj.

2. Also, cephalopodic, cephalopodous /sef'euh lop"euh deuhs/. belonging or pertaining to the Cephalopoda.

[1820-30; < NL Cephalopoda; see CEPHALO-, -POD]

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Any marine mollusk of the class Cephalopoda (e.g., cuttlefish, nautilus, octopus, and squid), which includes the most active and largest living invertebrates.

Cephalopods are bilaterally symmetrical and typically have a highly developed centralized nervous system. Their image-forming eyes are similar in structure to vertebrate eyes, and their heads are armed with tentacles that have rows of round suction disks. Most cephalopods can change skin colour to blend in with their surroundings. All can swim, propelling themselves backward by expelling water forcefully. Most are carnivores that feed on fish, crustaceans, and other mollusks.

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▪ class of mollusks

Introduction

 any member of the class Cephalopoda, of the phylum Mollusca, a small group of highly advanced and organized, exclusively marine animals. The octopus, squid, cuttlefish, and chambered nautilus are familiar representatives. The extinct forms outnumber the living, the class having attained great diversity in late Paleozoic and Mesozoic times. The extinct cephalopods are the ammonites, belemnites, and nautiloids, except for five living species of Nautilus.

General features and importance to humans

      The cephalopods agree with the rest of the Mollusca in basic structure, and the ancestors appear to have the closest affinity with the ancestors of the class Gastropoda. The best-known feature of the cephalopods is the possession of arms and tentacles, eight or 10 in most forms but about 90 in Nautilus. Except for the nautilus, all living members of the class show great modification and reduction of the characteristic molluscan shell.

      Cephalopods range greatly in size. The giant squids (Architeuthis species) are the largest living invertebrates; A. dux attains a length of more than 20 metres (60 feet), including the extended tentacles. The smallest cephalopod is the squid Idiosepius, rarely an inch in length. The average octopus usually has arms no longer than 30 centimetres (12 inches) and rarely longer than a metre (39 inches). But arm spans of up to nine metres (30 feet) have been reported in Octopus dofleini. The shell of the fossil ammonite Pachydiscus seppenradensis from the Cretaceous measures 205 centimetres (6 feet 8 inches) in diameter; it is considered to have been the largest shelled mollusk.

      Cephalopods occur in large numbers and form one of the greatest potential food resources of the oceans. They are eaten in most parts of the world and have been accepted as part of the general diet in North America and northern Europe. They also are indirectly important to humans since they furnish a large part of the diet of sperm whales and smaller whales, seals, fishes, and seabirds.

Natural history

Reproduction and life cycles

      The sexes are usually separate in the Cephalopoda. Sexual dimorphism is usually expressed in slight differences of size and in the proportions of various parts. In the argonaut and the blanket octopus (Tremoctopus) the males differ in appearance and size from the females.

      The female reproductive system is simple, consisting of the posterior ovary and paired oviducts. Nidamental glands exist in species that lay eggs encased in heavy gelatinous capsules.

      In males the reproductive system contains a series of chambers or sacs along the course of the vas deferens, which produce long tubes (spermatophores) to contain the spermatozoa. The final sac (Needham's organ) is used for storage of spermatophores. The spermatophores are complicated, containing sperm reservoir, cement body, cap, and a delicate triggering mechanism for releasing the tube and cementing it to the female's body, where the sperm are released when the eggs are mature and ready to be laid. Since spermatophores vary in appearance from species to species, they are important taxonomic characters.

      During courtship the male deposits spermatophores in the female, either within the mantle cavity or on a pad below the mouth, by means of a specially modified arm, the hectocotylus. The hectocotylized arm of Octopus bears a deep groove on one side, ending in a spoonlike terminal organ. In Argonauta and Tremoctopus the arm is highly modified and in mating is autotomized (autotomy) (self-amputated) and left within the mantle cavity of the female. In the squids a much larger section of the arm may be modified; often the suckers are degenerate and the distal half of the arm bears rows of slender papillae, although special pouches and flaps may often be found. The modified arm of Nautilus is termed the spadix.

