Marine Fishes of Arctic Canada 9781442667297

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MARINE FISHES OF ARCTIC CANADA

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Marine Fishes of Arctic Canada Edited by

BRIAN W. COAD

and

JAMES D. REIST

With contributions by

Peter Rask Møller, Claude B. Renaud, Noel R. Alfonso, Karen Dunmall, Michael Power, Chantelle D. Sawatzky, Fikret Berkes, J. Brian Dempson, Les N. Harris, and Heidi K. Swanson

CA NA D I A N MUS E UM O F NAT U RE and UN I V ER S I T Y O F TO R O NTO P R E SS T o r o n t o B u ffa lo  Lo n d o n

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© Canadian Museum of Nature 2018 Published by University of Toronto Press Toronto Buffalo London www.utorontopress.com Printed in Canada ISBN 978-1-4426-4710-7 Printed on acid-free paper Jacket and interior design by Peter Ross, Counterpunch Inc.

Library and Archives Canada Cataloguing in Publication Marine fishes of Arctic Canada / edited by Brian W. Coad and James D. Reist ; with contributions by Peter Rask Møller, Claude B. Renaud, Noel R. Alfonso, Karen Dunmall, Michael Power, Chantelle D. Sawatzky, Fikret Berkes, J. Brian Dempson, Les N. Harris, and Heidi K. Swanson. Includes bibliographical references and index. ISBN 978-1-4426-4710-7 (cloth) 1. Marine fishes – Canada, Northern. I. Coad, Brian W., editor II. Reist, J.D. (James Douglas), 1951–, editor QL626.M37 2018

597.17709719

C2017-901932-5

University of Toronto Press acknowledges the financial assistance to its publishing program of the Canada Council for the Arts and the Ontario Arts Council, an agency of the Government of Ontario.

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CONTENTS

Dedication, Brian W. Coad  viii

Keys

Acknowledgments, Brian W. Coad and James D. Reist  xii

Introduction, Brian W. Coad  88 Families, Brian W. Coad  91 Species in Individual Families, Brian W. Coad (Zoarcidae by Peter Rask Møller; Gadiforms by Brian W. Coad and Claude B. Renaud)  108

Introduction Purpose of the Book, Brian W. Coad  2 Biodiversity, Brian W. Coad and Noel R. Alfonso  3 New Information, Brian W. Coad  8 Environment, James D. Reist and Chantelle D. Sawatzky  9 History of Research, James D. Reist  30

Family and Species Accounts Introduction, Brian W. Coad  158

Family 1. Myxinidae – Hagfishes, Myxines (1), Claude B. Renaud  162

Climate, James D. Reist  53



Arctic Fish, Northern Cultures, and Traditional Ecological Knowledge, Fikret Berkes  57

Family 2. Petromyzontidae – Lampreys, Lamproies (1), Claude B. Renaud  164



Family 3. Rhinochimaeridae – Longnose Chimaeras, Chimères à long nez (2), James D. Reist  167



Family 4. Chimaeridae – Shortnose Chimaeras, Chimères (1), James D. Reist  171



Family 5. Scyliorhinidae – Cat Sharks, Roussettes (1), Brian W. Coad  173



Family 6. Etmopteridae – Lantern Sharks, Requins-lanternes (1), James D. Reist  175



Family 7. Somniosidae – Sleeper Sharks, Somniosidés (3), James D. Reist  178



Family 8. Rajidae – Skates, Raies (9), James D. Reist  185



Family 9. Acipenseridae – Sturgeons, Esturgeons (1), James D. Reist  203

Habitats, James D. Reist  41

Fisheries, James D. Reist  61 Scientific Names, Brian W. Coad  67 Fish Structure, Brian W. Coad  68 Collecting and Preserving Fishes,Brian W. Coad  70

Checklists of Species Marine Species,Brian W. Coad  74 Extralimital Species,Brian W. Coad  80 Brackish Water Species,Brian W. Coad  84



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Family 10. Notacanthidae – Deep-sea Spiny Eels, Poissons-tapirs à épines (2), Brian W. Coad  205



Family 11. Synaphobranchidae – Cutthroat Eels, Anguilles égorgées (2), Brian W. Coad  208



Family 29. Macrouridae – Grenadiers, Grenadiers (9), Claude B. Renaud and Brian W. Coad  333



Family 30. Moridae – Codlings, Moros (3), Claude B. Renaud  346



Family 31. Phycidae – Phycid Hakes, Phycidés (4), Claude B. Renaud  351



Family 12. Nemichthyidae – Snipe Eels, Poissons-avocettes (1), Brian W. Coad  211





