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Home , Black dogfish, Bramble shark, Centrophoridae, Chain catshark, Cirrhigaleus, Cookiecutter shark

Deepwater and Other of the U.S. Atlantic Exclusive Economic Zone

SARI J. KIRALY, JON A. MOORE, and PAULA H. JASINSKI

Introduction Conservation and Management on what is known regarding their life Act. Thirty-three of these were history characteristics, deep- sharks This paper reviews species descrip- listed by NOAA's National Marine are likely to be highly vulnerable to tions, including primary distribution Service (NMFS) as part of the over exploitation, especially given the (where known), of 51 deepwater and management unit in the Final Fishery rapid boom and bust tendencies of many other sharks ( 1) of the U.S. At- Management Plan for Atlantic , deep-sea fisheries (Moore, 1999). Also, lantic Ocean Exclusive Economic Zone , and Sharks (HMS FMP) over exploitation of shallow- fishery (EEZ) which currently are not included (USDOC, 1999) only in so far as to resources has driven some fishermen in any Federal fishery management plan prohibit finning, as these sharks are into deeper fishing grounds, and this can prepared under the Magnuson-Stevens sonfetimes taken incidentally in directed result in an increase in rates of , , and swordfish fisheries. certain deep-sea sharks in U.S. . Subsequently, in Amendment 1 to the For example, the monkfish, HMS FMP (USDOC, 2003) these spe- americanus, fishery off the northeastern Sari J. Kiraly is with the Highly Migratory Spe- cies Management Division, Office of Sustainable cies were removed from the management has ventured to depths of Fisheries, National Marine Fisheries Service, unit because a national ban on finning 700 m or more and can frequently capture NOAA, U.S. Department of Commerce, 1315 was instituted under the , Centroscyllilum fabricii, East-West Highway, Silver Spring, MD 20910 1 (e-mail: [email protected]), Jon A. Moore Prohibition Act. However, NMFS con- and various lantemsharks, Etinopterus is with the Atlantic University, Wilkes tinues to collect available fishing data on spp., as bycatch (co-author J. A. Moore, Honors College. 5353 Parkside Drive, Jupiter, FL 33458, and Paula H. Jasinski is with the Chesa- these species. personal observ.). Ecosystem manage- peake Office, Office of Conserva- For many of the sharks described ment of such deepwater fishing grounds tion. National Marine Fisheries Service, NOAA, here, little is known of either their biol- will require better knowledge of these P.O. Box 1346, Gloucester Point, VA 23062. ogy or distribution. Also, at present little deep-sea sharks. Obviously, for those information is known on the stock size or species where little to no information is reproductive characteristics of many of available on and stock size, ABSTRACT-Fifty-one deepwater and these species. For species where at least ecosystem management becomes an even other shark species of the U.S. Exclusive some reproductive information is known, more difficult task. Economic Zone in the and life-history characteristics include slower The other, non deepwater sharks of , which currently are not growth rates and lower reproductive rates, covered in this paper are included be- includedin any Federalfisherymanagement plan, are described, with a focus on pri- even when compared to shallow-water cause they play an important role in mary distribution. Many of these shark spe- sharks. When combined with the more their respective ecosystems as middle-to cies are not well known, while others which limited of cold, deep-sea top-level predators. Their inclusion un- are more common may be of particular environments, many of these deepwater derscores the importance of considering interest. Owing to concerns regardingpos- them in addressing sible increases infishing effort for some of sharks are at risk of serious decline before these species, as well as possible increases any fishery management scheme can be management. in bycatch rates as other fisheries move put into place. Since publication of the HMS FMP farther offshore, it is important that these The potential for an increased interest (USDOC, 1999) some of the species sharks be considered in marine ecosystem in fishing for these species warrants a listed in that document have been sys- management efforts. This will necessitate a better understanding of their biology and better understanding of them, and it is tematically reclassified based on new distribution. Primary distribution maps are important that they be included in any information. In some cases, there has included, based on geographic information ecosystem-based analysis pertaining to been confusion when referring to a spe- system (GIS) analyses of both published marine fisheries management. Based cies by common name. For example, and unpublished data, and a review of the more than one common name may be literature. The most recent systematic clas- sification and nomenclature for these spe- IH.R. 5461. P.L. 106-557, Shark Finning Prohi- used for the same species, or two sepa- cies is used. bition Act, 21 Dec. 2000, 114 Stat. 2772,4 p. rate species may share a nearly identical

65(4) I common name. No single standardized areas, and presence/absence data, and primary distribution maps: published reference for common names yet exists, represent the best available information literature, unpublished data and pho- especially for those species found below for these species. To visually illustrate tographs from the NMFS Northeast the 200 m depth zone (which serves as species primary distribution, the data Center's groundfish the cut-off point for one published list of were analyzed using a geographic infor- surveys, unpublished data from the ) (Robins et al., 1991). In those in- mation system (GIS), and shaded areas Northeast Fisheries Science Center's stances, the most frequently used and/or marking those boundaries are depicted fishery observer program, and catalogued the United Nations Food and Agriculture on the maps (Table 2). Biological and museum specimens from the following Organization (FAO) names are used in fishing-related information is taken in institutions: Academy of Natural Science this report. part from Castro (1983) and Compagno in Philadelphia (ANSP), CaliforniaAcad- (1 984), and updated with supplemental emy of Sciences (CAS), Field Museum of Species Accounts information from more recent literature, Natural History (FMNH), Florida State Primary distribution descriptions for personal observations, and personal Biological Collections (FSBC), Museum the shark species included here are based communications. of Comparative at Harvard on life history information, expert opin- The following data sources were used University (MCZ), Scripps Institution of ion regarding the importance of certain to construct the GIS and associated in La Jolla, Calif. (SIO),

Table 1.-Fifty-one deepwater and other sharks of the U.S. Atlantic Ocean Exclusive Economic Zone which currently are not included in any Federal fishery management plan prepared under the Magnuson-Stevens Fishery Conservation and Management Act. Nomenclature note: Apristuruslaurussoniis the incorrect name for the species commonly found off the U.S. What is in U.S. waters Is an undescribed species similar in appearance to A. laurussoni(Nakaya').The proper designation is Apristurussp.As of yet, there is no common name to apply to this species.

ECHiNORHINIDAE (Bramble Sharks) laticaudus Smith and Radcliffe, 1912 Apnisturusprofundorum (Goode and Bean, 1896) brucus (Bonnaterre, 1788) Spined , Pygmy shark Deepwater (Gulper Sharks) Apr/sturus riven' Bigelow and Schroeder, 1944 (Lanternsharks) cf. acus Garman, 1906 Broadgill catshark fabricii (Reinhardt, 1825) Undescribed #1 sp. (laurussonimlike) Black dogfish An undescribed species similar to the Needle dogfish Undescribed carshark #1 bigelowi Shirai andTachikawa, 1993 C. acus from the Pacific (McEachman and Fechhelm, An undescribed species similar in appearance to Blurred lanternshark 1998). A. Iaurussoni (Nakayal). Often confused with the similar E pusillus, which does Centrophorus granulosus (Bloch and Schneider, 1801) antillensis Springer, 1979 not occur in the western Atlantic Gulper shark Antillean roughtail catshark Etmopterus bullisi Bigelow and Schroeder, 1957 Centrophorus cf. harrissoni McCulloch, 1915 Ga/eus aae (Nichols, 1927) Lined lanternshark Undescribed gulper shark #2 Marbled catshark, Roughtall catshark Etmopterus gracilispinis Krefft, 1968 Usually identified as C. uyato, but that species name 2 Galeus springeri Konstantinou and Cozzt, 1998 Broadbanded lanternshark, Broadband dogfish is now considered invalid (Compagno ). Striped sawtail catshark Etmopterus hillanus (Poey, 1861) Centrophorus niaukangTeng, 1959 Scyl/orhinus boa Goode and Bean, 1896 lantemshark, Blackbelly dogfish gulper shark Boa catshark Etmopterus princeps Collett, 1904 Centrophorus cf. tessellatus Garman, 1905 Scyl/orhinus meadi Springer, 1966 , Rough sagre Undescribed gulper shark #3 Blotched catshark Etmopterus robinsi Schofield and Burgess, 1997 An undescribed species similar to the Tesellated Scyliorlinus retifer (Garman, 1881) West Indian lantemshark gulper shark C. tessellatus from the Pacific according , Chain dogfish Etmopterus schultzi Bigelow, Schroeder, and to McEachran and Fechhelm (1998). torrei/Howell-Rivero, 1935 Springer, 1953 profundorm (Smith and Radcliffe, 1912) Dwarf catshark Fringefin lanternshark Arrowhead gulper shark, Arrowhead dogfish, Flatnose PROSCYLLIDAE (Ribbontail ) dogfish Etmopterus Wirens (Bigelow, Schroeder, and Er/dacnis barbouri (Bigelow and Schroeder, 1944) Springer, 1953) ( Sharks) Cuban ribbontail catshark Green lantemshark asper (Merrett, 1973) barbouti is a frequently used . (Sleeper Sharks) Roughskin spiny dogfish, Roughskin (False Catsharks) coelolepis Bocage and Capello, 1864 This species was recently transferred from Squalus Pseudotriakis microdon Capello, 1867 Portuguese shark, to this . Centroscymnus owstoni Garman, 1906 Squalus cubensis Howell-Rivero, 1938 Scyl/orhinus licha is an older name for this species. CHLAMYDOSELACHIDAE (Frill Sharks) C. ctyptacanthus is a junior synonym of this species. Squalus mitsukurf/Jordan and Snyder, 1903 Chlamydoselachus anguineus Garman, 1884 Somnlosusmicrocephalus (Bloch and Schneider, 1801) Shortspine dogfish, Frill shark shark Often misidentified as S. blainvillei, which is now TRIAKIDAE (Smoothhound Sharks) Symnodon squamulosus (Gianther, 1877) considered an invalid name (Compagno2). Mustelus canis canis (Mitchill, 1815) PRSTIOPHORIDAE () Dusky smoothhound, Smooth dogfish S. obscurus is a junior synonym of this species Puistiophorus schroederi Springer and Bullis, 1960 Mustelus canis insularis Heemstra, 1997 according toYano and Tanaka (1984) and Wetherbee American , Bahamas sawshark Antillean smoothhound and Crow (1996). SCYLIORHINIDAE (Catsharks) MustelushigmaniSpringer and Lowe, 1963 (Kitefin Sharks) Apristurus canutus Springer and Heemstra, 1979 Smalleye smoothhound Dalatias lIcha (Bonnaterre, 1788) Hoary catshark Mustelus norr/s/Springer, 1940 Apristurus manis (Springer, 1979) Florida smoothhound, Narrowfin smoothhound brasiliensis (Quoy and Gaimard, 1824) Mustelus sinusmexicanus Heemstra, 1997 Apristurus microps (Gilchrist, 1922) Gulf smoothhound Isistiusplutodus Garrick and Springer, 1964 Smalleye catshark ODONTASPIDAE ( Tiger Sharks) Bigtooth cookiecutter shark, Largetooth cookie-cutter Apristurus parvipinnis Springer and Heemstra, 1979 ferox (Risso, 1810) shark Smallfin catshark Ragged-tooth shark, Smalltooth sand

'Nakaya, K. 1999. Personal commun., Hokkaido Univ., . 2Compango, L. 1999. Personal commun., South African Museum, Town.