      Little is known about the mating habits of most cephalopods. In the common octopus the male and female remain some distance apart while the male caresses the female with the tip of the hectocotylized arm. The male then inserts the tip of the arm into the mantle cavity of the female, where it remains for more than an hour, during which time the spermatophores travel down the spermatophoral groove of the arm. In the cuttlefish (Sepia), according to the Dutch zoologist L. Tinbergen, the pair swims side by side, the male indulging in some courtship behaviour with its arms. Eventually, mating takes place by the pair intertwining their arms and remaining together while the spermatophores are placed on the inner side of the female's mouth membrane. In loliginid squids a somewhat similar type of mating occurs, except that it takes place en masse in schools of thousands of individuals.

      Eggs (egg) may be laid shortly after mating or after a prolonged period of maturation during which time the sperm remain viable. In loliginids they are fertilized as they are ejected and before being fixed in the egg capsule. In the octopods they may be fertilized as they pass through oviductal glands near the end of the oviduct. In cuttlefishes the eggs are fertilized before the heavy capsule is formed. Egg laying in octopods is accomplished by the female individually fixing the eggs singly or in festoons by a short stalk or thread. In loliginids the eggs in fingerlike capsules often form immense moplike patches, the result of the communal spawning of perhaps hundreds of individuals. Spawning of oceanic squids is very poorly known. The number of eggs laid during a spawning period varies greatly; it may range from only a few dozen in octopuses with large eggs to more than 100,000 in the common octopus, laid over a period of about two weeks. In cuttlefishes the number of eggs is smaller, about 200 to 300 being laid in a season. In loliginids several thousand eggs may be laid by a single female, and the egg mop of the European common squid, resulting from the efforts of many individuals, may contain more than 40,000 eggs.

      The eggs of most cephalopods are enclosed within a capsule that may be gelatinous and transparent (the squids of the genus Loligo) or opaque and leathery (Octopus and cuttlefishes). The eggs of oceanic species may be laid in large sausagelike gelatinous masses or singly. The eggs of most coastal species are laid inshore and are attached singly or in clusters, primarily to rocks and shells on the bottom. Parental care is exhibited by some octopuses, in which the female broods over the eggs in the den, and in the argonaut (Argonauta), in which the eggs are carried in a special shell secreted by the female. In most squids and cuttlefishes the eggs are left uncared-for. Squids that attach their eggs to the bottom engulf them in a gelatinous mass that protects them from disease and deters predators. Cuttlefishes squirt their eggs with ink when they are laid to camouflage the otherwise white eggs.

      All cephalopod eggs have a remarkable amount of yolk, unlike that in the rest of the Mollusca, so that segmentation is incomplete and restricted to one end of the egg, where the embryo develops. The embryo of a cuttlefish (Sepia), squid (Loligo), or octopus (Octopus) has a yolk sac. In certain presumably archaic Teuthoidea there is less yolk, and the yolk sac is nearly absent. Development of the embryo is direct, without the distinctive larval stages and metamorphoses that occur in other mollusks.

      Incubation time varies, but in Octopus young hatch in about 50 days and in Loligo in about 40 days. At hatching, the young may closely resemble the adult and assume the adult habitat or they may differ from the adult and spend a considerable time in the plankton as part of the drifting life. The juveniles of many cephalopods were described as distinct genera before their juvenile status was discovered. In octopods with small eggs (e.g., Octopus vulgaris) the juveniles are planktonic, spending several weeks in the plankton; the “Macrotritopus” stage of Scaeurgus may greatly prolong its juvenile life until a favourable bottom substrate is found. In octopods with large eggs (e.g., Octopus briareus) the young resemble the adult and immediately assume a bottom-dwelling mode of life.

      In the order Sepioidea (cuttlefishes and bottle-tailed squids) the young closely resemble the adults and are only briefly planktonic. In the Teuthoidea (squids), especially the Oegopsida, the larvae may differ widely from the adult and the juvenile period may be quite long.

      Little is known about the life span of cephalopods. Studies have shown that in Octopus joubini raised from the egg in aquariums, sexual maturity and spawning were reached in five months; in a loliginid squid (Sepioteuthis sepioidea), likewise raised from the egg, sexual maturity and full growth were also attained in five months. It thus appears that the smaller inshore species may have a life span of no more than one year or, exceptionally, two or three. Nothing is known of the life span of the large oceanic squids, but it is presumed that giants such as Architeuthis attain their bulk only after a period of perhaps four to five years. In the smaller octopuses and squids, observational data indicate that many of the males die after mating and females after the first major spawning.