Family 13. Serrivomeridae – Sawpalates, Serrivomers (1), Brian W. Coad  213

Family 32. Gadidae – Cods, Morues (8), Claude B. Renaud  358





Family 14. Saccopharyngidae – Swallowers, Avaleurs (1), Brian W. Coad  215

Family 33. Bythitidae – Viviparous Brotulas, Donzelles vivipares (1), Brian W. Coad  376





Family 15. Eurypharyngidae – Gulpers, Grandgousiers (1), Brian W. Coad  217

Family 34. Lophiidae – Goosefishes, Baudroies (1), Brian W. Coad  378





Family 16. Clupeidae – Herrings, Harengs (2), Brian W. Coad  219

Family 35. Himantolophidae – Footballfishes, Poissons-football (1), Brian W. Coad  380



Family 17. Argentinidae – Argentines, Argentines (1), Brian W. Coad  224



Family 36. Oneirodidae – Dreamers, Rêveurs (3), Brian W. Coad  382



Family 18. Microstomatidae – Pencilsmelts, Microbecs (1), Brian W. Coad  226



Family 37. Ceratiidae – Seadevils, Poissons-pêcheurs (1), Brian W. Coad  386



Family 19. Platytroctidae – Tubeshoulders, Circés (5), Brian W. Coad  227



Family 38. Gigantactinidae – Whipnoses, Tacts géants (1), Brian W. Coad  388



Family 20. Alepocephalidae – Slickheads, Alépocéphales (6), Brian W. Coad  233





Family 21. Osmeridae – Smelts, Éperlans (3), Brian W. Coad  240

Family 39. Melamphaidae – Ridgeheads, Poissons-heaumes (1), Brian W. Coad  390



Family 40. Anoplogastridae – Ogrefishes, Ogres (1), Brian W. Coad  392



Family 41. Trachichthyidae – Slimeheads, Hoplites (1), Brian W. Coad  394



Family 42. Gasterosteidae – Sticklebacks, Épinoches (2), Chantelle D. Sawatzky and James D. Reist  396



Family 43. Scorpaenidae – Scorpionfishes, Scorpènes (3), Brian W. Coad  401



Family 44. Cottidae – Sculpins, Chabots (14), Brian W. Coad  407



Family 22. Salmonidae – Trouts and Salmons, Truites et Saumons (17), James D. Reist (with J. Brian Dempson, Karen Dunmall, Les N. Harris, Michael Power, and Heidi K. Swanson)  248



Family 23. Gonostomatidae – Bristlemouths, Cyclothones (2), Brian W. Coad  302



Family 24. Sternoptychidae – Marine Hatchetfishes, Haches d’argent (1), Brian W. Coad  305



Family 25. Stomiidae – Dragonfishes, Dragons à écailles (6), Brian W. Coad  307





Family 26. Notosudidae – Waryfishes, Guetteurs (1), Brian W. Coad  315

Family 45. Agonidae – Poachers, Poissons-alligators (3), Brian W. Coad  435





Family 27. Paralepididae – Barracudinas, Lussions (4), Brian W. Coad  317

Family 46. Psychrolutidae – Fathead Sculpins, Chabots veloutés (3), Brian W. Coad  440





Family 28. Myctophidae – Lanternfishes, Poissons-lanternes (8), Brian W. Coad  322

Family 47. Cyclopteridae – Lumpfishes, Poules de mer (5), Brian W. Coad  446



Family 48. Liparidae – Snailfishes, Limaces de mer (11), Brian W. Coad  454

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Family 49. Caristiidae – Manefishes, Caristes (1), Brian W. Coad  471

Glossary, Brian W. Coad  573



Family 50. Zoarcidae – Eelpouts, Lycodes (32), Peter Rask Møller  473

Illustration Credits, Brian W. Coad  590



Family 51. Stichaeidae – Pricklebacks, Stichées (9), Brian W. Coad  515



Family 52. Pholidae – Gunnels, Sigouines (1), Brian W. Coad  528



Family 53. Anarhichadidae – Wolffishes, Poissons-loups (4), Brian W. Coad  530



Family 54. Chiasmodontidae – Black Swallowers, Grands avaleurs (1), Brian W. Coad  539



Family 55. Ammodytidae – Sand Lances, Lançons (2), Brian W. Coad  541



Family 56. Trichiuridae – Cutlassfishes, Sabres de mer (1), Brian W. Coad  545



Family 57. Stromateidae – Butterfishes, Stromatées (1), Brian W. Coad  547



Family 58. Pleuronectidae – Righteye Flounders, Plies (9), Noel R. Alfonso   549



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Bibliography, Brian W. Coad and James D. Reist  587 Index  603

