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The Frilled Shark (Chlamydoselachus anguineus) is one of two extant species of shark in the family Chlamydoselachidae, with a wide but patchy distribution in the Atlantic and Pacific Oceans. This uncommon species is found over the outer continental shelf and upper continental slope, generally near the bottom though there is evidence of substantial upward movements. It has been caught as deep as 1,570 m (5,150 ft), whereas in Suruga Bay, Japan it is most common at depths of 50–200 m (160–660 ft). Exhibiting several "primitive" features, the frilled shark has often been termed a "living fossil". It reaches a length of 2 m (6.6 ft) and has a dark brown, eel-like body with the dorsal, pelvic, and anal fins placed far back. Its common name comes from the frilly or fringed appearance of the gill slits, of which there are six pairs with the first pair meeting across the throat.

Seldom observed, the frilled shark is speculated to capture its prey by bending its body and lunging forward like a snake. The long, extremely flexible jaws enable it to swallow large prey whole, while the many rows of small, needle-like teeth prevent escape. It feeds mainly on cephalopods, while also consuming bony fishes and other sharks. This species is aplacental viviparous: the embryos emerge from their egg capsules inside the mother's uterus, and are sustained to term primarily by yolk. The gestation period may be as long as three and a half years, the longest of any vertebrate. Between 2 and 15 young are born at a time; there is no distinct breeding season. Frilled sharks are occasionally captured as bycatch by commercial fisheries but have little economic value. The International Union for Conservation of Nature (IUCN) has assessed it as Near Threatened, since given its very low reproductive rate even incidental catches may deplete its population. This shark, or a supposed giant relative, has been suggested as a source for reports of sea serpents.

Distribution and habitatEdit

Rather uncommon, the frilled shark has been recorded from a number of widely scattered locations in the Atlantic and Pacific Oceans. In the eastern Atlantic, it occurs off northern Norway, northern Scotland and western Ireland, from France to Morocco including Madeira, and off Mauritania. In the central Atlantic, it has been caught at several locations along the Mid-Atlantic Ridge, from north of the Azores to the Rio Grande Rise off southern Brazil, as well as over the Vavilov Ridge off West Africa. In the western Atlantic, it has been reported from off New England, Georgia, and Suriname. In the western Pacific, it is known from southeastern Honshu, Japan, to Taiwan, off New South Wales and Tasmania in Australia, and around New Zealand. In the central and eastern Pacific, it has been found off Hawaii, California, and northern Chile.[1]The frilled sharks off southern Africa were described as a different species, C. africana, in 2009.

The frilled shark inhabits the outer continental shelf and upper to middle continental slope, seeming to favor upwellings and other biologically productive areas. Though it has been caught from a depth of 1,570 m (5,150 ft), it usually does not occur deeper than 1,000 m (3,300 ft). In Suruga Bay, it is most common at a depth of 50–200 m (160–660 ft), except from August to November when the temperature at the 100 m (330 ft) water layer exceeds 15 °C (59 °F) and the sharks shift into deeper water. On rare occasions, this species has been seen at the surface. The frilled shark is usually found close to the bottom, with one individual observed swimming over an area of small sand dunes. However, its diet suggests that it does make substantial forays upward into open water. This species may make vertical migrations, approaching the surface at night to feed. There is spatial segregation by size and reproductive condition.

DescriptionEdit

With its elongated, eel-like body and strange appearance, the frilled shark has long been likened to the mythical sea serpent. The head is broad and flattened with a short, rounded snout. The nostrils are vertical slits, separated into incurrent and excurrent openings by a leading flap of skin. The moderately large eyes are horizontally oval and lack nictitating membranes (protective third eyelids). The very long jaws are positioned terminally (at the end of the snout), as opposed to the underslung jaws of most sharks. The corners of the mouth are devoid of furrows or folds. The tooth rows are rather widely spaced, numbering 19–28 in the upper jaw and 21–29 in the lower jaw. The teeth number around 300 in all; each tooth is small, with three slender, needle-like cusps alternating with two cusplets. There are six pairs of long gill slits with a "frilly" appearance created by the extended tips of the gill filaments, giving this shark its name. The first pair of gill slits meet across the throat, forming a "collar".

