DOGFISH SHARKS Order Squaliformes Compagno, 1973C, J

Home , Bramble shark, Cirrhigaleus, Echinorhinus, Prickly shark, Squaliformes

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Order Squaliformes Compagno, 1973c, J.Linn.Soc.(Zool.), Land., 53, suppl. 1.

Synonymy : Order Cyclospondyli: Jordan & Evermann, 1896, Fowler, 1941, Smith, 1949. Order Euselachii, Suborder Squaloidea: Blot, 1969 (in part). Suborder Galei: Gill, 1872 (in part). Order Lamniformes, Suborder Squaloidei: Patterson, 1967 (in part). Suborder Plagiostomi Cyclospondyli: Hasse, 1879. Order Squalea, Suborder Squalida: White, 1936, 1937 (in part). Suborder Squali: Gill, 1862 (in part). Order Squaliformes: Rass & Lindberg, 1971, Applegate, 1974, Chu & Wen, 1979. Order Squaliformes, Suborder Squaloidei: Berg, 1940 (in part), Berg & Svedovidov, 1955 (in part), Arambourg & Bertin (1958) (in part). Suborder Squaliformes: Goodrich, 1909 (in part), Lozano y Rey, 1928, Bertin, 1939 (in part), Budker & Whitehead, 1971. Suborder Squalina: Matsubara, 1955, Fowler, 1968a. Order and Suborder Squaloidea: Schultz & Stern, 1948. Suborder Squaloidea: Romer, 1945, 1966 (in part), Bigelow & Schroeder, 1948, Norman, 1966. "Division" Squaloidei: Regan, 1906 (in part). "Group" Squaloidei: Garman, 1913 (in part). Suborder Squaloidei: Engelhardt, 1913 (in part), Lindberg, 1971, Nelson, 1976. Order Squatiniformes, Suborders Echinorhinoidei and Squaloidei: Glikman, 1967. Order Tectospondyli: Gill, 1893, Whitley, 1940. Order Tectospondyli, Suborder Squaloidei: Jordan 1923.

Field Marks : Cylindrical or compressed body, not raylike, 5 gill slits, 2 dorsal fins, no anal fin, snout normal, not sawlike.

Diagnostic Features : Trunk cylindrical, slightly depressed or somewhat compressed, not flattened and raylike. Head conical to depressed, not expanded laterally; five pairs of gill slits present on sides of head, with the posteriormost in front of pectoral fin origins; spiracles present and small to very large, well behind or close behind eyes and often above level of eyes; nostrils without nasoral grooves or circumnarial grooves, separate from mouth, anterior nasal flaps usually short and not reaching mouth, expanded into barbels in one genus (Cirrhigaleus) but without separate barbels on their outer edges; eyes on sides of head, without nictitating lower eyelids; snout short to long, flattened or conical, not greatly elongated into a flattened blade with lateral teeth and rostral barbels; mouth moderately large to short, arched and elongated to nearly transverse, below eyes; labial furrows well-developed, short to very long, on both jaws; teeth only moderately differentiated along the jaws, without enlarged anterior or posterior teeth and without a gap or small intermediate teeth between anterior and lateral teeth in the upper jaw. Two dorsal fins, with or without fin spines (Squaliolus with a spine on the first dorsal only), the first with origin varying from over the pectoral bases or gill slits to over the anterior halves of the pelvic bases; pectoral fins small to moderately large, not expanded and raylike, without triangular anterior lobes that cover the gill slits; pelvic fins small to moderately large, with vent continuous with their inner margins; anal fin absent; caudal fin with a long to moderate-sized dorsal lobe and the ventral lobe long to absent but always shorter than the dorsal lobe when present; vertebral axis elevated into the dorsal caudal lobe. Intestinal valve of spiral type.