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DEDICATION Donald Evan McAllister, 1934–2001

Brian W. Coad

Marine Fishes of Arctic Canada is dedicated with affection and respect to Don McAllister in recognition of his contributions to the study of Arctic marine fishes and in acknowledgment of his decades of work that formed the foundation and impetus of this book. Don was hired as Curator of Fishes at the National Museum of Canada (now the Canadian Museum of Nature, Ottawa) on 1 August 1958. His primary research endeavour for the next thirty-six years was centred on Arctic marine fishes. His aim was to complete a book in monographic form on this fauna to complement and improve on others that had appeared on Pacific coast fishes (1946, rewritten in 1973), Atlantic coast fishes (1966, rewritten in 1988), and freshwater fishes (1973). Don published forty-nine articles relevant to Arctic marine fishes, as well as others not listed here on freshwater fishes in Canada’s north. He was the first author on thirty-five of these and collaborated with twenty-four co-authors. The numbered bibliography below forms an outline of his research in this field. He began publishing on the Arctic in 1960 with a preliminary, annotated checklist of Canada’s marine fishes2 followed by keys to the Arctic fishes3. The latter benefited from Don’s first foray into Russian1, a language Don taught himself to read so that he could access works complementary to his studies on Canadian Arctic fishes. The Canadian Museum of Nature’s collections in 1958 comprised only 4,368 specimens from all of Canada and the rest of the world. Don planned and carried out fieldwork to enhance this limited material base, with particular emphasis on Arctic marine fishes, eventually making the museum’s collection of the latter the best in the world. He made several expeditions, among them one to the Herschel Island area in the Yukon with J.G. Hunter on the M.V. Salvelinus in 19605, where thirty species of fish were taken and new records and taxonomic decisions were made. Other expeditions were taken with museum colleagues Edward L. Bousfield in 19616 along the southeastern Alaska coast, and Arthur H. Clarke in 196310,11 on the east coast of Hudson Bay (the “Tnuk Expedition”). Don not only collected fishes from the shore and from boats but, as an avid scuba diver, also collected under water. Under Don’s direction, staff and contractors participated in or carried out many

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subsequent expeditions. Continuing work, and the need for work in the Arctic, based on the materials collected was conveyed to peers in various papers, notes, and presentations15,17–18,23,32,39–42. He was to have been an editor of this book before his untimely death, and his last article on the Arctic was a poster presentation on this project49. At 14 January 2013, the Canadian Museum of Nature’s collection of Arctic marine fishes stood at 4,777 collection lots and 57,623 specimens. In the days before online collection databases, Don summarized the museum’s holdings of Arctic fishes in two publications: one listing all the marine fish species, with curatorial assistant Michèle Steigerwald, in 198231; and one mapping solely Arctic fishes in a distributional atlas33. The former summarized the Arctic holdings at that date as 135 species, 4,410 collections, and 53,742 specimens, a tremendous increase from twenty years before, when one recalls that this was only part of the collection’s growth. The latter was also co-authored with Michèle Steigerwald and in particular with Jerry Hunter and Shirley Leach, both members of the Arctic Biological Station, DFO, with whom Don had a long-term collaborative relationship in accessioning and identifying collections from DFO research activities in the north. Type-specimens in the fish collection were detailed in 196512 and included, for example, the holotype of Cyclopterus lumpus hudsonius from Fort Churchill, Hudson Bay. Specimens were often collected, preserved, identified, labelled, catalogued, shelved, retrieved, measured, counted, and re-shelved alone in the early days. Don catalogued the new material using an original and innovative system he had developed that also recorded ecological data as well as the traditional tombstone data. He also pioneered computerization of the collections to facilitate data retrieval and mapping of distributions. The computer used in the 1970s featured a screen with a single text line and such limited power that a mapping program had to run overnight. Not infrequently, the paper feed stuck, and fifteen hours of inking would be confined to one very deep and dark spot on an expanse of white. A collection of line drawings based on selected specimens in the collection was accumulated over decades in competition with other scientists for the limited time of a staff artist or for scarce funds. This