The pectoral fins are short and rounded. The single, small dorsal fin is positioned far back on the body, about opposite the anal fin, and has a rounded margin. The pelvic and anal fins are large, broad, and rounded, and also positioned well back on the body. The caudal fin is very long and roughly triangular, without a lower lobe or a ventral notch on the upper lobe. There are a pair of thick skin folds of unknown function running along the belly, separated by a groove. The midsection is relatively longer in females than in males, with the pelvic fins pushed closer to the anal fin. The dermal denticles are small and shaped like the tip of a chisel, becoming enlarged and sharp on the dorsal margin of the caudal fin. This species is a uniform dark brown or gray. The frilled shark differs from its southern African relative, C. africana, in having more vertebrae (160–171 vs 147) and more turns in the spiral valve intestine (35–49 versus 26–28), as well as in various proportional measurements such as a longer head and shorter gill slits. The maximum known length is 1.7 m (5.6 ft) for males and 2.0 m (6.6 ft) for females.

Biology and ecologyEdit

Highly specialized for life in the deep sea, the frilled shark has a reduced, poorly calcified skeleton and an enormous liver filled with low-density lipids, allowing it to maintain its position in the water column with little effort. It is one of the few sharks with an "open" lateral line, in which the mechanoreceptive hair cells are positioned in grooves that are directly exposed to the surrounding seawater. This configuration is thought to be basal in sharks and may enhance its sensitivity to the minute movements of its prey. Many frilled sharks are found with the tips of their tails missing, probably from predatory attacks by other shark species. Parasites identified from this shark include a tapeworm in the genus Monorygma, the fluke Distomum veliporum, and the nematode Metaleptus rabuka.

FeedingEdit

The long jaws of the frilled shark are highly distensible with an extremely wide gape, allowing it to swallow whole prey over one-half its size. However, the length and articulation of its jaws means it cannot deliver as strong a bite as more conventionally built sharks. Most captured individuals are found with no or barely identifiable stomach contents, suggesting a fast digestion rate and/or long intervals between feedings. This species preys upon cephalopods, bony fishes, and smaller sharks. One 1.6 m (5.2 ft) long individual, caught off Chōshi, was found to have swallowed a 590 g (1.3 lb) Japanese catshark (Apristurus japonicus). Squid comprise some 60% of the diet of sharks in Suruga Bay; this includes not only slow-moving, deep-dwelling types such as Chiroteuthis and Histioteuthis, but also relatively large, powerful swimmers of the open ocean such as Onychoteuthis, Sthenoteuthis, and Todarodes.

How the ostensibly weak-swimming frilled shark captures active, fast-moving squid is a matter of speculation. One possibility is that it takes advantage of injured squid, or those that are exhausted and dying after spawning. Alternatively, it may surprise its prey by curving its body and, bracing itself with its posteriorly positioned fins, launching a quick strike forward in the manner of a snake. It may also be able to close its gill slits and create negative pressure to suck prey into its mouth. The many small, sharp, recurved teeth of the frilled shark are functionally similar to squid jigs and could easily snag the body or tentacles of a squid, particularly as they are rotated outwards when the jaws are protruded. Observations of captive frilled sharks swimming with their mouths open suggest that the small teeth, light against the dark mouth, may even fool squid into attacking and entangling themselves.

Life historyEdit

The frilled shark is aplacental viviparous; the developing embryos are mainly nourished by yolk, though the difference in weight between the egg and the newborn indicates that the mother also provides additional nutrition via unknown means. Adult females have two functional ovaries and one functional uterus, on the right. Unsurprisingly, there is no defined breeding season for either sex, as this shark inhabits depths at which there is little to no seasonal influence. A possible mating aggregation of 15 male and 19 female sharks has been recorded over a seamount on the Mid-Atlantic Ridge. The litter size ranges from 2 to 15, with an average of 6. Females ovulate eggs into the uterus about once every two weeks; vitellogenesis (yolk formation) and the growth of new ovarian eggs halt during pregnancy, apparently due to insufficient space inside the body cavity.

Newly ovulated eggs and early-stage embryos are enclosed in a thin, ellipsoid, golden-brown capsule. When the embryo is 3 cm (1.2 in) long, the head is pointed when seen from above or below, the jaws are barely developed, the external gills have begun to appear, and all the fins are present. The egg capsule is shed when the embryo grows to 6–8 cm (2.4–3.1 in) long, and is expelled from the female's body; at this time the embryo's external gills are fully developed. The size of the yolk sac remains mostly constant until around an embryonic length of 40 cm (16 in), whereupon it begins to shrink, mostly or completely disappearing by an embryonic length of 50 cm (20 in). The embryonic growth rate averages 1.4 cm (0.55 in) per month, and therefore the entire gestation period may last three and a half years, far longer than any other vertebrate. Newborn sharks measure 40–60 cm (16–24 in) long; males attain sexual maturity at 1.0–1.2 m (3.3–3.9 ft) long, and females at 1.3–1.5 m (4.3–4.9 ft) long.

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