2 Marine FisheriesReview Texas Cooperative Wildlife Collections at Texas A&M University (TCWC), Texas Natural History Collection (TNHC), Texas University Collections (TU), University of Alabama Ichthyologi- cal Collections (UAIC), University of -'~~~~~~~.... -_.S.gNt000000o000000''.' t000000... ,ISE.0...I000000 Florida in Gainesville (UF), University of North Carolina Institute of Marine Figure 1.-Black dogfish, Centroscyllium fabricii. taken at Science (UNCMIS), University of Puerto about 915 m depth near Hydrographer during deep- Rico at Managua (UPRM), National water monkfish survey aboard F/V Mary K. NMFS photo. Museum of Natural History (USNM) at the Smithsonian Institution, and Yale University's Peabody Museum of Natural PrimaryDistribution Within thze U.S. Table 2.-Primary distribution maps. History (YPM). EEZ (Map 1) In the Map no. Name Page no,

All length measurements of specimens in water depths of 300-900 m, from 1 Bramble Shark 21 are given as total length. the Florida Panhandle east of DeSoto 2 Black Dogfish 22 3 Blurred Lanternshark 23 Canyon at 86.5°W to west of Mississippi 4 Lined Lanternshark 24 Bramble Sharks (Echinorhinidae) Canyon at 90.5°W. 5 Broadband Dogfish 25 6 Caribbean Lanternshark 26 Bramble shark 7 Great Lanternshark 27 Lanternsharks (Etmopteridae) 8 West Indian Lanternshark 28 Echinorhinus brucus 9 Fringefin Lanternshark 29 Black dogfish 10 Green Lanternshark 30 The bramble shark is a large, sluggish, 11 Portuguese Shark 31 bottom dwelling shark. It is primarily a Centroscylliulmfabricii 12 Roughskin Dogfish 32 13 33 deepwater species, widely distributed in The black dogfish (Fig. 1) is a small, 14 Velvet Dogfish 34 the deep temperate and tropical waters bottom-dwelling shark that sometimes 15 Kitefin Shark 35 16 Cookie-cutter Shark 36 of continental and insular shelves and forms large schools, which are possi- 17 Undescribed Gulper Shark #1 37 upper slopes. Although the species ap- bly segregated by sex and size (ICES, 18 Gulper Shark 38 19 Undescribed Gulper Shark #2 39 to be most abundant at depths of 1995). It is a deepwater species found 20 Taiwan Gulper Shark 40 300-900 m, it is occasionally found in in temperate to boreal waters over the 21 Taiwan Gulper Shark 41 22 Undescribed Gulper Shark #3 42 shallower water (<18 m). Bramble shark outer continental shelves and slopes of 23 Arrowhead Gulper Shark 43 catches are often reported in the eastern the North Atlantic Ocean. Catches are 24 Cuban Dogfish 44 180-1,600 m, 25 Shortspine Dogfish 45 Atlantic and western Indian , but reported from depths of 26 American Sawshark 46 only five specimens have been reported most usually deeper than 250 m. This 27 Hoary Catshark 47 from the North American east in species typically occurs at shallower 28 Ghost Catshark 48 29 Smallfin Catshark 49 the past century (one off Cape , depths in the northern part of its range. 30 Broadgill Catshark 50 Mass., one off , and three near It is found off the United States from 31 Undescribed Catshark 51 32 Antillean Roughtail Catshark 52 the Mississippi Delta) (Goode and Georges Bank, off New England, to Cape 33 Marbled catshark 53 Bean, 1896; Musick and McEachran, Hatteras, N.C., and possibly also in the 34 Striped Sawtail Catshark 54 35 Boa Catshark 55 1969; Schwartz, 1993; McEachran and northem Gulf of Mexico. 36 Blotched Catshark 56 Fechhelm,1998). Biology This species feeds on small 37 Chain Dogfish 57 38 Dwarf Catshark 58 Biology Very little is known about , , bony fishes, 39 Cuban Ribbontail Catshark 59 the habits of the bramble shark. It report- and (Compagno, 2002a). De- 40 Smooth Dogfish 60 41 Antillean Smoothhound 61 edly feeds on smaller sharks, bony fishes, velopment is ovoviviparous and litters of 42 Florida Smoothhound 62 and . Development is ovovivipa- up to 50 pups have been reported (ICES, 43 Gulf Smoothhound 63 rous; i.e. hatch in the before 1995). Pups are approximately 14 cm (6 the embryos are fully developed, and the in) at birth. Females mature at 58-70 cm embryos then continue to be nourished by (23-28 in) although the species is known New England, south to Cape Hatteras, the sac. Litters of 15-24 pups have to reach a maximum size of 120 cm (48 N.C., at 350N. been reported, with pups approximately in) (ICES, 1995). 40 cm (16 in) at birth. Males mature at Fishing Black dogfish are taken as Blurred lanternshark 150-174 cm (60-70 in) and females at bycatch in deepwater trawl and longline Etmopterms bigelowi 213-230 cm (85-92 in). The species is fisheries. The blurred lantemshark is a recently known to reach a maximum size of 310 PrimaryDistribution Within the U.S. described small, deepwater shark dis- cm (124 in). EEZ (Map 2) In the Atlantic Ocean in tributed in tropical to temperate waters Fishing Thebramble sharkis usually water depths of 180-1,600 m, from the in the Atlantic, southwestern Indian, caught in water depths >300 m. northern extent of the EEZ boundary off and southeastern Pacific Oceans (Shirai

65(4) 3 __I ______I-_ Ii - - _ _ ' - __ _ ' -, . 9 I I I _ = * -::fI I Figure 2.-Broadband dogfish, Etmopterus gracilispinis,from 2002 Bear cruise DE02-06 at depth of 0-1194 m in open water. Photo: Chris Kenaly (Harvard Univ. specimen MCZ 161542).

and Tachikawa, 1993). It formerly was confused with the , Etmopterus pusillus, which does not occur in the western North Atlantic (Shirai and Tachikawa, 1993). The blurred lanternshark is both mesopelagic and benthopelagic, occurring over the outer continental shelves and slopes at depths ranging from 100 m to below 1,000 m. In U.S. waters it is distributed in the northeastern Gulf of Mexico from Florida to . Biology The blurred lanternshark is known to feed on and small fishes. Development is ovoviviparous. Females mature at 50 cm (20 in) and males at 42 cm (17 in). It is known to reach a maximum size of 66 cm (27 in) Figure 3.-Head of broadband dogfish, Etmopterus gracilispinis, from 2002 Bear Seamount cruise DE02-06 at depth of 1194 m in open water. Photo: Jon Moore (Shirai and Tachikawa, 1993). Blurred (Harvard Univ. specimen MCZ 161542). lanternsharks show a size vs. depth segregation, with smaller individuals occurring at shallower depths and larger ones in deeper waters. in the Atlantic from Florida to North Fishing The lined lantemshark is Fislhing The blurred lantemshark is Carolina. This species also most likely caught in deepwater trawls. caught in deepwater trawls. occurs off Puerto Rico and the Virgin Is- PrimaryDistribution Within the U.S. Primary DistributionWithini the U.S. lands, given its occurrence off the nearby EEZ (Map 4) In the Atlantic Ocean EEZ (Map 3) In the Gulf of Mexico Leeward . and Gulf of Mexico in water depths of in water depths of 100-1,000 m, from Biology The lined lanternshark 200-850 m: in the Atlantic extending the at 81.3°W, west to the feeds on small crustaceans and squid from Cape Hatteras, N.C., at 35°N, south Mississippi at 89.5°W. (McEachran and Fechhelm, 1998). It is around Florida, including the of capable of swallowing relatively large Florida, and into the Gulf ofMexico west Lined lantemshark squid whole, possibly by distending its to the Texas/Mexico border. Etinopterus bullisi jaws. Although development is presumed The lined lanternshark is a small. to be ovoviviparous, nothing else is Broadband dogfish (also referred to slender shark that inhabits the tropical to known of its reproduction. The majority as Broadbanded lantermshark) temperate western Atlantic Ocean. It is a of captured specimens have been juve- Etmopterus gracilispinis deepwater species, usually found along niles measuring 18-26 cm (7-10 in). The Broadband dogfish (Fig. 2, 3) have the continental slope in water depths of maximum size known is 26 cm (10 in) been reported from the western North 250-850 m. In U.S. waters it is distrib- (Compagno, 2002a). The size of an adult Atlantic Ocean and from both sides of the uted in the northem Gulf of Mexico and lined lantermshark is not yet known. South Atlantic (Compagno, 2002a). It is

4 Marine FisheriesReview widely distributed in the Gulf of Mexico from Florida to Louisiana, and along the east coast from Florida to the southern New England slope south of Nantucket, Mass. (Moore et al., 2003). This species is mesopelagic, occurring at depths of 70-480 m, and benthopelagic along the outer and upper slope, where it occurs at depths ranging from Figure 4.-Great lantermshark, Etmopterus princeps, collected by a comrnmercial fishernan during exploratory fishing at depth of 1464 m in Welker Canyon on 100 to 1,000 m. Similar to several other Georges Bank. Specimen is 58 cm long. Photo: Jon Moore. (Harvard Univ. specimen small, deepwater sharks, it is believed to YPM 10419). ascend the at night. Biology Development is ovovivipa- rous. A newborn specimen measuring 13 (10 in), although some individuals have PrimaryDistribution Within the U.S. cm (5 in) has been reported, while a 26 been reported at 28 cm (1 in). EEZ (Map 7) In the Atlantic Ocean at cm (10 in) male was determined to be im- Fishing The Caribbean lanternshark depths of 500-2,200 m, from the northern mature. Specimens of broadband dogfish is caught in deepwater trawls. extent of the EEZ boundary off New Eng- have been recorded to 33 cm (13 in). Primary Distribution Within the U.S. land south to Delaware at 38.7°N. Fishing The broadband dogfish is EEZ (Map 6) In the Caribbean Sea in caught in deepwater trawls. water depths of 300-700 m surrounding West Indian lanternshark PrimaryDistribution Within the U.S. Puerto Rico and the U.S. Virgin Islands, Etmtopterus robinsi EEZ (Map 5) In the Atlantic Ocean extending east and west to the limits of The West Indian lanternshark is a and Gulf of Mexico in water depths of the EEZ; in the Atlantic Ocean and Gulf small, recently described shark found in 70-1,000 m: in the Atlantic from the of Mexico in water depths of 300-700 m: the tropical western NorthAtlantic Ocean northern extent of the EEZ off New in the Atlantic from the Virginia/Mary- (Schofield and Burgess, 1997). It appears England, south to Cape Hatteras, N.C., land border at 38°N, extending south to be confined to deep waters of the upper at 35°N. Also rare individuals are found around Florida, including the Straits of continental and insular slopes, and has to occur from Cape Hatteras south to Florida, into the Gulf of Mexico west been reported from depths of 400-800 northern Florida to 29°N (Compagno, to 83.2°W; in the Gulf of Mexico from m, with most reports deeper than 550 m. 2002a; Moore et al., 2003); in the Gulf of the Florida Panhandle east of DeSoto In U.S. waters it is found off southern Mexico from the Florida Panhandle east Canyon at 86°W, west to the Mississippi Florida and likely occurs off Puerto Rico of DeSoto Canyon at 86.5°W west to the River Delta at 91 oW. and the Virgin Islands, given its occur- area of the Mississippi Canyon at 90.5°W. rence off the nearby Leeward Islands. Great lanternshark Biology Very little is known of the Caribbean lanternshark Etmopterus princeps biology of this species. Males reach ma- (also referred to as Blackbelly dogfish) The great lanternshark (Fig. 4) is a turity at about 26 cm (10 in) (Schofield Etmopterus hillianus small, slender shark (although the larg- and Burgess, 1997). Development is The Caribbean lanternshark is a small est lanternshark species in the western presumably ovoviviparous. The maxi- shark found in the tropical to subtropical Atlantic Ocean) found throughout the mum reported size is about 34 cm (14 in) Caribbean Sea and the western North temperate North Atlantic Ocean. Off (Schofield and Burgess, 1997). Atlantic Ocean. It appears to be confined the United States it is caught from off Fishing The West Indian lantern- to deep waters of the upper continental Georges Bank south to New Jersey shark is caught in deepwater trawls. and insular slopes, and has been reported (Moore et al., 2003). Confined to deep Primary Distribution Within the U.S. from depths of 300-700 m (Schofield waters of the continental slopes, the great EEZ (Map 8) In the Atlantic Ocean and Burgess, 1997). In U.S. waters it is lanternshark has been reported from and Gulf of Mexico in water depths of found from southern Florida to Virginia, depths of 500-2,200 m. 400-800 m: in the South Atlantic begin- in the northern Gulf of Mexico off the Biology The diet of this shark con- ning at 30°N out to the EEZ boundary, Delta, and off Puerto sists of and decapod crustaceans including the Blake , extending Rico and the Virgin Islands. (ICES, 1995). Development is ovovi- south around Florida, including the Biology Males and females reach viparous and pups are born at about 18 Straits of Florida, and into the Gulf of maturity at about 20 cm (8 in) and 30 cm (7 in) (ICES, 1995). A 55 cm (22 in) Mexico west to 83.8°W. cm (12 in), respectively (Schofield and male was reported as mature, although Burgess, 1997). Development is ovovi- individuals can grow to a length of about Fringefin lanternshark viparous. Litters consist of up to five 89 cm (36 in). Etmopterus schultzi pups that are approximately 9 cm (4 in) Fishing The great lanternshark is The fringefin lanternshark is a small, at birth. The average size is about 25 cm caught only in deepwater trawls. deepwater shark found in the tropical