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proved to be an invaluable base for Marine Fishes of Arctic Canada and for other works on Canadian fishes. Don was a firm believer in compiling bilingual lists of names to foster understanding and communication. He learnt French well enough to be exempt from the civil service’s language examinations and collaborated with colleagues in Québec such as Vianney Legendre, along with Jerry Hunter, to produce in 1975 a list of the French, English, and scientific names of the marine fishes of Arctic Canada22. In 1987 this list became quadrilingual with Inuktitut, French, English, and scientific names38, and in 1990 he expanded the list to encompass all Canadian species of fish, from all three coasts and from fresh waters43. He reported 137 species on the Arctic coast. Some of the rarer species had no common name, and for these Don coined new ones, some of which proved mildly controversial. The dearth of information on Arctic fishes led Don to build up a large collection of reprints and copies of published works along with books and translations. He developed an extensive reprint-exchange program through contacts with colleagues around the world, particularly Russian scientists. This information was disseminated in bibliographies in 196613 and revised with Michèle Steigerwald in 198737. The former was 16 pages long with 278 references, and the latter 108 pages with 1,111 references. The online bibliography for this book exceeds 5,300 references. Don also published a bibliography with Iola Gruchy on Smelts, an important family of fishes in Arctic waters19. Don had long held strong beliefs on the need for the conservation of species and was active in working with the Committee on the Status of Endangered Wildlife in Canada (COSEWIC). In 1985 this work was summarized in a publication with Brad Parker and Paul McKee, on the rare, endangered, and extinct fishes in Canada34, mostly on freshwater fishes but also mentioning two Arctic marine species. Status reports on these two species, the Bering Wolffish, Anarhichas orientalis, and the Blackline Prickleback, Acantholumpenus mackayi, were co-authored with J. Houston in 199044–45. Both species were considered vulnerable. His environmental concerns for Arctic waters were expressed in papers on the James and Hudson Bays hydro projects46,48 and on global warming47. Completion of a series of revisionary papers on Arctic taxa was planned to place the systematics of the Arctic ichthyofauna on a firm basis but was continually delayed by Don’s forays into other fields and by the demands and opportunities of his position as Curator of Fishes. His work in other fields has been reviewed in obituaries by Cook, Gruchy, and Coad (2001), Cook and Coad (2002), Cook, Coad, and Renaud (2002), Coad (2011), and Cook, Coad, Renaud, Gruchy, and Alfonso (2011). Nevertheless Don carried out systematic studies on various taxa in Arctic and adjacent waters. He first tackled the Smelts (Osmeridae), producing a revision of the family in 19637 that included new taxa from Japan and data on Arctic Canadian samples. Systematic data on the Smelts were analysed using numerical taxonomy in 1967 14, and comments were made on the utility of this new technique. With Russian colleague V.A. Kljukanov, Don reviewed the taxonomy, systematics, and biology of the family for the book Clofnam: Check-List of the Fishes of the North-Eastern Atlantic and of the Mediterranean, which extends its coverage into Arctic waters20.That



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short segment contained a dissenting opinion of Don, the junior author, ever of independent thought. Don investigated the variation and characters of sculpins in several papers. The first with Arctic marine content was on Myoxocephalus thompsonii and its “parent” species M. quadricornis, which included systematic and ecological analyses4. The genera Artediellus, Icelus, and Triglops were examined in 19638, and keys, descriptions, and systematic conclusions were given from new material. His paper on mandibular pore patterns in the Family Cottidae included various Arctic taxa and demonstrated their utility as characters16. With curatorial assistant Jadwiga Aniskowicz in 1976, Don published the results of studies on the vertebral number in the Myoxocephalus quadricornis complex25, concluding provisionally that coastal marine forms should be called Myoxocephalus quadricornis hexacornis. A series of papers were published on the Eelpouts (Family Zoarcidae) of northern waters. An early study with E.I.S. Rees in 19649 was a revision of the worldwide genus Melanostigma, now known to have one species in Arctic Canada. The paper included a new genus, Oidiphorus. A new species of Arctic eelpout, Lycodes sagittarius, from the Beaufort Sea (Alaska) and the Kara Sea (former USSR), was described by Don in 197524, and, with M. Eric Anderson and Jerry Hunter, the deep-water Eelpouts from Arctic Canada and Alaska were revised in 198130. This latter paper gives a key to the Arctic Canadian zoarcids and detailed descriptions, illustrations, maps, and biology. One species, Lycodes frigidus, had a range extension of over 6,500 km, and this Glacial Eelpout was recorded in 1986 with N.J. Prouse36 in the Arctic Canadian Basin at 85°48' N, 350 km from the North Pole. Recognition of his contributions to polar ichthyology came in the description of new species of Eelpouts named Oidiphorus mcallisteri by M. Eric Anderson (1988) and Lycodes mcallisteri by Peter Rask Møller (2001). Another systematically difficult family in Arctic water is that of the Snailfishes (Liparidae), and it was revised with Ken Able in 198129. The layout of this work is similar to the one on zoarcids30 and shows the kind of monographic treatment that Don planned for all the species in the Arctic waters of Canada. Collaboration with a Russian colleague, Anatoly Andriashev, in 1977 and 1978 had clarified the confused status of the presumed Arctic zoarcid fish Ophidium parrii, which turned out to be a common species of Snailfish27–28. Don’s last contribution to the systematics of Arctic fishes was on biochemical evidence of speciation in the cod genus Gadus in collaboration with his doctoral student Claude Renaud and with Larry Speers, Carleton University, Ottawa, and Sami Qadri of the University of Ottawa35. Don also accumulated data on the ecology of these fishes along Canada’s least accessible coast, co-operating extensively with Jerry Hunter of the Arctic Biological Station, and compiling the files used in this book. An overview of the ecology of Arctic fishes was published in 197726. The authors of Marine Fishes of Arctic Canada are pleased to dedicate it to the memory of their colleague and hope that it will stimulate further work on the systematics, biology, and conservation of this ichthyofauna that for so long engrossed Don’s thoughts and endeavours.