Key to Families

1a. First dorsal fin originating behind pelvic fin origins (Fig. 1) - Bramble Echinorhinus Fig.1 sharks ...... Family Echinorhinidae (one genus, Echinorhinus)

1b. First dorsal fin originating in front of pelvic fin origins (Fig. 2)

2a. Body very high and compressed, triangular in cross- section; dorsal fins extremely partial lateral view Fig.2 high (Fig. 3) - Rough sharks ...... Family Oxynotidae (one genus, Oxynotus)

2b. Body low and more cylindrical in cross-section; dorsal fin low (Fig. 4) - Dogfish sharks ...... Family Squalidae (see key to genera) Oxinotus Fig.3

Centroscymnus Fig.4 - 25 -

3.1 FAMILY ECHINORHINIDAE Gill, 1862 ECHIN

Echinorhinoidae Gill, 1862, Ann.Lyceum Nat.Hist.N.Y., 7:406.

Synonymy : None.

FAO Names : En - Bramble sharks; Fr - Squales bouclés; Sp - Tiburones espinosos.

Field Marks : Short-nosed, cylindrical sharks with no anal fin, 2 very small, spineless, posterior dorsal fins, the first behind the pelvic origins, and coarse denticles or enlarged thorns.

Diagnostic Features: Trunk stout and cylindrical, without abdominal ridges. Head moderately depressed; last (5th) gill slits abruptly expanded in width; spiracles very small, well behind eyes; nostrils far apart from each other; mouth broadly arched, with very short labial furrows that do not encircle mouth; lips not papillose; teeth alike in both haws, strongly compressed and bladelike, with a cusp and up to three side cusplets in adults, but with a cusp only in young. Two small spineless dorsal fins, both smaller than the pelvic fins, the first with its base over the pelvic bases; caudal fin without a subterminal notch.

Habitat, Distribution and Biology: This family includes two uncommon species of large, poorly known wide-ranging, deepwater sharks in cold-temperate to tropical seas. They have a spotty but virtually circumglobal distribution on continental and insular shelves and slopes from 11 to 900 m depth, on or near the bottom. Bramble sharks reach a maximum size of 3 to 4 m. They feed on a variety of benthic and neritic fishes, including other sharks, ling, hake, flatfishes, lingcod, lizardfishes, rockfishes, topsmelt, herring, and elephantfishes (Callorinchus), as well as crabs, octopuses and squids. They have a moderate-sized mouth and a very large pharynx, and are thought to suck in their prey by suddenly expanding their mouths and pharynxes when in range. They are very sluggish. harmless sharks, never recorded as attacking people. Bramble sharks are ovoviviparous and lack a yolk- sac placenta.

Interest to Fisheries : Although these sharks attain a large size (3 to 4 m maximum total length), they are uncommon to rare in most areas where they occur and hence are of minimal interest to fisheries. They generally occur as a bycatch of other fisheries, including those for other sharks, and are taken on line gear, deepset gillnets, and more commonly in bottom trawls. They are used for fishmeal and medicinal purposes. Remarks : This family is sometimes placed as a synonym of Squalidae, but morphological study indicates that it is very distinct from the Squalidae. It has a single living genus, Echinorhinus.

Echinorhinus Blainville, 1816 ECHIN Echln

Subgenus Echinorhinus Blainville, 1816 (genus Squalus), Bull. Soc. Philomat.Paris, 8:121.

Type Species : "spinosus" (Squalus spinosus Gmelin, 1789), by monotypy, equals Squalus brucus Bonnaterre, 1788. Synonymy : Genus Goniodus Agassiz, 1838; Subgenus Rubusqualus Whitley, 1931 (Genus Echinorhinus); Genus Echynorhynus Nobre, 1935 error or emendation).

Remarks : I follow Garrick (1960) in recognizing two species in this genus.

Key to Species :

1a. Denticles of body numerous, close-packed, relatively small, stellate, and not fused into plates with multiple cusps ...... E cookei

1b. Denticles of body few, sparse, relatively large, not stellate, and with some fused into plates with multiple cusps ...... E brucus

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Echinorhinus brucus (Bonnaterre, 1788) ECHIN Echin 1

Squalus brucus Bonnaterre, 1788, Tabl.encyclop.method.trois reg.Nat., Ichthyol., Paris, 11. Holotype: lost. Type Locality: "L'Océan" (eastern North Atlantic).

Synonymy : Squalus spinosus Gmelin, 1789; Echinorhinus obesus Smith, 1849; Echinorhinus (Rubusqualus) mccoyi Whitley, 1931.