65(4) 5 Figure 5.-Portuguese shark, Centroscymnits coelolepis, (top, specimen no. YMP 10137) and black dogfish, Centroscylliumfabri- cii, (bottom, specimen no. YPM 10134) caught by exploratory deepwater fisherman near Block Canyon at depth of 1006 m. Photo: Jon Moore.

and subtropical western North Atlantic the Caribbean Sea and Gulf of Mexico Puerto Rico and the U.S. Virgin Islands, Ocean. It appears to be common along to and off the east coast of extending east and west to the limits of upper continental slopes at depths rang- Florida. It appears to live in dense schools the EEZ; in the Atlantic Ocean and Gulf ing from 200 to 1,000 m. In U.S. waters confined to moderately deep waters, typi- of Mexico in water depths of 100-1000 it is distributed in the northern Gulf of cally 100-1,000 m. In U.S. waters it is m: in the Atlantic beginning at 30°N Mexico from Florida to Texas, and in the relatively common in the northern Gulf out to the EEZ boundary, including the Atlantic from Florida to South Carolina of Mexico from Texas to Florida, and off Blake Plateau, extending south around (although it may possibly extend north to Puerto Rico and the Virgin Islands. Florida, including the Straits of Florida, Cape Hatteras). Biology The green lanternshark and into the Gulf of Mexico west to the Biology The species is known to feed feeds primarily on squid. The cephalo- Texas/Mexico border. on squid. No data on its reproductive pod beaks and eyes commonly found development are available. Males mature in stomach contents are large enough Sleeper Sharks (Somniosidae) at 27 cm (11 in) and females at 28-30 cm to indicate that the shark's jaws must (11-12 in). The maximum reported size have been greatly stretched at the time Portuguese shark of a fringefin lantemshark is 30 cm (12 of swallowing. It has been suggested Centroscymnus coelolepis in) (Compagno, 2002a). that dense schools of these sharks attack The Portuguese shark (Fig. 5) inhabits Fishing The fringefin lantemshark is prey much larger than themselves, biting very deep waters along the continental caught in deepwater trawls. off chunks with their sharp lower teeth. slope and rise nearly worldwide. This PrimnaryDistribution Within the U.S. Development is ovoviviparous and shark is found at depths of 150-3,700 EEZ (Map 9) In the Atlantic Ocean the period is believed to last m, with most captures occurring in water and Gulf of Mexico in water depths of around one year. Litters consist of 1-3 depths >600 m (Compagno, 2002b) and 200-1,000 m: in the Atlantic extending pups, which measure nearly 9 cm (4 in) at temperatures of 5-6°C (41-430 F). from the North CarolinalSouth Carolina at birth. Maturity is reached at about Due to its preferred depth range, few border at 33°N, south around Florida, 20-23 cm (8-9 in) and the average size is catches have been reported in North including the Straits of Florida, and into 20-25 cm (8-1 0 in). The maximum size American waters. Sporadic captures off the Gulf of Mexico west to the Texas/ observed is 26 cm (10 in) (Compagno, the United States range from the Straits Mexico border. 2002a). of Florida to Georges Bank. Fishing The green lanternshark is Biology Stomach contents of Portu- Green lanternshark caught in deepwater trawls. guese sharks have consisted of small Etmopterus virens PrimaryDistribution Within the U.S. fishes, squid, and cetacean flesh. Devel- The green lanternshark is found in EEZ (Map 10) In the Caribbean Sea in opment is ovoviviparous, with average the western North Atlantic Ocean, from water depths of 100-1,000 m surrounding litters of 13-16 pups measuring 27-30

6 Marine FisheriesReview cm (11-12 in) at birth. Japanese stud- ate to polarNorth Atlantic waters. This is boundary off New England south to ies indicate that Portuguese sharks may the only shark species regularly encoun- Cape Cod, Mass., including the Gulf have a tri-annual reproductive cycle. The tered in the polar waters of the Atlantic of Maine and Cape Cod Bay. Rare indi- average size is 90-107 cm (35-42 in), Ocean. Individuals have been reported viduals have been found off New Jersey although the maximum reported size is in the Gulf of Maine and as far south as and Georgia. 1995). Georgia (Herdendorf and Berra, 1995). 128 cm (51 in) (ICES, Velvet dogfish of Portuguese This species has been found in water Fishing The majority squamulosus shark specimens are caught on longlines temperatures ranging from 0.6 to 16°C set at depths >350 m, but this species is (33-61°F), with most specimens taken The velvet dogfish is a little-known also occasionally taken in trawls. The in temperatures of 2-7°C (36-45°F). In deepwater shark found in tropical to species is commercially fished in the the summer, the Greenland shark tends to subtropical waters of the Atlantic, south- eastern Atlantic Ocean. reside at depths of 200-750 m, although west Indian, and western Pacific Oceans. Primary Distribution Within the U.S. some have occurred as deep as 2,200 m It is mesopelagic and benthopelagic EEZ (Map 11) In the Atlantic Ocean (Herdendorf and Berra, 1995). During over continental slopes at depths rang- and Gulf of Mexico in water depths of the winter months individuals ascend ing from 39 to 2,200 m, but usually at 160-3,700 m: in the Atlantic from the the water column, often approaching the depths between 400 m and 900 m (Yano northern extent of the EEZ boundary off surface at the edge of the ice. and Tanaka, 1984; Wetherbee and Crow, New England extending south around Biology The Greenland shark feeds 1996). In U.S. waters this species is found Florida, including the Straits of Florida, on bony fishes (e.g. , char, halibut, in the northern Gulf of Mexico in the west to the Dry Tortugas at 83.4°W. herring, lumpfish, and salmon), seals, vicinity of the , , squid, crabs, and other benthic and possibly off Puerto Rico and the Roughskin dogfish . Stomach content analyses Virgin Islands. Scymnodon obscurus is a Centroscymnus owstoni reveal fast-moving fishes often found frequently used synonym for this species The roughskin dogfish inhabits deep with their tails bitten off. This species (Yano and Tanaka, 1984; Wetherbee and waters along the continental and sea- often gathers in large numbers around Crow, 1996). mount slopes nearly worldwide at sealing or whaling operations, feeding Biology Velvet dogfish feed on fishes depths of 400-1,500 m. Rare captures on or carrion. Size at maturity is and bottom invertebrates. of devel- of this species in U.S. waters are from 240 cm (96 in) (Scott and Scott, 1988). opment is unknown. Males mature at 52 the northern Gulf of Mexico from the Development is ovoviviparous. Pups cm (21 in) and females at 75 cm (30 in) vicinity of the Mississippi River Delta measure about 38 cm (15 in) at birth, (Yano andTanaka, 1984). The maximum to Pensacola, Fla., and the Florida Keys. and up to 10 pups have been reported in reported size is 84 cm (34 in). Centroscymnnus cryptacanthusis a com- one litter. Tagging studies have shown Fishing Velvet dogfish are infrequent monly used junior synonym for this the Greenland shark to be a very slow- bycatch in longline and trawl fisheries. species (Compagno, 2002b). growing fish-medium-size specimens Primary Distribution Within the U.S. Biology The diet of roughskin dog- appear to grow only 1 cm (0.4 in) or EEZ (Map 14) In the Gulf of Mexico fish consists of small fishes. Very little is less per year. The average size is 340 cm in water depths of 400-900 m, from known about development in the species. (11.1 ft) and 285 kilograms (kg) (627 lb), the Florida Panhandle east of DeSoto Males mature at 72-84 cm (29-34 in) and whereas the largest specimen on record Canyon at 88.5°W, west to the Missis- females at 102 cm (41 in). The maximum measured 640 cm (21.0 ft) and weighed sippi River Delta at 86.4°W. reported size is 104 cm (42 in). 1,022 kg (2,250 lb). Fishing Roughskin dogfish are oc- Fishing The Greenland shark has Kitefin Sharks (Dalatiidae) in trawls or longlines been fished along the of , casionally caught Kitefin shark 400 m. , and Greenland for its oil. In set at depths greater than Dalatias licha Primary Distribution Within the U.S. Greenland it is targeted using longlines at EEZ (Map 12) In the Gulf of Mexico 250-550 m depths. In the winter fisher- The kitefin shark is a small, deepwater in water depths of 400-1,500 m, from men often use light to lure sharks to the shark found worldwide, usually over the the Straits of Florida at 80.3°W, west to surface where they tend to be extremely outer continental and insular shelf and at 82.4°W, and from the Florida sluggish and offer little resistance. slope at depths of 37-1,800 m, but mostly Panhandle east of DeSoto Canyon at Individuals are occasionally caught as at depths below 200 m. This species is 88.5°W, west to the Mississippi River bycatch in squid trawl fisheries off the rarely found in U.S. waters-catches Delta at 86.40W. northeastern United States. have been reported on Georges Bank, Primary Distribution Within the U.S. the Blake Plateau off Florida, the north- Greenland shark EEZ (Map 13) In the Atlantic Ocean ern Gulf of Mexico off the Mississippi microcephalus from inshore coastal waters and embay- River Delta, and off Puerto Rico and The Greenland shark is a large, wide- ments, except , out to 2,200 the Virgin Islands. Catch records in the bodied shark commonly found in temper- m, from the northern extent of the EEZ Mediterranean suggest that the kitefin is

65(4) 7 Figure 7.-Cookie-cutter shark, Isistius brasiliensis. Photo: Karsten Hartel (Har- vard Univ. specimen MCZ 60365).

Figure 6.-Cookie-cutter shark, Isi- 1988). Average size and weight for the Alabama) and the area north of the Ba- stius brasiliensis. Photo' Karsten Hartel (Harvard Univ. specimen species is about 120 cm (47 in) and 8 hamas. However, it is also likely found MCZ 60365). kg (18 lb), respectively, with the largest off Puerto Rico and the Virgin Islands. specimen on record measuring 182 cm Most catches occur after dark between (72 in) (ICES, 1995). the surface and 550 m, indicating a pos- primarily a solitary shark that does not Fishing The kitefin shark is usu- sible nighttime vertical migration from exhibit schooling behavior. ally taken in deepwater trawls or on deeper water (Compagno, 2002c). The Biology The kitefin shark is a ver- longlines. species is also thought to exhibit school- satile deep-sea predator that feeds on Primary Distribution Within the U.S. ing behavior. numerous bony fishes, rays, crabs, EEZ (Map 15) In the Atlantic Ocean Biology The cookie-cutter sharkhas and squid. Adults tend to consume and Gulf of Mexico in water depths of very powerful jaws and large teeth. It more crustaceans and sharks and fewer 200-1,800 m: in the Atlantic off Florida feeds extensively on large squid, but may cephalopods than do juveniles, and in at 30.4°N, east out to the EEZ boundary, also attack even larger prey. Evidence the spring and winter rely heavily on including the Blake Plateau, and south indicates that it feeds by taking bites sharks as an alternative food source. to the EEZ boundary; in the Gulf of from large pelagic fishes (tunas, wahoo, Submersible observations indicate that Mexico from the Florida Panhandle east dolphin, marlins, etc.) as well as por- this species is also likely to be a of DeSoto Canyon at 85.5°W, west to the poises and . It has been suggested (Clark and Kristof. 1990). Development Mississippi River Delta at 89°W; in the that the shark is able to grab a quick bite is ovoviviparous, with litters consisting Caribbean Sea at depths of 200-1,800 after being approached, and subsequently of 10-16 pups. The gestation period is m surrounding Puerto Rico and the U.S. rejected, by larger in search of estimated to be two years (Silva, 1988), Virgin Islands, extending east and west prey. The ventral surfaces of the head and with pups measuring 42-45 cm (17-18 to the limits of the EEZ. body (except for the dark collar around in) at birth (Silva, 1988). In the , the gill area), as well as the ventral fin von Bertalanffy growth parameters were Cookie-cutter shark surfaces, are luminescent and emit a calculated as L. = 127, k =.198, and Isistius brasiliensis bright greenish glow. The number of light to = -1.949; Holden's method gave L, = The cookie-cutter shark (Fig. 6, 7) organs is highly variable-some speci- 130, k =.195, and to = -2.000 for males inhabits deep waters of the tropical and mens may have very few or emit no light and L. = 165, k =.147, and to = -2.000 subtropical belts of the Atlantic, Pacific, at all. Cookie-cutter shark development for females (Silva, 1988). Males reach and Indian Oceans. It is a very small is presumed to be ovoviviparous. Six maturity at 77-121 cm (31-48 in) and shark species, usually characterized as or seven large eggs have been reported females at 120 cm (47 in). Studies in the epipelagic to bathypelagic (caught any- from females, but embryos have not yet Azores indicate that there may be depth where from the surface down to 3,700 been reported. Females mature at 38-44 segregation by sex, with females peak- m) (Retzer, 1990; Compagno, 2002c). cm (15-18 in) and males are thought to ing in abundance at a depth of 230 m The reported captures come from the reach maturity around 31-37 cm (12-15 and males peaking at 412-448 m (Silva, northern Gulf of Mexico (from Texas to in). The cookie-cutter sharkranges in size