Behaviour (animal behaviour)

      Cephalopods are unique among the invertebrates in the degree of cephalization and cerebralization attained. The uniting of the major ganglionic centres of the central nervous system constitutes a brain of considerable complexity. Studies undertaken at the Zoological Station in Naples by the British zoologists J.Z. Young, Martin J. Wells, and others have demonstrated that Octopus is capable of learning (animal learning) and has considerable intelligence. The behaviour of squids and octopuses differs considerably because of their different modes of life. Laboratory behavioral studies have dealt mainly with learning processes and have centred around food acceptance, reward and punishment, maze work, and shape discrimination. By means of surgical techniques it has been possible to determine the various functional centres of the brain of Octopus and the transmission and receiving pathways.

      Research of a detailed nature has been concerned with colour change. Most cephalopods possess colour pigment cells (chromatophores (chromatophore)) and reflecting cells (iridocytes) in the skin. The chromatophores are expanded by nerves controlled by the brain, and the colours are exposed (brown, black, red, yellow, or orange red). Colours and colour patterns are exhibited according to specific behavioral (avoidance behaviour) conditions—e.g., attack on prey, camouflage, rest, and alarm (alarm signal) or defense. Alarm patterns are the most readily recognized, consisting of strong contrasting light and dark areas, bars and peripheral dark outlines, or vivid displays of spots, like huge eyes.

      Other behavioral patterns are found in changes in skin texture, including the erection of branched or spikelike papillae and curling of the arms. These actions often are attempts by cephalopods to conceal or camouflage themselves through imitating bottom objects such as sand, coral, or seaweed.

      The ink of cephalopods is used for both defense and escape. In Octopus under attack by a moray eel, the cloud of ink seems to paralyze for some time the eel's senses of sight and smell. In squids the ink is ejected as a spindle-shaped mass about the size of the squid itself, the ink coagulating in the water. With this “dummy” left behind, the squid contracts its chromatophores, becomes nearly transparent, and jets away.

      Many cephalopods (but not Nautilus and Octopus) possess special light organs (photophores (photophore)), which emit chemical light or bioluminescence. Light is produced by the enzymatic reaction of luciferin and luciferase or, in bottle-tailed squids (sepiolids), indirectly, through cultures of luminescent bacteria. Photophores distributed over the body are employed at night or in the mid depths in various ways: mating play, recognition of the sexes, aid in schooling, attracting prey, defense, and camouflage. The light organs of the squid Histioteuthis are highly complicated, consisting of reflector, light source, directive muscles, lens, diaphragm, window, and colour screens.

      Octopuses, squids, and cuttlefishes display considerable skill and cunning in hunting, stalking patiently, or luring prey (predation) within reach of their arms or tentacles. Both cuttlefishes and octopuses may use the tips of their arms as wormlike lures to attract small fishes, and octopuses have been reported to thrust stones between the valves of clams to prevent their closing. This has not been verified by later observers, but such intelligence is not beyond belief.

Locomotion

      Cephalopods move by crawling, swimming, or jet propulsion, mainly the latter. The mantle, which has a passive role in the majority of mollusks, has become involved in locomotion in cephalopods, having almost entirely lost its rigid shell and become highly muscular. Its expansion and contraction produce a locomotory water current by drawing water into the mantle cavity and expelling it through the funnel. The rapid ejection of this jet of water enables the animal to execute quick backward and forward movements.

      Water is drawn into the mantle cavity by the relaxation of the circular muscles and resultant expansion of the mantle. It enters around the neck region or aperture of the mantle (through the funnel in some deep-sea octopuses). In the oceanic squids the system is more efficient, with a nonreturn valve that prevents water from entering the wrong way through the funnel. When the mantle is contracted the aperture is closed by locking mechanisms and contraction of the anterior ring muscle, and the water is forced out through the funnel. This flexible tube is constructed similarly to a jet nozzle and may be turned in any direction, giving the animal great flexibility of motion for steering.