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Arctic Marine Fish Bibliography of Donald Evan McAllister A complete bibliography of works by Don McAllister, numbering over 625, can be found in Coad (2011). 1. McAllister, Don E. 1960. Preliminary translation of keys of: Observations on the eel-like lycodids [Lycenchelys Gill (Pisces, Zoarcidae) and related forms] in the seas of U.S.S.R. and neighbouring waters. Fisheries Research Board of Canada Translation Series, 278: 1–4. Translated from A.P. Andriashev, Trudy Zoologicheskova Instituta Akademii Nauk SSSR, 18 (1955): 349–84. 2. McAllister, D.E. 1960. List of the marine fishes of Canada. National Museum of Canada Bulletin, 168 [Biological Series, 62]: iv + 76 pp., 1 table. 3. McAllister, D.E. 1960. Keys to the marine fishes of Arctic Canada. Natural History Papers, National Museum of Canada, 5, 1–21, 27 figures. 4. McAllister, D.E. 1961. The origin and status of the deepwater sculpin, Myoxocephalus thompsonii, a Nearctic glacial relict. National Museum of Canada Bulletin, (Contributions to Zoology), 172, 1959 [Biological Series, 65]: 44–65, 3 figures, 6 tables. 5. McAllister, D.E. 1962. Fishes of the 1960 “Salvelinus” Program from western Arctic Canada. National Museum of Canada Bulletin, (Contributions to Zoology), 185, 1962 [Biological Series, 69]: 17–39, 4 figures. 6. Bousfield, E.L., & McAllister, D.E. 1962. Station list of the National Museum marine biological expedition to southeastern Alaska and Prince William Sound. National Museum of Canada Bulletin, (Contributions to Zoology), 183, 1960–1 [Biological Series, 68]: 76–103, 2 figures, 5 maps. 7. McAllister, D.E. 1963. A revision of the smelt family, Osmeridae. National Museum of Canada Bulletin, 191 [Biological Series, 71]: iv + 53 pp., 14 figures, 2 tables. 8. McAllister, D.E. 1963. Systematic notes on the sculpin genera Artediellus, Icelus, and Triglops on Arctic and Atlantic coasts of Canada. National Museum of Canada Bulletin, (Contributions to Zoology), 185, paper 4: 50–9, 8 figures, 3 tables. 9. McAllister, D.E., & Rees, E.I.S. 1964. A revision of the eelpout genus Melanostigma with a new genus and with comments on Maynea. National Museum of Canada Bulletin, (Contributions to Zoology), 199, paper 5 (1963): 85–110, 9 figures, 3 tables. 10. McAllister, D.E. 1964. Dans l’Arctique ... au bout du monde. Le Jeune Scientifique, 3(1): 17–19, 4 figures. 11. McAllister, D.E. 1964. Fish collections from eastern Hudson Bay. Canadian Field-Naturalist, 78(3): 167–78, 8 figures, 1 table. 12. McAllister, D.E. 1965. Type specimens of fishes in the National Museum of Canada with brief historical notes. Natural History Papers, National Museum of Canada, 31: 1–13, 2 figures. 13. McAllister, Don E. 1966. Bibliography of the marine fishes of Arctic Canada. Institute of Fisheries, University of British Columbia, Vancouver, Museum Contribution, 8: 1–16. 14. McAllister, D.E. 1967. Numerical taxonomy and the smelt family, Osmeridae. Canadian Field-Naturalist, 80(4) (1966): 227–38, 2 figures, 4 tables.