8 MarineFishzeries Review from 14 to 50 cm (6-20 in); the largest ranging worldwide, inhabiting temperate in the Smithsonian are from depths of on record is 50 cm (20 in). and tropical mesopelagic waters at depths 732-915 m. Fishing The cookie-cutter shark is ranging from 200 to 800 m (Glukhov and Biology No biological information is caught at the surface and in mid-water Kuzmichev, 1984). Pygmy sharks tend available until this species is distinguished trawls after dark. It does not appear to to undergo diurnal vertical migrations, from the Japanese gulper shark. be attracted to lights. migrating upward to depths of 200 m at Fishery This species is taken with Primary Distribution Within the U.S. night to feed. This species is known to either trawls or longlines. EEZ (Map 16) In the Atlantic Ocean occur in U.S. waters off Virginia, and in Primary Distribution Within the U.S. and Gulf of Mexico in water depths of the northern Gulf of Mexico (McEachran EEZ (Map 17) In the Caribbean Sea in 200-3,700 m or out to the EEZ boundary, and Fechhelm, 1998; Moore et al., water depths of 200-900 m surrounding whichever is closer, extending from Cape 2003). Puerto Rico and the U.S. Virgin Islands, Canaveral, Fla., at 280N, south around Biology The is extending east and west to the limits of Florida, including the Straits of Florida, known to feed on squid, , the EEZ; in the Atlantic Ocean and Gulf and into the Gulf of Mexico west to the and lightfishes. Although embryos have of Mexico in water depths of 200-900 Texas/Mexico border. not been observed, development is pre- m: in the Atlantic off Virginia and North sumed to be ovoviviparous. Males reach Carolina from 37.5°N extending south to Bigtooth cookie-cutter shark maturity at 15-22 cm (6-9 in), while fe- 34.5°N, off southern Florida from 26.5°N (also referred to as Largetooth males mature at 17-20 cm (7-8 in). This extending south around Florida, includ- cookie-cutter shark) shark is the smallest on record, with an ing the Straits of Florida, and into the Isistius plutodus average size of 15-22 cm (6-9 in) and a Gulf of Mexico west to the Dry Tortugas The bigtooth cookie-cutter shark is maximum size of about 27 cm (11 in). at 83.8°W; in the Gulf of Mexico from a very rare, small shark characterized Fishing The spined pygmy shark is the Florida Panhandle east of DeSoto as epipelagic to bathypelagic, caught caught in mid-water trawls at depths of Canyon at 85.3°W, extending west to the at depths of 880-6,440 m. Only a few 200-800 m. Mississippi River Delta at 90.5°W. specimens are presently known, from Primary Distribution Within the U.S. Gulper shark Okinawa, , , Sahara Re- EEZ Available information is insuf- Centrophorusgranulosus public, and the holotype caught in the ficient to determine primary distribution Gulf of Mexico near the Mississippi at this time. The gulper shark is a deepwater spe- River Delta (Compagno, 2002c). cies found along the outermost conti- Biology The habits of the bigtooth Gulper Sharks (Centrophoridae) nental shelves and upper slopes in the cookie-cutter shark are presumed to be Gulper sharks are moderately sized Atlantic, western Indian, and western similar to those of the cookie-cutter shark. sharks, on the order of 1-1.7 m (40-68 Pacific Oceans. Specimens have been However, its more powerfuljaws, bigger in) in length. This is a group in need caught at depths of 100-1,200 m from mouth, and gigantic lower teeth (propor- of further taxonomic work. With - the Carolinas in the Atlantic, to the tionately the largest of any living shark) tentially three undescribed species in Mississippi River Delta region in the enable it to take larger bites out of its prey. U.S. waters, it is best for management northem Gulf of Mexico. This species The holotype had a plug of fish flesh in purposes to lump all species of Centro- is also abundant off Puerto Rico and the its stomach. In addition, its short snout phorus together until further informa- Virgin Islands (Russell et al., 1988). In and anteriorly positioned eyes allow for tion is available to aid in distinguishing the past, a number of other larger gulper binocular vision, and may be useful in all these species. The distributions shark species have been confused with locating prey. There are no data available appear to be similar enough so that four this species. on reproduction. The maximum known areas of concentration are found in U.S. Biology Development is ovovivipa- size of this species is 42 cm (17 in). waters: in the mid Atlantic Ocean, Gulf rous, with litters consisting of 4-6 pups Fishing A bigtooth cookie-cutter of Mexico, and Caribbean Sea, and off that measure approximately 35 cm (14 shark specimen was caught in a mid- southem Florida. in) at birth. Males mature at 85-92 cm water trawl. (34-37 in) (Yano and Kugai, 1993). The shark #1 Primary Distribution Within the U.S. Undescribed gulper maximum reported size is about 96 cm Centrophoruscf. acus EEZ Available information is insuf- (38 in). ficient to determine primary distribution A possible new species of gulper shark Fishing The gulper shark is usually at this time. similar to the Japanese gulper shark from caught in deepwater trawls. the Pacific Ocean, Centrophorms acus, PrimaryDistribution Within the U.S. Spined pygmy shark is found in the Caribbean Sea and Gulf EEZ (Map 18) In the Caribbean Sea in (also referred to as Pygmy shark) of Mexico (McEachran and Fechhelm, water depths of 100-1,200 m surround- Squaliolus laticaudus 1998). At least two specimens are known ing Puerto Rico and the U.S. Virgin The spined pygmy shark is a minute, from the Dry Tortugas. It is presumed to Islands, extending east and west to the cigar-shaped shark. The species is wide occur at depths below 200 m; specimens limits of the EEZ; in the Atlantic Ocean

65(4) 9 Figure 8.-Large female Taiwan gulper shark, Centrophorus niaukang, from fishery survey catch at Norfolk Canyon in 1998. NMFS photo.

and Gulf of Mexico in water depths of Biology Given the present difficulty , and North Atlantic Ocean 100-1,200 m: in the Atlantic off Virginia in defining the species, very little biologi- (Munoz-Chapuli and Ramos, 1989), typi- and North Carolina from 37.5°N, extend- cal information is available. Diet consists cally occurring at depths of 200-900 m. ing south to 34.5°N, off southern Florida of squid and bony fishes. Males mature In U.S. waters it has been caught at depths from 26.5°N, extending south around at 81-94 cm (32-38 in) and females at of 145-660 m within an area extending Florida, including the Straits of Florida, 75-89 cm (30-36 in) (Yano and Kugai, from Cape Hatteras, N.C., to just north and into the Gulf of Mexico west to the 1993). The largest recorded length for of Norfolk Canyon off Virginia (Moore Dry Tortugas at 83.8°W; in the Gulf of a is 100 cm (39 in) et al., 2003). Mexico from the Florida Panhandle east (McEachran and Fechhelm, 1998). Biology Taiwan gulper sharks feed of DeSoto Canyon at 85.3°W, extending Fishing The little gulper shark on bony fishes, skates, and squid. west to the Mississippi River Delta at is caught with deepwater trawls and Development is ovoviviparous, with 90.5 0W. longlines. pups measuring 35-40 cm (14-16 in) Primary Distribution Within the U.S. at birth. Males mature at 110 cm (44 Undescribed gulper shark #2 EEZ (Map 19) In the Caribbean Sea in in) and females at 146 cm (58 in). The (also referred to as Little gulper shark) water depths of 200-800 m surrounding maximum reported size is 169 cm (68 Centrophoruscf harrissoni Puerto Rico and the U.S. Virgin Islands, in) (co-author J. A. Moore, personal The little gulper shark is a small, extending east and west to the limits of observ.). Two records for U.S. waters slender shark that inhabits waters of the the EEZ; in the Atlantic Ocean and Gulf involved captures of multiple individu- continental slopes in the Gulf of Mexico of Mexico in water depths of 200-800 als, 8 sharks just north of Cape Hatteras and Caribbean Sea. Individuals have been m: in the Atlantic off Virginia and and 40 near the head of Norfolk Canyon. collected in U.S. waters from the north- North Carolina from 37.50N. extending These appeared to be feeding aggrega- eastern Gulf of Mexico (off northwestern south to 34.5°N, off southern Florida tions in shallower water at night. One Florida to Louisiana), in the Straits of from 26.5°N, extending south around unique aspect to the U.S. captures of Florida, and around Puerto Rico and the Florida, including the Straits of Florida, this species is that all 50 specimens have Virgin Islands. The typical depth range and into the Gulf of Mexico west to the been mature, gravid females. Those cap- for this species is 200-800 m. although Dry Tortugas at 83.8°W; in the Gulf of tured in February and March contained individuals have been reported in waters Mexico from the Florida Panhandle east eggs and very small embryos and those as shallow as 50 m and as deep as 1,400 of DeSoto Canyon at 85.3°W, extending captured in September had embryos m. This long-snouted species is most west to the Mississippi River Delta at at various stages of development (17- similar to the Australian dumb gulper 90.50W. 40 cm, 8-16 in), including near-term shark, Centrophorusharrissoni, and has pups (co-author J. A. Moore, personal previously been reported as Centropho- Taiwan gulper shark observ.). Given that these groups are rus uyato, although this latter name is Centrophorusniaukang segregated by sex and all examined now considered invalid (Compagno2 ). The Taiwan gulper shark (Fig. 8-10) is individuals were both large-no less a larger-bodied gulper shark distributed than 149 cm (60 in)-and gravid, it may 2Compango. L. 1999. Personal commun., South over the outer continental shelf and slope be that these groups represent breeding African Museum, Cape Town. in the South Sea, southwestern aggregations.

10 MarineFisheries Review Figure 9.-Male near-term pup of Taiwan gulper shark, Centrophorus niauikang, from fishery survey catch at Norfolk Canyon in 1999. Photo: Jon Moore.