      Squids also possess terminal or lateral fins used in slow movement or hovering. Locomotion is by the rapid undulation of the outer edges of the fins. Movement through the water is aided by lateral expansions (swimming keels) on the outer surface of the third pair of arms. Some squids (Onychoteuthis, Thysanoteuthis) are able to “fly” for several hundred feet, driven into the air by powerful thrusts from their jets and gliding on their expanded fins and arm keels. This normally occurs when the squids are pursued by predatory fishes and dolphins. When the squid jets rapidly through the water, the fins are often curled tightly around the mantle. Cuttlefishes, because of the large cuttlebone, are less active animals and spend most of their life lying on or hovering slightly above the bottom. Both jet and fins are used, the latter more frequently.

      The benthic octopods are bottom crawlers, gliding about the bottom and in and out of crannies with amazing agility through the use of their eight arms and hundreds of suckers. Because of the extraordinary flexibility of the body, they are able to pass through openings hardly larger than the diameter of one eye. When disturbed, octopods swim rapidly by jet propulsion. Mass swimming migrations have been reported.

      The finned octopods and the bottle-tailed squids have paddle-shaped fins that probably are most useful for hovering and slow swimming. The female argonaut (Argonauta) propels herself by jet propulsion while encased in her paper-thin shell. The chambered nautilus (Nautilus) is a less active swimmer, partly due to the inefficient funnel, which is composed of two flaps instead of a single fused tube, and to a reduced mantle cavity.

      The active swimmers and most bottom dwellers apparently possess no hydrostatic organ. The cuttlefish, however, which swims or hovers above the bottom or rests on the bottom, adjusts its buoyancy through the amount of gases contained in the porous cuttlebone. Nautilus, which swims slowly above the bottom or in midwater, accomplishes this similarly, adjusting the gases in the chambered shell. Inactive oceanic squids, such as some cranchiids, concentrate ions lighter than seawater in the body chamber, while others, such as Bathyteuthis, concentrate buoyant oil in the chambers associated with the digestive gland.

Ecology

      The Cephalopoda are exclusively marine animals. There are numerous littoral species, but few have been reported from brackish water except for the squid Lolliguncula brevis, which occurs along the Florida coast in bays where the salinity is as low as 8.5 parts per thousand (about one-fourth that of the open ocean). Cephalopods are excluded from the Baltic Sea by lower salinities but have been found in areas of the Suez Canal where salinities are higher than in the oceans. There is some evidence that in parts of the open sea that differ only slightly in salinity there may be a varying cephalopod fauna. Grace E. Pickford, an American zoologist, has found that the vertical distribution of young in Vampyroteuthis is governed by water density, and, according to the American zoologist Clyde F.E. Roper, a similar situation exists in the deep-sea squid Bathyteuthis.

      The Coleoidea are carnivorous and live principally on other cephalopods, crustaceans, mollusks, and small fishes. In the food cycle of the squid Illex, in which the adults feed upon young mackerel and the adult mackerel feed upon young squid. Squids also are cannibalistic. Octopus feeds upon bivalve mollusks and on decapod crustaceans, sometimes causing severe losses to the lobster fishery by entering the traps and eating the captive lobsters. The smaller oceanic squids probably feed primarily upon small fish, copepods, heteropods, and caridean shrimp. The Cirrata, which have reduced musculature and radula, indicating reduced activity and masticatory power, probably feed on bottom dwellers or small plankton.

      The Cephalopoda are fed upon by many marine mammals, large fish, and seabirds. Sperm whales and other toothed whales feed primarily on squids.

      The 19th-century French naturalist Alcide d'Orbigny (Orbigny, Alcide Dessalines d') asserted that the Cephalopoda are in general sociable (Social behaviour in animals), and this statement is certainly true of Nautilus, which are found together in large numbers. Some authorities, studying the Mediterranean cephalopods, have concluded that only certain pelagic cephalopods are gregarious (Todarodes, Ocythoe). Schools of the large oceanic squid Thysanoteuthis rhombis have been reported at Madeira. Although there is no proof that numbers constitute gregariousness, octopus colonies have been reported.

      The breeding season has a marked effect on the local distribution of certain cephalopods. The common cuttlefish comes into shallow water in the spring and summer to breed, and similar migrations have been observed in some squids (Loligo, Alloteuthis, Illex).

      The geographic distribution of cephalopods is incompletely known. Some open-ocean pelagic and bathypelagic forms are cosmopolitan in warm and temperate waters (Onychoteuthis banksi, Cranchia scabra). Others may be limited by genera to particular oceans or to continental waters. Even some species of bathypelagic habitat are limited to one ocean. The Octopoda, as a result of their bottom-dwelling habits, show stronger restrictions in their distribution, but Octopus vulgaris and O. macropus, both species with planktonic larvae, have gained worldwide distribution. In general, the pelagic and planktonic cephalopods conform in their distribution to other pelagic animals.