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15. McAllister, D.E. 1968. Northern field projects of the National Museum of Natural Sciences, 1968. The Arctic Circular, 18(1/2): 33. 16. McAllister, D.E. 1968. Mandibular pore pattern in the sculpin family Cottidae. National Museum of Canada Bulletin (Contributions to Zoology), 223(4): 58–69, 1 figure, 2 tables. 17. McAllister, D.E. 1969. Proposed activities of the National Museum of Natural Sciences, 1969. The Arctic Circular, 19(2): 38–9. 18. McAllister, D.E. 1970. Arctic activities of the National Museum of Natural Sciences, 1969 and 1970. The Arctic Circular, 20(1): 17–18. 19. Gruchy, I.M., & McAllister, Don E. 1973. A bibliography of the smelt family, Osmeridae. (Fisheries Research Board of Canada Technical Report No. 368, 1972): ii + 104 pp. 20. Kljukanov, V.A., & McAllister, D.E. 1973. Osmeridae. In J.C. Hureau & T. Monod (Eds.), Clofnam: Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean / Catalogue des poissons de l’Atlantique du nord-est et de la Méditerranée (vol. 1, pp. 158–9). Volume 1: xxii + 683 pp. Paris: UNESCO. 21. McAllister, Don E. 1974. Zoogeography of the marine fishes of Arctic Canada. In Programme de la 54e réunion annuelle de la Société Américaine d’Ichthyologie et de Herpétologie / Program of the 54th Annual Meeting of the American Society of Ichthyologists and Herpetologists, Musée National des Sciences Naturelles / National Museum of Natural Sciences, 17 June to 22 June / du 17 au 22 juin 1974, Ottawa [abstract] p. 22. 22. Legendre, Vianney, Hunter, J.G., & McAllister, Don E. 1975. French, English and scientific names of marine fishes of Arctic Canada / Noms français, anglais et scientifiques des poissons marins de l’Arctique canadien. Syllogeus, 7: 1–15, 1 figure, 1 map. 23. McAllister, Don E. 1975. The ecology of the marine fishes of Arctic Canada. In Proceedings of the Circumpolar Conference on Northern Ecology, September 15–18, 1975, Ottawa [abstract], p. 33. 24. McAllister, Don E. 1975. A new species of Arctic eelpout, Lycodes sagittarius, from the Beaufort Sea, Alaska, and the Kara Sea, USSR (Pisces: Zoarcidae). National Museum of Natural Sciences, National Museums of Canada, Publications in Biological Oceanography, 9: 1–16, 3 figures, 2 tables. 25. McAllister, Don E., & Aniskowicz, Jadwiga. 1976. Vertebral number in North American sculpins of the Myoxocephalus quadricornis-complex. Journal of the Fisheries Research Board of Canada, 33(12): 2792–9, 3 figures, 2 tables. 26. McAllister, D.E. 1977. Ecology of the marine fishes of Arctic Canada. In Proceedings of the Circumpolar Conference on Northern Ecology, September 15–18, 1975, National Research Council of Canada, Ottawa. Section II, Marine Ecology, pp. 49–65; viii + 648 pp, 2 figures. 27. McAllister, D.E., & Andriashev, A.P. 1977. Ophidium parrii Ross, 1826 (Pisces): Application for suppression Z.N.(S.) 1578. Bulletin of Zoological Nomenclature, 34(1): 58–60. 28. Andriashev, A.P., & McAllister, Don E. 1978. Status of the presumed zoarcid fish Ophidium parrii and its identity with Liparis koefoedi, Liparidae. Copeia, 1978(4): 710–12, 1 figure. 29. Able, Kenneth W., & McAllister, Don E. 1981. Revision of the snailfish genus Liparis from Arctic Canada. Canadian Bulletin of Fisheries and Aquatic Sciences, 208(1980): viii + 52 pp., 27 figures, 7 tables.