Fishing The Taiwan gulper shark is caught with deepwater trawls and long- lines. The large size of this species makes it an attractive fishery target in the eastern Atlantic and South China Sea. PrimaryDistribution Within the U.S. EEZ(Map 20) In the Caribbean Seain water depths of 150-700 m surrounding Puerto Rico and the U.S. Virgin Islands, Figure 10.-Female Taiwan gulper shark, Centrophorms niainkang, 158 cm long, extending east and west to the limits of from Norfolk Canyon at a depth of 460-660 m, captured off Virginia during NMFS the EEZ; in the Atlantic Ocean and Gulf cruise AL98-04. Photo: Jon Moore. of Mexico in water depths of 150-700 m: in the Atlantic off Virginia and North Carolina from 37.5°N extending Florida to the Mississippi River Delta. It Arrowhead gulper shark south to 34.5°N, off southern Florida occurs at depths of 200-750 m. (also referred to as Dogfish, from 26.5°N extending south around Biology The maximum reported size flatnose dogfish) Florida, including the Straits of Florida, is 89 cm (36 in). Virtually no other bio- Deaniaproficndorum and into the Gulf of Mexico west to the logical information is available until this Dry Tortugas at 83.8°W; in the Gulf of species is distinguished from the Pacific The arrowhead gulper shark is a Mexico from the Florida Panhandle east . poorly known deepwater species, widely of DeSoto Canyon at 85.3°W extending Fishery This species is taken with distributed in bottom waters of the upper west to the Mississippi River Delta at either trawls or longlines. continental slopes worldwide. In U.S. 90.5°W. Particularly important feeding Primary Distribution Within the U.S. waters this species is caught off Virginia and potential pupping grounds have been EEZ (Map 22) In the Caribbean Sea in and North Carolina and also in the vicin- identified along the outer continental water depths of 200-750 m surrounding ity of the Mississippi River Delta. It has shelf and upper slope in the Atlantic in Puerto Rico and the U.S. Virgin Islands, been reported at depths of 300-1,800 m water depths of 150-700 m off Cape extending east and west to the limits of (Compagno, 2002d). Hatteras, N.C., beginning at 36°N and the EEZ; in the Atlantic Ocean and Gulf Biology This species is known to extending to north of Norfolk Canyon of Mexico in water depths of 200-750 feed on crustaceans, squid, and lantern- to 37.3°N (Map 21). m: in the Atlantic off Virginia and fishes. Development is ovoviviparous, North Carolina from 37.5°N, extending with 5-7 pups per litter. Males reach Undescribed gulper shark #3 south to 34.5°N, off southern Florida maturity at 43 cm (17 in) and females at Centrophorus cf. tessellatus from 26.5°N, extending south around about 70 cm (28 in). The average size is A possible new species of gulper shark Florida, including the Straits of Florida, about 50 cm (20 in), although individu- similar to the Pacific mosaic gulper shark, and into the Gulf of Mexico west to the als can reach a maximum size of 76 cm Centrophorustessellates, this species is Dry Tortugas at 83.8°W; in the Gulf of (30 in). found in the Caribbean Sea and Atlantic Mexico from the Florida Panhandle east Fishing The arrowhead gulper shark Ocean, and also in the Gulf of Mexico of DeSoto Canyon at 85.3°W, extending is caught in deepwater trawls. (McEachran and Fechhelm, 1998), where west to the Mississippi River Delta at Primary Distribution Within the U.S. specimens are known from northwestern 90.5°W. EEZ (Map 23) In the Atlantic Ocean

65(4) 11 and Gulf of Mexico in water depths of depth segregation by size has been re- Fishing Shortspine dogfish are 300-1,800 m: in the Atlantic off Virginia ported, with smaller, younger individuals usually caught in mid-water or bottom and North Carolina from 37.5°N, south found in shallower water than were the trawls. to 34.5°N; in the Gulf of Mexico from adults. Development is ovoviviparous, PrimaryDistribution Within the U.S. the Florida Panhandle east of DeSoto with observed litters of 10 embryos. EEZ (Map 25) In the Caribbean Sea Canyon at 85.3°W, extending west to the Maturity is reached at 50 cm (20 in). The in water depths of 100-750 m sur- Mississippi River Delta at 90.5 0W. average size is 75 cm (30 in), although rounding Puerto Rico and the U.S. individuals can reach 110 cm (43 in) Virgin Islands, extending east and west Spiny Dogfish Sharks (Squalidae) (Compagno, 2002e). to the limits of the EEZ; in the Atlantic Fishing The Cuban dogfish is caught Ocean and Gulf of Mexico in water Roughskin spiny dogfish in bottom trawls at depths greater than depths of 100-750 m: in the Atlantic (also referred to as 50 m. It is harvested for its liver, which extending from Cape Hatteras, N.C., at Roughskin spurdog) is used in the production of oil and 35°N, south around Florida, including Cirrhigaleusasper . the Straits of Florida, and into the Gulf The roughskin spiny dogfish is widely Primary Distribution Within the U.S. of Mexico west to the Texas/Mexico distributed in tropical to subtropical EEZ (Map 24) In the Caribbean Sea in border. waters 200-650 m deep over the upper water depths of 100-400 m surrounding continental slope (Compagno, 2002e). Puerto Rico and the U.S. Virgin Islands, Sawsharks (Pristiophoridae) Most specimens have been isolated cap- extending east and west to the limits of tures off North Carolina south to Florida, the EEZ; in the Atlantic Ocean and Gulf American sawshark (also referred in the northern Gulf of Mexico. or around of Mexico in water depths of 100-400 to as Bahama sawshark) Puerto Rico and the Virgin Islands. m: in the Atlantic extending from Cape Pristiophorusschroederi Biology The roughskin spiny dog- Hatteras, N.C., south around Florida, The American sawshark is a poorly fish is a poorly known species. Its diet including the Straits of Florida, and into known deepwater species, inhabiting includes squid and small fishes. Devel- the Gulf of Mexico west to the Texas/ waters of the continental and insular opment is ovoviviparous, with litters Mexico border. slopes. It is known only from waters of 10-22 pups reported. Although size off southeast Florida, , and the at maturity has not been determined, Shortspine dogfish (also referred Bahamas (Compagno, 2002f). Catches specimens 85 cm (33 in) long have been to as Shortspine spurdog) have occurred at water depths of 400- reported as mature. The average size is Squalus mitsukurii 1,000 m. around 90 cm (35 in), although indi- Shortspine dogfish are found through- Biology The American sawshark is viduals can reach at least 1 18 cm (46 in) out deep tropical and temperate waters easily recognized by its snout, which (Compagno, 2002e). along the continental shelves and upper is prolonged into a long blade. The Fishing Theroughskin spiny dogfish slopes. Catches have been reported from snout is equipped with teeth, actually has been caught with both hook and line North Carolina to Florida, throughout enlarged dermal denticles, on each side and gear in deep waters. the Gulf of Mexico, and off Puerto and two long barbels on the underside. Primary DistributionWithin the U.S. Rico and the Virgin Islands. This spe- The American sawshark is not to be con- EEZAvailable information is insufficient cies is taken typically by bottom trawls fused with the , the latter being a to detenrnine primary distribution at this at depths ranging from 100-750 m. shark-like ray of the batoid family Pris- time. It is often misidentified as Squalus tidae. However, the sawshark may use blainvillei, which is now considered its saw to stun and disable prey just as Cuban dogfish an invalid name (Compagno, 2002e). do. Development is assumed Squalus cubensis There is some question, however, as to to be ovoviviparous and newborns mea- The Cuban dogfish inhabits tropical to whether Squalus mitsukurii represents a sure 30 cm (12 in). The largestAmerican subtropical waters of the western Atlantic single widespread species or a complex sawshark specimen was recorded at Ocean. In U.S. waters it is very abundant of several species. 81 cm (32 in). throughout the Gulf of Mexico, along Biology The diet of the shortspine Fishing The American sawshark is the Atlantic coast from Florida to North dogfish includes crustaceans, squid, and caught in deepwater trawls. Carolina, and around Puerto Rico and small fishes. Development is ovovivipa- Primary Distribution Within the U.S. the Virgin Islands (Russell et al, 1988). rous with a two-year gestation period. EEZ (Map 26) In the Atlantic Ocean It is a bottom-dwelling species found Litters typically consist of 2-15 pups that and Gulf of Mexico in water depths along the continental shelf and uppermost measure 22-26 cm (9-10 in) at birth and of 400-1,000 m: in the South Atlantic slopes, forning dense schools at depths reach maturity at 60-70 cm (24-28 in). extending from Jupiter , Fla., at of 100-400 m (Compagno, 2002e). The average size is about 75 cm (30 in), 27°N, south around Florida, including Biology The habits and diet of this although individuals can reach 125 cm the Straits of Florida, and west to the Dry shark have not yet been reported. Some (50 in) (Compagno, 2002e). Tortugas at 83.4°W.

12 Marine FisheriesReview Figure 12.-Ghost catshark pup, Apristurus manis, from searnount top at depth of 1128-1804 m 2000. Bear Seamount cruise DEOO-11. Photo: Jon Moore (Harvard Univ. specimen MCZ 158897). Figure 11.-Live shot of an Apris- turus catshark, most likely the ghost catshark, Apristurus manis, taken at Manning Seamount. Video frame- grab from the 2003 in the Sea cruise' Photo: Mountains in the deep. Captures of this species in U.S. near Veatch Canyon, off Sea research group. waters are limited to the continental (Nakaya and Stehmann, 1998; Moore slope between Block Canyon off south- et al, 2003). ern New England and the northern Biology Development is presum- boundary of the EEZ, and also on Bear ably oviparous. Size at maturity is Catsharks (Scyliorhinidae) Seamount (Moore et al., 2003). Very estimated at 70 cm (28 in) although the little biological information is available maximum reported size is 73 cm (29 Hoary catshark for this species. in) (Nakaya and Stehmann, 1998). No Apristurus canutus Biology Development is oviparous, other information is available for this This poorly known deepwater shark i.e. eggs are laid enclosed in leathery species. was not described until 1979. The species cases. Size at maturity is estimated at Fishing The smalleye catshark is is benthic, occurring along continental 70 cm (28 in) (Nakaya and Stehmann, caught in deepwater trawls. and insular slopes in the Caribbean 1998), while the maximum size is 88 PrimaryDistribution Within the U.S. Sea and in the Straits of Florida. Most cm (35 in) (co-author J. A. Moore, pers. EEZ Available information is insuf- specimens have been accidental captures observ.). No other information is avail- ficient to determine primary distribution from West Indian waters 500-1,000 m able for this species. at this time. deep. Captures of this species in U.S. Fishing The ghost catshark is caught Smallfin catshark waters are limnited to the Straits of Florida in deepwater trawls. and around Puerto Rico and the Virgin Primary Distribution Within the U.S. Apristurus parvipinnis Islands. EEZ (Map 28) In the Atlantic Ocean The smallfin catshark is benthic along Biology Males mature at 39 cm (16 in water depths of 600-1,900 m off the upper continental and insular slopes in), while the maximum size is 46 cm (18 southem New England and Long , of the Gulf of Mexico and the Caribbean in). No other information is available. along the continental slope from Block Sea. In U.S. waters it is commonly found Fishing Thehoary catsharkis caught Canyon to Veatch Canyon. However, in deep waters of the Straits of Florida in deepwater trawls. specimens have been found up to the and Gulf of Mexico where specimens PrimaryDistribution Within the U.S. northem boundary of the EEZ, and also have been collected at depths ranging EEZ (Map 27) In the Atlantic Ocean on Bear Seamount. In the future, what from 600 to 1,200 m. It may possibly and Gulf of Mexico in water depths of is considered primary distribution for be present around Puerto Rico and the 500-1,000 m: in the Atlantic extending the species may be extended to the EEZ Virgin Islands. from Jupiter Inlet, Fla., at 27°N, south boundary. Biology Development is oviparous. around Florida, including the Straits of The average size is 45-50 cm (18-20 Florida, and west to the Dry Tortugas Smalleye catshark in), although the maximum known size at 83.40 W. Apristurus microps is 52 cm (21 in). No other information This deepwater shark was previ- is available. Ghost catshark ously known from Newfoundland and Fishing The smallfin catshark is Apristurus manis the temperate eastern Atlantic Ocean. caught in deepwater trawls. This poorly known deepwater shark Most specimens have been acciden- PrimaryDistribution Within the U.S. (Fig. 11, 12) was not described until tal captures in waters 1,000-2,200 EEZ (Map 29) In the Atlantic Ocean 1979. It is a benthic dweller on the m deep off Great Britain and South and Gulf of Mexico in water depths of continental slopes of Britain and New Africa (Nakaya and Stehmann, 1998). 600-1,200 m: in the Atlantic extending England. Most specimens have been ac- Capture of this species in U.S. waters from Ft. Lauderdale, Fla., at 26.6°N, cidental captures in waters 600-1,900 m is limited to a single specimen taken south around Florida, including the Straits

65(4) 13 Figure 13.-, Apristunrs profundorurn, (top) compared to ghost catshark, Apristurus manis (bottom) taken on 2004 Bear Seamount cruise DE04-09 from seamount top at depth of 1110-1284 m. Photo: Jon Moore.