FAO Names : En - Bramble shark; Fr - Squale bouclé; Sp - Tiburón de clavos.

Field Marks : No anal fin, dorsals spineless and far back, first behind pelvic origins, large scattered thornlike denticles on body and fins.

Diagnostic Features : Dermal denticles on body and fins varying from small to very large, with many large, widely spaced, thorn or buckler-like denticles with bases not stellate and over a centimetre wide; some of these large denticles are fused in groups of 2 dermal denticles underside of head to 10 and may form large plates over 25 mm across.

Geographical Distribution : Western Atlantic: Virginia, Massachusetts, USA; Argentina. Eastern Atlantic: Scottish and Irish Atlantic Slopes and North Sea to Medi- terranean, Morocco, Canary Islands, Senegal, Ivory Coast; Namibia to Cape of Good Hope, South Africa. Western Indian Ocean: South Africa, southern Mozambique, ?Oman, India. Western Pacific: Japan (southeastern Honshu), Australia (South Australia), New Zealand, ?Kiribati.

Habitat and Biology : A large, sluggish bottom shark sometimes occurring in shallow water but primarily a deepwater species, occurring on the continental and insular shelves and upper slopes at depths from 18 to 900 m. Ovoviviparous, number of young per litter from 15 to 24; may breed in April in Indian waters. Eats smaller sharks (spiny dogfish), bony fishes (including ling, catfish, and lizardfish), and crabs. Size : Maximum total length about 3.1 m. Young born between 29 and 90 cm, adult females reported at 213 to 230 cm, adult males reported at 150 to 174 cm.

Interest to Fisheries: Apparently relatively common only in the eastern Atlantic, especially from the North Sea to Portugal, where it is caught in bottom trawls and on line gear. Utilized in the eastern Atlantic for fishmeal; liver oil has high value for medicinal purposes in South Africa. Relatively unimportant as a fisheries species.

Literature : Bigelow & Schroeder (1948); Tortonese (1958); Musick & McEachran (1969); Silas & Severaj (1972), Hureau & Monod (1973); Cadenat & Blache (l981). - 27 -

Echinorhinus cookei Pietschmann, 1928 ECHIN Echin 2

Echinorhinus cookei Pietschmann, 1928, Anz.Akad.Wien, 65:297. Holotype: Formerly in Bernice P. Bishop Museum, Oahu, Hawaiian Islands, apparently lost. Type Locality: South coast of Kauai, Hawaiian Islands. Neotype: National Museum of New Zealand, NMNZ 2774, Pallaser Bay, Cook Strait, New Zealand, designated by Garrick (1960:110).

Synonymy : None.

FAO Names : En - Prickly shark; Fr - Squale boucIé du Pacifique; Sp - Tiburón negro espinoso.

Field Marks: No anal fin, dorsals spineless and far back, first behind pelvic origins; denticles relatively close-set and only moderately large, not formed as large, flat thorns. Diagnostic Features : Dermal denticles on body moderate-sized, close-set, and with relatively small, stellate bases, less than 5 mm in diameter; denticles not expanded into large bucklers or thorns, not fused together. underside of head dermal denticles

Geographical Distribution : Western Pacific: Taiwan (Province of China), New Zealand. Central Pacific: Palau (Belau) and Hawaiian Islands. Eastern Pacific: Central California to Gulf of California, also Peru.

Habitat and Biology : A large, sluggish bottom shark, occurring on continental and insular shelves and upper slopes at depths from 11 to at least 424 m; in California caught in submarine canyons. Eats a variety of fishes, including spiny dogfish, the young of the sixgill shark (Hexanchus griseus), hake, flounders, rockfish, lingcod, topsmelt, herring, elephantfish (Callorinchus), and catshark (Apristurus) eggcases, also octopuses and squids.

Size : Maximum about 4 m, size at birth below 45 cm, mature males reaching at least 220 cm, females immature at 254 cm and adult at 299 cm (sex not reported for 4 m specimen).

Interest to Fisheries : Unimportant, occasionally taken by line gear, gillnets and bottom trawls.

Literature : Garrick (1960); Garrick & Moreland (1968); Varoujean (1972); B. Saunders (pers. comm.).

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