      The vertical distribution is also incompletely known. Nautilus moves vertically through the water, living near the bottom, and has been obtained at a depth of about 550 metres (1,800 feet). It is fished in the Philippines when it comes into shallow water. Of the cuttlefishes, the Sepiidae are littoral, whereas the Sepiolidae dwell on or near the bottom, down to considerable depths. Among the squids the Myopsida are coastal forms, whereas the Oegopsida are oceanic, living from the surface to depths in excess of 5,000 metres (16,400 feet). The Octopoda occur from the surface of the open ocean (Tremoctopus) to the ocean floor (Pareledone, Bentheledone, Cirroteuthis) in excess of 5,000 metres (16,400 feet); most are bottom-dwelling forms restricted to the continental shelf and its slope.

Form and function

      Cephalopods vary from elongate, streamlined oceanic organisms to saccular, slow-moving bottom and drifting forms. Their body plan is indicative in many ways of the habitat in which they dwell and their mode of life.

      The viscera of a generalized cephalopod are covered by a dome-shaped or elongated sheath of muscle, the mantle, which is connected with the head anteriorly. Ventrally, the mantle is free and encloses the mantle cavity, the space into which the gills project and the excretory and reproductive systems open.

      Anterior to the mantle is the head-foot, which bears a ventral muscular tube, the funnel. Surrounding the mouth are eight long, prehensile arms, provided with suckers in octopuses and argonauts (order Octopoda), or eight arms and two tentacles, equipped with horny ringed suckers bearing teeth or hooks in squids and cuttlefishes (orders Teuthoidea and Sepioidea). The primitive Nautilus has about 90 small suckerless appendages.

      The fossil nautiloids and ammonites (represented today only by Nautilus) were primitive, less-specialized forms, probably leading a rather inactive sluggish life. The modern octopuses, squids, and cuttlefishes have acquired an active, vigorous life that has led to marked departures in structure and function from the type represented by Nautilus. Modern forms are divided into three basic life-styles: the sluggish life in the great depths of the sea, a floating life in the midwaters, and a more active, aggressive existence near the surface. The nautiloids and ammonites were probably shallow-water animals living near the bottom and, like the slow-moving Nautilus, relied for protection on a coiled or curved calcareous external shell.

      All cephalopods have an internal cartilaginous covering of the consolidated ganglia of the nervous system. In all except ammonites and nautiloids, it constitutes a cranium. Various other skeletal supports are found at the base of the fins and in the “neck,” gills, and arms.

      The alimentary system consists of a buccal mass with a pair of jaws (mandibles) and a rasping tongue (radula), esophagus, salivary glands, stomach, cecum, digestive gland (“liver”), intestine, and anus. In the Octopoda the esophagus may be expanded to form a crop, while in the deep-sea Cirrata the radula may be degenerate or lacking. Except in Nautilus and certain deep-sea Octopoda there is an ink sac located near the anus. This secretes a dark fluid, the sepia or ink, which is forcibly ejected through the funnel.

      All members of the Coleoidea (octopuses, squids, and cuttlefishes) possess a closed circulatory system of blood vessels; in Nautilus it is partly lacunar (i.e., made up of sinuses). The blood contains a blue respiratory pigment, hemocyanin (a copper compound), dissolved in the plasma. There are three hearts (heart), the main systemic heart and the two branchial hearts, one at the base of each gill. The rhythmical contractions and expansions of the mantle cause a circulation of water over the gills where gas exchange takes place between the seawater and blood. The featherlike gills, consisting of a central axis with a row of lamellae on either side, are suspended in the mantle cavity from the mantle wall.

      The excretion of nitrogenous wastes is carried out exclusively by the kidneys. There are four kidneys in Nautilus and two in the coleoids.