of Florida, and into the Gulf of Mexico Primary DistributionWithin the U.S. larger size. The average adult size is 42 west to the Texas/Mexico border. EEZ In the Atlantic Ocean in water cm (17 in), with the largest recorded depths of 1,100-1.750 m, in Hudson specimen measuring Deepwater catshark 48 cm (19 in). Canyon, along the continental slope Fishing Apristurus profiindorum The broadgill catshark is to Block Canyon, and also on Bear caught in deepwater trawls. This poorly known, but not uncommon, Seamount. Primary Distribution Within the U.S. deepwater shark (Fig. 13) is in need of re- EEZ (Map 30) In the Gulf of Mexico Broadgill description to aid in distinguishing it from catshark in water depths of 700-1,500 m, from Apristurus riveri other species within this genus. Several the Straits of Florida at 79.8°W, extend- features, such as velvety texture to the This is another poorly known deep- ing west to the Mississippi River Delta denticles, no enlarged denticles at the base water shark. It is a benthic species, at 89.40W. of the anal and second dorsal fins, wider occurring along continental and insular and shorter snout, smaller dorsal fins, and slopes in water depths of 700-1,500 m. Undescribed catshark squared off posterior edges to the pectoral The species ranges from Florida, Cuba, Apristlrus cf. laurussoni and anal fins, all help distinguish this spe- and Hispaniola to the northern Gulf of Although this species is commonly cies from the ghost cat shark, Apristurus Mexico and along the Central American caught off the United States, many manis, the more common Apristurus coast south to (Compagno, authors have incorrectly identified this species off New England. Capture of this 2002g). Captures of this species in U.S. undescribed species of deepwater shark species in U.S. waters is limited to the waters are limited to the Straits of Florida as the , Apristurus holotype taken near Hudson Canyon, a and eastern Gulf of Mexico west to the laurussoni, which it closely resembles specimen from Block Canyon, and several Mississippi River Delta. This species is (Nakaya3 ). This undescribed species is individuals taken from Bear Seamount also likely to be present around Puerto found along the U.S. east coast, in the (Moore et al., 2003). It is caught in water Rico and the Virgin Islands. Gulf of Mexico, and in the Caribbean Sea depths of 1,100-1.750 m. Biology Little is known of the habits off the coasts of Honduras and Venezuela Biology Virtually no biological of the broadgill catshark. Development (Nakaya and Sato, 1998). It currently is information is available for this species. is oviparous and cases are smooth- under study by K. Nakaya. This species The holotype is a subadult male 51 cm surfaced, translucent, greenish with in- is benthic, usually over mud or other soft (20 in) long and maximum size known distinct bands of lighter color, measuring bottoms, along the continental slopes in is 59 cm. about 5.5 cm (2 in) long by 1.3 cm (0.5 Fishing The deepwater catshark is 3 in) wide. Females are believed to mature Nakaya, K. 1999. Personal commun., Hokkaido caught in deepwater trawls. at 40 cm (16 in) and males at a slightly Univ., Lab. Mar. Zoology, Japan.

14 Marine FisheriesReview water depths of 700-1,500 m. Captures of coastfromBelize to Nicaragua (Compag- Fishing The striped sawtail catshark this species in U.S. waters occur offMas- no, 2002g). It is common throughout its is usually caught in deepwater trawls. sachusetts and Delaware, and throughout range, although distribution is irregular, Primary Distribution Within the U.S. the entire northem Gulf of Mexico from and inhabits waters 250-750 m deep with EEZ (Map 34) In the Caribbean Sea in 0 Florida to Texas (Compagno, 2002g; a temperature range of 6-1 °C (43-52°F) water depths of 450-700 m surrounding Moore et al., 2003). (Springer, 1979). Adults have been found Puerto Rico and probably the U.S. Virgin Biology Very little biological in- to reside in deeper water than do juve- Islands, extending east and west to the formation is available for this species. niles. The occurrence of this species in limits of the EEZ (Nakaya and U.S. waters is in the Atlantic Ocean from Development is oviparous Boa catshark 1998). The maximum reported size North Carolina south to Florida, along the Sato, Scyliorhinus boa is 68 cm (27 in). Gulf coast of Florida, and, rarely, west of Fishing This species of catshark is Pensacola to the Mississippi River Delta Even though this deepwater shark caught in deepwater trawls. (Compagno, 2002g). was described over 100 years ago, less Primary Distribution Within the U.S. Biology The marbled catsharkfeeds than two dozen specimens exist, and EEZ (Map 31) In the Atlantic Ocean on various species of deepwater . the biology of this species is still very and Gulf of Mexico in water depths of Type of development has not yet been poorly known. This species is a benthic 700-1,500 m throughout the entire EEZ, determined, but is believed to be ovovi- dweller in water depths of 200-700 m in from the northern extent of the EEZ viparous due to the presence of eggs with- the Caribbean Sea along the continental boundary offNew England, south around out cases found inside a female. Gravid slope from Honduras south to Surinam, Florida, including the Straits of Florida, females are very seldom seen, although and also off the West Indian islands (His- and into the Gulf of Mexico west to the large numbers of females have been paniola, Jamaica, Puerto Rico, Lesser Texas/Mexico border. caught. Maturity is reached at about 27 Antilles, and Barbados). Captures of cm (11 in), with the average size approxi- this species in U.S. waters are limited to Antillean roughtail catshark mately 35 cm (14 in). The maximum size Puerto Rico, although it most likely also Galeus antillensis is estimated at 40 cm (16 in). occurs off the Virgin Islands. This small shark was formerly con- Fishing The marbled catsharkis usu- Biology A male was reportedly sidered a subspecies of the marbled ally caught in deepwater shrimp trawls. mature at 35 cm (14 in) (Springer, 1966). catshark, Galeus arae, but has recently Primary DistributionWithin the U.S. Development is probably oviparous, but been elevated to full species status. The EEZ (Map 33) In the Atlantic Ocean little is known about the reproduction and species is benthic on West Indian insular and the Gulf of Mexico in water depths biology of this species. The maximum slopes at depths of 150-700 m. Captures of 250-750 m: in the Atlantic from size is 54 cm (22 in). in U.S. waters are limited to areas off Cape Hatteras, N.C., at 36°N, extend- Fishing The boa catshark is usually Puerto Rico and the Virgin Islands. ing south around Florida, including the caught in deepwater trawls. Biology Very little biological in- Straits of Florida, and into the Gulf of PrimaryDistribution Within the U.S. formation is available. Development is Mexico west to the Mississippi River EEZ (Map 35) In the Caribbean Sea in oviparous. Size at maturity is greater than Delta at 90°W. water depths of 200-700 m surrounding 35 cm (14 in), the maximum reported size Puerto Rico and probably the U.S. Virgin Striped sawtail catshark being 46 cm (18 in) (Konstantinou and Islands, extending east and west to the Galeus springeri Cozzi, 1998). limits of the EEZ. The Antillean roughtail This recently described, small shark Fishing Blotched catshark in deepwater trawls. was formerly confused with theAntillean catshark is caught Scyliorhinus meadi PrimaryDistribution Within the U.S. roughtail catshark, Galeus antillensis, EEZ (Map 32) In the Caribbean Sea in with which it co-occurs (Konstantinou This small benthic shark is taken from water depths of 150-700 m surrounding and Cozzi, 1998). The species is a benthic upper continental slope waters at depths Puerto Rico and the U.S. Virgin Islands, dweller on West Indian insular slopes in of 300-600 m, ranging from offshore of extending east and west to the limits of water depths of 450-700 m (Konstanti- North Carolina to Florida, the Santaren the EEZ. nou and Cozzi, 1998). Captures of this , and the Bahamas (Burgess et species in U.S. waters are limited to al., 1979). In U.S. waters, captures of this Marbled catshark Puerto Rico, although it most likely also species occur off North Carolina south to (also referred to as Roughtail catshark) occurs off the Virgin Islands. the Straits of Florida. Galeus arae Biology Development is oviparous. Biology The few specimens col- The marbled catshark is a small, slen- All captured males have been immature; lected have all been immature, ranging der shark that is benthic along the conti- one female, at43 cm (17 in), was gravid. from 18 to 49 cm (7-19 in). Diet consists nental slope from North Carolina south- The maximum reported size is 44 cm of cephalopods, , euphausids, ward through the northeastem Gulf of (18 in) (Konstantinou and Cozzi, 1998). and bony fishes (Burgess et al., 1979; Mexico and along the Central American Little else is known about its biology. Parsons, 1985). Development is prob-

65(4) 15 ably oviparous, but little is known about brownish/amber in color, and possess a observed. Size at maturity is 26 cm (10 the reproduction and biology of this long tendril at each corner, which wraps in), while the largest recorded dwarf species. around bottom structures (e.g. hydroid catshark measured 32 cm (13 in). Fishing The blotched catshark is colonies, soft corals, lost fishing nets). Fishing The dwarf catshark is caught usually caught in deepwater trawls. These bottom structures are utilized by only in deepwater trawls. PrimaryDistribution Within the U.S. a number of egg-laying individuals-a Primary Distribution Within the U.S. EEZ (Map 36) In the Atlantic Ocean hydroid colony from Washington Canyon EEZ (Map 38) In the Caribbean Sea in and Gulf of Mexico in water depths of was found with as many as 300 eggs water depths of 200-600 m surrounding 300-600 m: in the Atlantic extending attached, and an abandoned trawl recov- Puerto Rico and the U.S. Virgin Islands, from Cape Fear, N.C., south around Flor- ered from Hudson Canyon similarly had extending east and west to the limits of ida, including the Straits of Florida, and many eggs attached (Able and Flescher, the EEZ; in the Atlantic Ocean and Gulf west to the Dry Tortugas at 83.5 0W. 1991). Eggs are laid in pairs at the rate of Mexico in water depths of 200-600 of one pair every 14-17 days (Castro et m: extending from Jupiter Inlet, Fla., at Chain dogfish al., 1988). Incubation time is estimated at 27°N, south around Florida, including (also referred to as Chain catshark) about one year (Castro et al., 1988). One the Straits of Florida, and west to the Dry Scyliorhinus retifer trawl off Nags Head, N.C., produced a Tortugas at 83.4°W. The chain dogfish is a small, slender large number of newly hatched or small catshark that ranges from Georges Bank, chain dogfish, suggesting that nursery Ribbontail Catsharks (Proscyllidae) offNew England, to Nicaragua, including areas may be highly localized. The pups the entire Gulf of Mexico. It is a benthic measure about 10 cm (3.9 in) at hatching, Cuban ribbontail catshark species found along the continental and (in the laboratory) grow 0.18-0.22 Eridacnisbarbouri shelf and slope at depths of 50-750 m mm/day during the first two years (Castro This species is taken as accidental (Sminkey and Tabit, 1992), usually in et al., 1988). The average size is about 38 captures from the upper continental temperatures near 8.5-11°C (47-520F), cm (15 in). The largest recorded chain slope in water depths of 400-650 m, although nursery areas can be as cold as dogfish measured 59 cm (24 in) (Castro off southem Florida, the Bahamas, and 7.4 0C (Castro et al., 1988), The species et al., 1988). Cuba (Compagno, 2002h). This is one appears to be most abundant in deep Fishing The chain dogfish is taken of the smallest shark species known and waters off Virginia and North Carolina. It by trawling in water depths greater than rivals the minute size of the spined pygmy is often taken in shallower water (50-220 75 m and at temperatures around 10°C shark. Triakis barbouri is a frequently m) in the northem parts of the range, (50°F). used synonym of this species. but more often inhabits deeper waters Primary Distribution Within the U.S. Biology Size at maturity is 27-28 cm (230-450 m or more) south of Cape Hat- EEZ (Map 37) In the Atlantic Ocean (11 in). Development is ovoviviparous teras (Able and Flescher, 1991; Castro et and Gulf of Mexico in water depths of with two pups (one per oviduct), which al., 1988). Submersible observations have 50-750 m throughout the entire EEZ, are 10 cm (4 in) at birth. The maximum found individuals in contact with bottom from the northern extent of the EEZ reported size is 34 cm (14 in). Almost structures (e.g. cerianthid , boundary off New England south around nothing else is known about the biology , and coils of cable) (Able and Florida, including the Straits of Florida, of this species. Flescher, 1991). and into the Gulf of Mexico west to the Fishing The Cuban ribbontail Biology Diet consists of (in descend- Texas/Mexico border. catshark is caught only in deepwater ing order) crustaceans, annelid worms, trawls. squid, and bony fishes, which make up Dwarf catshark Primary DistributionWithin the U.S. 90% of the food ingested (NMFS4 ). Scyliorhinus torrei EEZ (Map 39) In the Atlantic Ocean There may be geographic differences in The dwarf catshark is a small, slender and Gulf of Mexico in water depths of size at maturity: sharks off Virginia and shark known from off the southeast coast 400-650 m: in the South Atlantic ex- north were found to mature at 38-39 cm of Florida, the Bahamas, Cuba, Puerto tending from Cape Canaveral, Fla., at (15-16 in) (Able and Flescher, 1991; Rico, and the Virgin Islands. It is a ben- 28°N, south around Florida, including Sminkey and Tabit, 1992), while sharks thic species along the upper continental the Straits of Florida, into the Gulf of off Florida and South Carolina matured at slope, and has been caught at depths of Mexico to 81.5°W. 50-52 cm (20-21 in) (Castro et al., 1988). 200-600 m. Captures in U.S. waters are Development is oviparous. Egg cases are limited to southeastem Florida, Puerto False Catsharks (Pseudotriakidae) 5-6 cm (2 in) long by 2 cm (0.9 in) wide, Rico, and the Virgin Islands. Biology Analysis of stomach con- False catshark tents indicates a diet of squid and possibly Pseudotriakismicrodon 4 Unpublished data on file at the Dy- cufflefish. Nothing else is known about This large deepwater shark is ap- namics Program, NOAA, NMFS, NEFSC, Woods Hole Lab., 166 Water St., Woods Hole, MA its habits and reproduction, as neither parently wide-ranging in the Atlantic, 02543. eggs nor newly hatched pups have been Pacific, and Indian Oceans, but records