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30. McAllister, Don E., Anderson, M. Eric, & Hunter, J.G. 1981. Deepwater eelpouts, Zoarcidae, from Arctic Canada and Alaska. Canadian Journal of Fisheries and Aquatic Sciences, 38(7): 821–39, 5 figures. 31. Steigerwald, Michèle Bélanger, & McAllister, Don E. 1982. List of the Canadian marine fish species in the National Museum of Natural Sciences, National Museums of Canada / Liste des espèces de poissons marins du Canada au Musée national des sciences naturelles, Musées nationaux du Canada. Syllogeus, 41: 1–30, 2 figures. 32. McAllister, D.E. 1983. Biogeography of marine fishes of Arctic Canada. In Proceedings of the thirty-sixth meeting of the Canadian Conference for Fisheries Research held at Winnipeg, January 3 and 4, 1983 (pp. 14–16). 33. Hunter, J.G., Leach, Shirley T., McAllister, Don E., & Steigerwald, Michèle Bélanger. 1984. A distributional atlas of records of the marine fishes of Arctic Canada in the National Museums of Canada and Arctic Biological Station. Syllogeus, 52: 1–35, 3 text figures, 21 plates. 34. McAllister, Don E., Parker, Brad J., & McKee, Paul M. 1985. Rare, endangered and extinct fishes in Canada. Syllogeus, 54: 1–192, 60 figures. 35. Renaud, Claude B., Speers, Lawrence I., Qadri, Sami U., & McAllister, Don E. 1986. Biochemical evidence of speciation in the cod genus Gadus. Canadian Journal of Zoology, 64(7): 1563–6, 1 figure, 3 tables. 36. Prouse, N.J., & McAllister, Don E. 1986. The glacial eelpout, Lycodes frigidus, from the Arctic Canadian Basin, new to the Canadian ichthyofauna. Canadian Field-Naturalist, 100(3): 325–9, 2 figures. 37. McAllister, Don E., and Steigerwald, Michèle Bélanger. 1987. Bibliography of the marine fishes of Arctic Canada, 1771–1985. (Canadian Manuscript Report of Fisheries and Aquatic Sciences No. 1909, 1986): v + 108 pp., 2 figures. 38. McAllister, Don E., Legendre, Vianney, & Hunter, J.G. 1987. Liste des noms inuktitut (esquimaux), français, anglais et scientifiques des poissons marins du Canada arctique / List of Inuktitut (Eskimo), French, English and Scientific names of marine fishes of Arctic Canada. (Rapport manuscrit canadien des sciences halieutiques et aquatiques No. 1932): v + 106 pp., 1 figure. 39. McAllister, D.E. 1987. Priorities for ichthyological research in the Canadian Arctic Archipelago. In Program and Abstracts: The Canadian Arctic Islands, Canada’s Missing Dimension, an international meeting sponsored by the National Museum of Natural Sciences, Ottawa, Canada, 21–4 November 1987 [abstract], p. 77.



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40. McAllister, D.E. 1989. Fishes and benthic invertebrates of Hudson Strait. Atelier sur le Detroit d’Hudson Strait Workshop, 9–10 Novembre/November 1989, Holiday Inn Pointe-Claire, Québec. [Abstract], supplementary pages, 36A–36D. 41. McAllister, Don E. 1990. Priorities for ichthyological research in the Canadian Arctic Archipelago. In Harrington, C.R. (Ed.), Canada’s missing dimension: Science and history in the Canadian Arctic islands, vol. 2 (pp. 512–15), 5 figures. Canadian Museum of Nature, Ottawa. 42. McAllister, D.E. 1990. Diversity and biology of invertebrates and fish. In J.A. Percy (Ed.), Compte Rendu d’un Atelier: Études de l’écosytème marin du détroit d’Hudson / Proceedings of workshop: Marine ecosystem studies in Hudson Strait, 9–10 November, 1989, Montréal, Québec (pp. 120–6). (Canadian Technical Report of Fisheries and Aquatic Sciences No. 1770): xiv + 175 pp. 43. McAllister, Don E. 1990. A list of the fishes of Canada / Liste des poissons du Canada. Syllogeus, 64: 1–310, 32 figures. 44. Houston, J., & McAllister, D.E. 1990. Status of the Bering wolffish, Anarhichas orientalis, in Canada. Canadian Field-Naturalist, 104(1): 20–3, 3 figures. 45. Houston, J., & McAllister, D.E. 1990. Status of the Blackline Prickleback, Acantholumpenus mackayi, in Canada. Canadian Field-Naturalist, 104(1): 24–8, 3 figures. 46. McAllister, Don E. 1991. Questions on ocean impacts of James and Hudson Bay hydro projects. Sea Wind, Bulletin of Ocean Voice International, Ottawa, 5(3): 22–30, 2 figures. 47. McAllister, Don E. 1991. Measuring Arctic Ocean icepack shrinkage and warming: The planetary solar mirror and global thermostat. Sea Wind, Bulletin of Ocean Voice International, Ottawa, 5(4): 19–24, 1 figure. 48. Alfonso, Noel, & McAllister, Don E. 1994. Biodiversity and the Great Whale Hydroelectric Project / La biodiversité et le projet hydroélectrique Grande-Baleine. (Great Whale Environmental Assessment, Background Paper No. 11): vii + 75 pp. Great Whale Public Review Support Office, Montréal. / (Dossier-synthèse no. 11, Évaluation environnementale du projet Grande-Baleine): vii + 81 pp., Bureau de soutien de l’examen public du projet Grande-Baleine, Montréal. 49. Coad, Brian W., Reist, James D., Renaud, Claude B., McAllister, Don E., Møller, Peter Rask, Alfonso, Noel, & Berkes, Fikret. 2001. A field guide to the Arctic marine fishes of Canada. Poster presented at Arctic Ocean Sciences Board, twentieth meeting (AOSB-XX), 23–4 April 2001, Iqaluit.