      The central nervous system is highly developed, the major ganglionic centres being concentrated in the head. In some of the cuttlefishes (Sepioidea) the cerebral centres are subdivided for specialization, with the pedal ganglia divided into brachial and epipodal elements that innervate the arms and funnel respectively. In active squids the mantle is innervated by giant paired dorsal axons (axon). Much of the present knowledge of mechanisms of nerve impulse conduction has come from the study of these giant axons. The sense organs of the cephalopods are eyes (eye, human), rhinophores (olfactory organs), statocysts (organs of equilibrium), and tactile organs. In Nautilus the eyes are open pits without lenses. In the Coleoidea the eyes are complex and approach those of some lower vertebrates in efficiency.

Evolution and paleontology

      The first cephalopods were probably provided with a simple, caplike shell. With elongation of the shell and the formation of septa or partitions, the nautiloid shell could be formed (Late Cambrian, 523 to 505 million years ago). The primitive elongate shell of Orthoceras became unmanageable and coiling resulted, as in the Gastropoda.

      The Ammonoidea (ammonoid) are usually considered to have evolved from Devonian (408 to 360 million years ago) straight-shelled forms ( Bactrites) with certain nautiloid traits. Coiled ammonites appeared in the Late Devonian epoch (Goniatites). The subclass became extinct in the Cretaceous.

      Modern cephalopods possess an internal and partly degenerate shell, straight except in Spirula. The state of the shell in modern forms is due to the progressive overgrowth of it by the mantle, probably accompanying the evolution of an active swimming life. The first evidence of the modification of the shell is in Aulococeras in the Triassic period (245 to 208 million years ago). The belemnites, with their modified, internal shell, gave rise to Spirula (coiled shell), to cuttlefishes (calcified phragmoconal septa, forming a thick shell), and to the squids (only the proostracum as a horny “pen”). These forms appeared in the Jurassic period (208 to 144 million years ago) and were probably derived from belemnite-like ancestors.

      In the Octopoda the shell persists as cartilaginous stylets or fin supports. Palaeoctopus newboldi, the oldest known octopod, from the Cretaceous of Syria, was already too advanced to provide a clue to the derivation of the Octopoda. The Vampyromorpha are considered to be a possible connecting link between the Teuthoidea and the Octopoda.

Classification

Distinguishing taxonomic features

      In fossil cephalopods, reliance is placed upon shell details (general shape, type of coiling, external sculpture, and sutures). In living forms, except for the Sepioidea, the shell is strongly degenerate or missing and the characters used consist of details of the soft parts: presence or absence of an eyelid, tentacles retractile or contractile or both, shape and size of fins, number of arms, number of sucker rows, presence or absence of teeth and hooks on arm and tentacular suckers, radular dentition, structure of funnel organ, spermatophores, details of the hectocotyles, number of gill lamellae, and the presence, patterns, and types of photophores.

Annotated classification

      The following classification has gained considerable acceptance among modern specialists. Groups indicated by a dagger (†) are known only as fossils.

Class Cephalopoda (cephalopod)

 Mollusks in which typical molluscan foot surrounds head and forms arms and tentacles; mantle surrounds mantle cavity and is part of locomotory system; central nervous system highly developed, forming true brain encased in cartilaginous cranium; mouth contains pair of parrotlike jaws (or beak); body usually somewhat streamlined; eyes highly developed, most closely resembling in acuity those of some vertebrates; about 650 living species.

      Subclass Nautiloidea (nautiloids)

 Cambrian to present; now living only in the Indo-Pacific region, particularly East Indies; external coiled or straight chambered shell present, chambers connected by median siphuncle; smooth septa; sutures simple, little or no external sculpture; tentacles suckerless, adhesive; living and supposedly fossil forms with 4 gills; funnel formed of 2 nonfused flaps; about 5 living species, in genus nautilus.

      †Subclass Ammonoidea (ammonoid) (ammonites)

 Devonian to Cretaceous; fossils only; external, coiled or straight chambered shell with marginal siphuncle, last chamber protected by single horny plate or 2 calcareous plates; septa wrinkled; complex sutures; external sculpture.

      Subclass Coleoidea (octopuses, squids, belemnites, cuttlefishes)

 Triassic to present; shell internal, reduced, vestigial, or lacking; 2 sets of gills; 8 or 10 arms, having suckers or hooks.

      †Order Belemnoidea (belemnoid) (belemnites)

 Triassic to Tertiary; fossils only; shell consisting of solid rostrum, small chambered phragmocone and anterior, broad proostracum; 6 to 10 arms bearing hooks in 1 or 2 rows; total length 5 to 210 cm.