I6 Marine FisheriesReview si -,

Figure 14.-Frill shark, Chlarnydoselachiusanguineus, specimen cited in Moore et al. (2003) checklist, from southern New England slope. Photo: Rob Nawojchik (Harvard Univ. specimen MCZ 153745). of captures are sparsely distributed and it of 100-1,300 m, but sometimes in is uncommon to rare where it does occur shallower, inshore waters (Compagno, (Compagno, 1988). This species is usu- 2002i). It is widely distributed but ally taken as accidental captures on the sparsely occurring in the eastern Atlantic upper continental slope in water depths Ocean from Norway to Mauritania and of 200-1,500 m, although it occasionally from to , and in ventures inshore (Compagno, 1988). the Pacific Ocean from Chile, southern It typically is benthic over continental , Japan, , and and insular slopes. The only records of . In the western Atlantic this species in U.S. waters are single it is found off and French Figure 15.-Frill shark head, Chla- specimens-one that washed ashore at Guiana and south of Massachusetts. The nmydoselachus anguineuts, specimen Amagansett, Long Island, N.Y, in 1883 sole capture in U.S. waters is the speci- cited in Moore et al. (2003) check- from the continental slope south list, from southern New England (Goode and Bean, 1896) and another men slope. Photo: Rob Nawojchik (Har- that swam into a nearshore pound net in of Martha's Vineyard, Mass. (Moore et vard Univ. specimen MCZ 153745). Manasquam, N.J., in 1936 (Bigelow and al., 2003). Schroeder, 1953). Scyliorhinus licha is Biology The diet of this shark con- an older name for this species. sists of cephalopods, other sharks, and Bay of Fundy (only the occasional rare Biology The diet of this species bony fishes (Kubota et al., 1991; Com- stray) to Florida as well as the Gulf of consists of mostly bony fishes and pagno, 2002i). Males mature at 97 cm Mexico. This is a common shark in bays cephalopods (Yano and Musick, 1992). (39 in) and females at 135 cm (54 in). and inshore waters, usually found at Males mature at 260 cm (104 in) (Yano, Development is ovoviviparous, with depths of less than 20 m. It is frequently 1992), and females at 212 cm (85 in). 2-12 embryos produced (Tanaka et al., encountered from Cape Cod, Mass., to Development is ovoviviparous, with two 1990; Compagno, 2002i). The gestation Charleston, S.C., where it is the second embryos produced. Pups are 120 cm (48 period may be 3.5 years (Tanaka et al., most abundant shark after the spiny dog- in) at birth (Yano, 1992). The maximum 1990), with the pups measuring 55 cm fish, Squallus acanthias.This subspecies recorded size is 296 cm (118 in) (Yano (22 in) at birth (Tanaka et al., 1990). The prefers muddy or sandy bottoms. There is and Musick, 1992). maximum recorded size is 196 cm (78 in) some evidence that this species is divided Fishing The false catshark is usually (Compagno, 2002i). into several discrete populations. The caught on longlines, but occasionally Fishing The frill shark is usually most well known population ranges from swims into pound nets. caught on longlines. the Carolinas north along the coast to PrimaryDistribution Within the U.S. Primary Distribution Within the U.S. New England and southern . The EEZ Available information is insuf- EEZ Available information is insuf- species is relatively uncommon between ficient to determine primary distribution ficient to determine primary distribution North Carolina and Florida but can be at this time. at this time. found in abundance off the Florida coast. In addition, smooth dogfish catches occur Frill Sharks (Chlamydoselachidae) Smoothhound Sharks (Triakidae) frequently in many areas of the Gulf of Mexico and the Caribbean Sea. Off the Frill shark Smooth dogfish (also referred to Atlantic coast the species migrates in Chilamydoselachusanguineus as Dusky smoothhound) response to changes in water temperature, canis canis The frill shark (Fig. 14, 15) is a Mustelus moving from north to south and inshore relatively rare deepwater shark, usually In North American waters the range to offshore with the seasons. Its range benthopelagic, occurring in water depths of the smooth dogfish encompasses the of bottom temperatures in the Mid-

65(4) 17 Atlantic is 5.3-27.7°C, with most Primary Distribution Within the U.S. Biology The diet of this shark con- captures from waters with temperatures EEZ (Map 40) In the Atlantic Ocean sists mainly of crustaceans, but also ceph- of 10-22°C (NMFS5). Individuals winter and Gulf of Mexico in waters ranging alopods and fishes (Compagno, 2002j). primarily in the area between southem from shallow estuaries and inshore areas Development is viviparous, with litters North Carolina and the , out to a depth of 450 m, throughout the of 1-7 pups, which measure 21-24 cm moving up the coast to New England in entire EEZ, from the northern extent of (8-10 in) at birth. Maturation occurs at a the spring. Unpublished NMFS data 5 also the EEZ boundary offNew England south size of 43-48 cm (17-19 in) (Heemstra, show that some individuals are in deeper around Florida, including the Straits of 1997). The maximum reported size is 64 waters (80-460 m) off New York and Florida, and into the Gulf of Mexico west cm (26 in) (Heemstra, 1997). New England during the winter. to the Texas/Mexico border. Fishing The smalleye smoothhound Biology The smooth dogfish has is easily taken with hook and line using a very slender body and a prominent Antillean smoothhound squid or shrimp bait. It is also caught by behind the eye. The species feeds Mustelus canis insularis trawlers. primarily on crustaceans (72%), includ- This is a relatively common deepwater PrimaryDistribution TVithin the U.S. ing crabs, , and shrimp (NMFS4 ). shark found around islands in the Carib- EEZ Available information is insuf- Other prey includes molluscs (7%), fishes bean Sea. The subspecies is found from ficient to determine primary distribution (4%), squid (3%), and worms (2%) Barbados to Cuba, the Bahamas, and at this time. (NMFS4 ). However, the smooth dogfish in water depths of 100-800 is both an opportunistic feeder and a scav- m (Heemstra, 1997). Captures in U.S. Florida smoothhound (also referred enger and will consume whatever prey is waters are limited to Puerto Rico and to as Narrowfin smoothhound) easily available. It is primarily nocturnal the Virgin Islands, where it is abundant Mustelus norrisi and tends to be a very active shark, con- (Russell et aL, 1988). This subspecies The very slender Florida smooth- stantly patrolling the bottom for food. prefers rugged bottoms, in contrast to hound is usually found in shallow coast- Development is viviparous; i.e. embryos the continental subspecies (Heemstra, al waters with sand or mud bottoms. develop in the uterus, initially dependent 1997). The species typically moves inshore to on stored yolk, and are later nourished Biology Development is viviparous, waters 5-7 m deep or less during the by the mother through a placental con- with pups measuring 34 cm (13 in) at winter months, although specimens nection. This species enters bays and birth (Heemstra, 1997). Maturation have been caught in waters as deep as estuaries, which serve as nursery grounds occurs at a size of 80-90 cm (32-36 in). 100 m. Off Florida this species moves for young of the year (Rountree andAble, The maximum reported size is 117 cm offshore (>55 m) between May and 1996). Pups measure 34-39 cm (13-15 (47 in) (Heemstra, 1997). October (Heemstra, 1997). The Florida in) at birth, with litters usually consist- Fishing TheAntillean smoothhound smoothhound is common along the west ing of 4-20 pups (Heemstra, 1997). The is easily taken with hook and line using coast of Florida, off Texas, and also in gestation period lasts about ten months squid or shrimp bait. It is often caught the southern Caribbean Sea and the and most births occur in early summer. in large numbers by trawlers. A fishery western Atlantic Ocean south to Brazil The growth rate of this species is believed exists for this subspecies off Cuba. (Heemstra, 1997). to be very fast, with maturation occur- Primary Distribution Within the U.S. Biology This smoothhoundfeeds on ring after only one or two years and at a EEZ (Map 41) In the Caribbean Sea in crabs, shrimp, and small fishes. Develop- size of 82-90 cm (33-36 in). Although water depths of 100-800 m surrounding ment is viviparous, with litters in late the average size is about 122 cm (48 in), Puerto Rico and the U.S. Virgin Islands, winter or early spring usually consisting individuals as large as 155 cm (62 in) extending east and west to the limits of of 7-14 pups that measure 30 cm (12 m) have been reported. the U.S. EEZ. at birth. Males reach maturity at about Fishing The smooth dogfish is easily 58 cm (23 in) and females at 65 cm (26 taken with hook and line using squid or Smalleye smoothhound in). The average size is 75 cm (30 in) shrimp bait. Because of its abundance, Mustelus higmani for males and 90 cm (35 in) for females, it interferes with shrimp trawling opera- A small, active shark (the smallest although individuals exceeding 100 cm tions and affects and stocks. smoothhound species in U.S. waters), (39 in) have been reported (Heemstra, It is often caught in large numbers by the smalleye smoothhound is found 1997). shrimp trawlers. The species is exten- mainly off northern South America, Fishing The Florida smoothhound sively used as a laboratory and occurring from close inshore to a depth is often taken in nets, usually very close often displayed in aquaria. of 900 m (Heemstra, 1997). The spe- to . cies also enters brackish estuaries and Primary Distribution Within the U.S. (Compagno, 2002j). Capture in EEZ (Map 42) In the Gulf of Mexico 5 Unpublished data on file at the Ecosystems Sur- U.S. waters is limited to one specimen in waters ranging from shallow estuaries veys Branch, NOAA, NMFS, NEFSC, Woods Hole Lab., 166 Water St., Woods Hole, MA taken in the northern Gulf of Mexico and inshore areas out to a depth of 100 m 02543. off Florida. throughout the EEZ from and including

18 Marine Fisheries Review the Straits of Florida, west to the Texas/ Acknowledgments ______. 2002e. Squalidae. Dogfish sharks. FAO Species Identification Sheets for Fish- Mexico border. The authors wish to express appre- ery Purposes. Westem Central Atlantic. FAO, ciation for the information shared by Rome 1:379-385. Gulf smoothhound ______. 2002f. Pristiophoridae. Sasharks. Mustelus sinusmexicanus colleagues of Jon A. Moore. We wish FAO Species Identification Sheets for Fish- to thank L. Compagno, J. Galbraith, H. ery Purposes. Western Central Atlantic. FAO, This is an active offshore shark found Konstantinou, C. Milliken, K. Nakaya, Rome 1:417-418. along the Gulf of Mexico coast on the ______. 2002g. Scyliorhinidae. Catsharks. K. Yano, and P. Yoos for graciously FAO Species Identification Sheets for Fish- outer shelf. The species is known to occur providing unpublished information uti- ery Purposes. Western Central Atlantic. FAO, at water depths of 30-250 m (Heemstra, Rome 1:444-455. lized in this report. Special thanks are ______. 2002h. . Finback (rib- 1997), although most records are from given to L. Compagno for also allowing bontail catsharks). FAO Species Identification waters 42-91 m deep (Compagno, Moore to read a number of unpublished Sheets for Fishery Purposes. Westem Central 2002j). Captures in U.S. waters occur Atlantic. FAO, Rome 1:456-457. manuscripts. ______. 2002i. Chlamydoselachidae. Frilled from Panama City, Fla., to Texas. The authors also wish to express ap- sharks. FAO Species Identification Sheets for Biology Development is viviparous, preciation to Jose Castro for his techni- Fishery Purposes. Western Central Atlantic. with litters of 8 pups, which measure FAO, Rome 1:372-373. cal review of the manuscript, and to ____-___ 2002j. Triakidae. 39-43 cm (16-17 in) at birth. Matura- Christopher Rilling for his assistance (smoothhounds, topes). FAO Species Jdenti- tion occurs at a size of 80 cm (32 in), fication Sheets for Fishery Purposes. Westem with the geographic information system Central Atlantic. FAO, Rome 1:458-465. although the maximum reported size is (GIS) maps. Glukhov, A. A., and A. P. Kuzmichev. 1984. New 140 cm (56 in). record of Squaliolus laticaudus (Squalidae) Fishing The gulf smoothhound is Literatured Cited and Neocyttus helgae (Zeidae ) in the North- eastAtlantic. Vopr. Ikhtiol. 4:669-671. easily taken with hook and line using Able, K. W., and D. Flescher. 1991. Distribu- Goode, G. B., and T. H. Bean. 1896. Oceanic squid or shrimp bait. It is also caught by tion and habitat of chain dogfish, Scyliorhzi- , a treatise on the deep-sea and trawlers. nus retifer, in the Mid-Atlantic Bight. Copeia pelagic fishes of the world, based chiefly upon (1991(l):231-234. the collections made by the steamers Blake, PrimaryDistribution Within the U.S. Bigelow. H. B., and W. C. Schroeder. 1953. Albatross, and Fish Hawvk in the northwestem EEZ (Map 43) In the Gulf of Mexico Fishes of the Gulf of Maine. U.S. Fish Wildl. Atlantic, with an atlas containing 417 figures. Serv., Fish. Bull. 53:1-577b. Spec. Bull. U.S. Natl. Mus. 2:1-553. in waters ranging from shallow estuaries Burgess,G.H.,G.W.Link,Jr.,andS.W.Ross.1979. Heemstra. R C. 1997. A review of the smooth- and inshore areas out to a depth of 100 m: Additional marine fishes new or rare to Caro- hound sharks (genus Mustelus, family Tria- from Panama City, Fla., at 86°W, west to lina waters. Northeast Gulf Sci. 3(2):74-87. kidae) of the westem Atlantic Ocean, with Castro, J. I. 1983. Sharks of North American descriptions of two new species and a new the Texas/Mexico border. Waters. Texas A&M Univ. Press, College Sta- subspecies. Bull. Mar. Sci. 60(3):894-928. tion, Texas, 180 p. Herdendorf, C. E., and T. M. Berra. 1995. A Sand Tiger Sharks (Odontaspidae) ______P. M. Bubugis, and N. A. Over- Greenland shark from the wreck of the SS strom. 1988. The reproductive biology of the Central America at 2,200 meters. Trans. Am. chain dogfish, Scyliorlzinis retifer. Copeia Fish. Soc. 124(6):950- 953. Ragged-tooth shark (also referred 1988(3):740-746. ICES. 1995. Report of the Study Group on the to as Smalltooth ) Clark, E., and E. Kristof. 1990. Deep-sea elas- Biology and Assessment of Deep-sea Fisher- Odontaspisferox mobranchs observed from submersibles of ies Resources. Intemational Council for the Bermuda, Grand Cayman, and Freeport, Exploration of the Sea (ICES), C.M. 1995/ This is a large deepwater shark found Bahamas. In H. L. Pratt Jr.. S. H. Gruber, and Assess:4, 91 p. T. Taniuchi (Editors), Elasmobranchs as living Konstantinou, H., and J. R. Cozzi. 1998. Galeus over continental and insular shelves and resources: advances in the biology, , springeri, a new species of sawtail catshark upper slopes in water depths of 13-420 systematics, and the status of the fisheries, p. from the Caribbean Sea (, 269-283. U.S. Dep. Commer, NOAA Tech. Scyliorhinidae). Copeia 1998(1):151-158. m. Although it is widely distributed and Rep. NMFS 90. Kubota, T., Y. Shiobara, and T. Kubodera. 1991. nearly cosmopolitan, it is rarely captured. Compagno, L. J. V. 1984. FAO Species Cata- Food habits of the Chlainydose- The one U.S. record is a single speci- logue. Vol. 4. Part I and 2. An annotated and lachus anguineus collected from Suruga illustrated catalogue of shark species known Bay, central Japan. Bull. Jpn. Soc. Sci. Fish. men taken off North Carolina (Sheehan, to-date. FAO Fish. Synop. 125. FAO U.N., 57(1): 15-20. 1998). Rome, 655 p. McEachran, J. D., and J. D. Fechhelm. 1998. . 1988. Sharks of the Order Carcha- Fishes of the Gulf of Mexico, Vol. 1. Univ. Biology Very little biological infor- rhiniformes. Princeton Univ. Press, 486 p. Texas Press. Austin. mation is available for this species. Its ___._ 2002a. Etmopteridae. Lantern sharks Moore, J. A. 1999. Deep-sea finfish fisheries: diet consists of small bony fishes, squid, (black dogfishes). FAO Species Identification Lessons from history. Fisheries (Bethesda, and shrimp. Young measure 105 cm (42 Sheets for Fishery Purposes. Western Central MD) 24(7):16-21 Atlantic. FAO, Rome 1:393-401. ______I K. E. Hartel, J. E. Craddock, and in) at birth. Males mature at 275 cm (110 FAG-. 2002b. Somniosidae. Sleeper sharks. J. K. Galbraith. 2003. An annotated list of in), although the maximum reported size FAO Species Identification Sheets for Fish- deepwater fishes from off the New England ery Purposes. Western Central Atlantic. FAO, region, with new area records. Northeast. Nat. is 360 cm (144 in). Rome 1:402-407. 10(2):159-248. Fishing Ragged-tooth sharks are ______. 2002c. Dalatiidae. Kitefin sharks. Munoz-Chapuli, R., and F. Ramos. 1989. Review caught with hook and line and in trawls. FAO Species Identification Sheets for Fish- of the Centrophorms sharks (Elasmobrachii, ery Purposes. Western Central Atlantic. FAO, Squalidae) of the eastern Atlantic. Cybium Primary Distribution Within the U.S. Rome 1:410-414. 13(1):65-81. EEZ Available information is insuf- . 2002d. Centrophoridae. Gulper Musick, J. A., and J. D. McEachran. 1969. The sharks. FAO Species Identification Sheets for squaloid shark Echinorhinis brncus off Vir- ficient to determine primary distribution Fishery Purposes. Western Central Atlantic. ginia. Copeia 1969:205-206. at this time. FAO, Rome 1:386-392. Nakaya, K., and K. Sato. 1998. Taxonomic

65(4) 19 review of Apristurus laurussoni (Saemunds- Atlantic, with a redescription of Etmopterus Tanaka, S., Y. Shiobara, S. Hioki, H. Abe, G. son, 1922) from the eastern North Atlantic hillianus. Bull. Mar. Sci. 60(3):1060-1073. Nishi, K Yano, and K. Suzuki. 1990. The (: Scyliorhinidae). Cybium Schwartz, F. J. 1993. A North Carolina capture reproductive biology of the frilled shark, 22(2):149-157. of the bramble shark. Echinorlzinus brucus, Clhlamydoselachus anguineus, from Suruga and M. Stehmann. 1998. A new species Family Echinorhinidae, the fourth in the west- Bay, Japan. Jpn. J. Ichthyol. 37(3):273-291. of deep-water catshark, Apristuirs aplhyodes ern North Atlantic. J. Elisha Mitchell Sci. Soc. USDOC. 1999. Final Fishery Management Plan n. sp., from the eastern North Atlantic (Chon- 109:158-162. for Atlantic tunas, swordfish, and sharks. U.S. drichthyes: : Scyliorhini- Scott, W. B., and M. G. Scott. 1988. Atlan- Dep. Commer., NOAA, NMFS, Vol. I-HII. dae). Arch. Fisch. Mar. Res. 46(l):77-90. tic fishes of Canada. Univ. Toronto Press, 2003. Final Amendment I to the Parsons. G. R. 1985. Notes on the life history of Toronto, 731 p. Fishery Management Plan for Atlantic tunas, the catshark, Scyliorhzinus mzeadi. Fish. Bull. Sheehan, T. E 1998. First record of the ragged- swordfish, and sharks. U.S. Dep. Commer., 83:695-696. tooth shark, Odontaspis ferox, off the U.S. NOAA, NMFS. Retzer, M. E. 1990. New records and range exten- Atlantic Coast. Mar. Fish. Rev. 60(l):33-34. Wetherbee, B. M., and G. L. Crow. 1996. First sions of twelve species of fishes in the Gulf of Shirai, S., and H. Tachikawa. 1993. Taxonomic record of the squaloid shark Scymnodon squa- Mexico. Northeast Gulf Sci. 11(2):137-142. resolution of the Etmopterus pusillus spe- mulosus from the Hawaiian Islands. Ichthyol. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. cies group (Elasmobranchii, Etmopteridae), Res. 43(3):334-339. Brooker, E. A. Lachner. R. N. Lea, and W. B. with description of E. bigeloivi, n. sp. Copeia Yano, K. 1992. Comments on the reproductive Scott. 1991. Common and scientific names of 1993(2):483-495. mode of the false cat shark, Pseudotriakis fishes from the United States and Canada. Fifth Silva, H. M. 1988. Growth and reproduction of microdon. Copeia 1992(2):460-468. Ed., Am. Fish. Soc.. Spec. Publ. 20, 183 p. kitefin shark Dalatias licha (Bonn, 1788) in and K. Kugai. 1993. Taiwan gulper Rountree, R. A., and K W. Able. 1996. Seasonal Azorean waters. ICES Council Meeting 1988 shark, Centrophornsniaukang, from the Oki- abundance, growth, and foraging habits of Collected Papers, C.M.1988/G:21. nawa Islands, Japan. Bull. Seikai Natl. Fish. juvenile smooth dogfish, Mustelus canis, in a Sminkey. T. R., and C. R. Tabit. 1992. Reproduc- Res. Inst. 71:41-49. New Jersey . Fish. Bull. 94:522-534. tive biology of the chain dogfish, Scyliorhirnus ______and J. A. Musick. 1992. Comparison Russell, G. M., E. J. Gutherz, and C. A. Barans. retifer, from the Mid-Atlantic Bight. Copeia of morphometrics of Atlantic and Pacific 1988. Evaluation of demersal longline gear 1992(1):25 1-253. specimens of the false catshark, Pseudotria- off South Carolina and Puerto Rico with Springer, S. 1966. A review of the western Atlan- kis microdon, with notes on stomach contents. emphasis on deep-water . Mar. tic catsharks, Scyliorhinidae, with descrip- Copeia 1992(3):877-886. Fish. Rev. 50(1):26-31. tions of a new genus and five new species. and S. Tanaka. 1984. Review of the Schofield, P. J.. and G. H. Burgess. 1997. Etnop- Fish Wildl. Serv., Fish. Bull. 65:581-624. deep sea squaloid shark genus Scymnodon off terus robinsi (Elasmobracnhii, Etmopteridae). . 1979. A revision of the catsharks, Japan, with a description of a new species. J. a new species of deep-water lantern shark family Scyliorhinidae. U.S. Dep Commer., Jpn. Ichthyol. 30(4):341-360. from the Caribbean Sea and western North NOAA Tech. Rep. NMFS Circ. 422:1-152.

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TITLE: Deepwater and Other Sharks of the U.S. Atlantic Ocean Exclusive Economic Zone SOURCE: Mar Fish Rev 65 no4 2003 WN: 0300401780001

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