      Order Sepioidea (cuttlefishes (cuttlefish) and bottle-tailed squids)

 Tertiary to present; worldwide with family exceptions; shell coiled and chambered (Spirulidae), straight with vestigial chambering (Sepiidae), vestigial, or lacking; eyes covered with transparent membrane; 8 sucker-bearing arms and 2 tentacles retractile into pockets; total length 2.5–90 cm.

      Order Teuthoidea (squids (squid))

 Tertiary to present; shell thin, horny gladius; 8 arms, 2 tentacles, which are contractile only; worldwide; total length 1.5 to at least 1,800 cm (0.75 in. to 60+ ft).

      Suborder Myopsida

 Eye covered by transparent membrane; neritic, inshore animals.

      Suborder Oegopsida

 Eye open to water, completely surrounded by free eyelid; open-ocean animals living from the surface down to at least 3,000 m.

      Order Vampyromorpha

 Purplish-black gelatinous animals with 1 or 2 pairs of paddle-shaped fins at various stages of growth; 8 arms and 2 small retractile filaments not homologous with tentacles; deep web between the arms; worldwide; 1 species.

      Order Octopoda (octopuses (octopus))

 Cretaceous to present; shell lacking or vestigial, nonhomologous shell or egg case in female argonautids; fins absent or present; body generally saccular, with 8 mobile, highly contractile sucker-bearing arms; worldwide; total length 5–540 cm (2 in. to 18 ft); maximum arm spread to about 900 cm (30 ft).

      Suborder Palaeoctopoda (finned octopod)

 Cretaceous, some living.

      Suborder Cirrata (Cirromorpha)

 Holocene; soft-bodied, deep-webbed forms with cirri on arms and small to large paddle-shaped fins; primarily deep-sea.

      Suborder Incirrata (common octopus)

 Holocene; compact, saccular to round bodied, finless forms with muscular, contractile arms; somewhat secretive; pelagic to deep-sea and shallow waters.

Critical appraisal

      The elucidation of higher classification of the cephalopods is fraught with difficulties. Early specialists divided the living cephalopods into Octopoda and Decapoda without relation to their internal structure; these were both placed in the Dibranchia, in contrast to all fossil forms, which were considered as Tetrabranchia because Nautilus has four gills rather than two. This unnatural classification, accepted by the French zoologist Alcide d'Orbigny in 1838, was gradually modified through the efforts of the Swiss zoologist Adolph Naef and the German zoologist Georg Grimpe and later workers to the form given above. Phylogenetic linkages are still highly theoretical, but, with X-ray techniques that reveal soft parts in fossils and with new paleontological finds, the unfolding of the evolutionary history is in sight.

Gilbert L. Voss Clyde F.E. Roper

Additional Reading

General treatises on the biology, evolution, ecology, physiology, and behaviour of the cephalopod group include Marion Nixon and J.B. Messenger (eds.), The Biology of Cephalopods (1977); P.R. Boyle (ed.), Cephalopod Life Cycles, 2 vol. (1983–87); and Clyde F.E. Roper, Michael J. Sweeney, and Cornelia E. Nauen, Cephalopods of the World: An Annotated and Illustrated Catalogue of Species of Interest to Fisheries (1984), which also has an illustrated glossary and keys to identification of orders and families. K.N. Nesis, Cephalopods of the World: Squids, Cuttlefishes, Octopuses, and Allies (1987; originally published in Russian, 1982), is a review of anatomy, with keys to the identification of species. W. Bruce Saunders and Neil H. Landman (eds.), Nautilus: The Biology and Paleobiology of a Living Fossil (1987), is a comprehensive, scholarly treatise on the evolution, distribution, ecology, physiology, biology, shell structure, and aquarium maintenance of Nautilus. Peter Douglas Ward, The Natural History of Nautilus (1987), compares the growth, buoyancy, physiology, and ecology of Nautilus with those of other living cephalopods. Clyde F.E. Roper and Kenneth J. Boss, “The Giant Squid,” Scientific American, 246(4):96–100, 104–105 (April 1982), is a nontechnical summary of current knowledge. M.J. Wells, Octopus: Physiology and Behaviour of an Advanced Invertebrate (1978), is a highly technical but very readable review.

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