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ACKNOWLEDGMENTS Brian W. Coad and James D. Reist

A work of this nature depends heavily on the contributions of many people over a long period of time. Scientists, naturalists, and consultants sent specimens of fishes from Arctic waters to the Canadian Museum of Nature (CMN) for identification and for deposit in the collections. Reprints of scientific papers, books, and consultant reports were also freely given. These materials were built up over 130 years, most actively during the past 60 years, and form one of the main bases of this work. Many were based upon field expeditions to the Canadian Arctic during this time frame (see the “History” section in the introduction), often under rigorous conditions. It is not possible to list all the contributors here, but we hope that Marine Fishes of Arctic Canada is a worthy acknowledgment of their efforts. The writing phase of this book received help from a number of people and institutions that need to be thanked specifically. The main external funding agencies were the Nunavut Wildlife Management Board (Jim Noble and Dan Pike, both now retired), and the Inuvialuit Fisheries Joint Management Committee (Bob Bell, now retired), which are gratefully acknowledged here. Brian Coad, Claude Renaud, and Noel Alfonso were supported by the CMN; James Reist, Chantelle D. Sawatzky, J. Brian Dempson, and Les N. Harris by Fisheries and Oceans Canada (DFO), Winnipeg; Karen Dunmall, University of Manitoba and DFO; Michael Power, University of Waterloo, Ontario; and Heidi K. Swanson, University of Alberta, Edmonton; H. Powles and Andries Blouw (both now retired), who aided in early development and funding of the book by DFO; Peter Møller by the Zoological Museum, University of Copenhagen; and Fikret Berkes by the University of Manitoba, Winnipeg. These institutions contributed salary and overhead to the project for extended periods. The maintenance, correction, and large amounts of input to the database on fish distributions, as well as the production of maps, were carried out by Noel R. Alfonso, CMN. Teams that input mapping data and located copies of scientific articles comprised principally Naomi Keyzer-de Ville, Diane Pathy, Melanie Gaudet, Stuart Lithwick, and Nadia Arbach at the CMN; and Neil J. Mochnacz and Megan Puchniak at the Arctic Fish Ecology and Assessment Research Section, DFO, Winnipeg.

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Mark Graham is thanked for access to his slide collection and particularly for his support as Director of the Research Services Division and later as Vice-President of Research and Collections, CMN, during the long course of this project, latterly assisted by Wendy McPeake. Staff at University of Toronto Press helped bring this book to fruition: executive editor Suzanne Rancourt, managing editor Leah Connor, copy editor Angela Wingfield, and production manager Ani Deyirmenjian. Thanks also to Peter Ross and Linda Gustafson at Counterpunch for their work on the design of this book. Artists and illustrators are listed in “Illustration Credits,” and also include Shannon MacPhee, Julie Henry, Sheila Atchison, Chantelle Sawatzky, and Michell Kamula, all DFO, for introductory chapters. Illustrations in the section “Arctic Fish, Northern Cultures, and Traditional Ecological Knowledge” in the introduction were obtained through the efforts of Erin Gurski and Mélissa Duncan, Canadian Museum of History, Gatineau. We should also like to thank the following for their contributions, which varied from small, but crucial, items to large and expansive datasets. The Canadian Museum of Nature: Mike Wayne and Anne-Marie Barter at the library for assistance in libraries and for access to documents on native foods; Sylvie Laframboise for collections assistance and patient verification of records; Alison Murray (formerly of CMN) for checking collections held by the Redpath Museum, Montréal; Roger Bull for identification of a shark specimen through bar-coding; A. Thérien for stitching assistance; and Mélanie Gaudet-Leblanc and Daniel Nadon (formerly of CMN) for technical support. Fisheries and Oceans Canada: Andries Blouw (now retired) for early aid in developing administrative linkages to support the publication; Brian Dempson for information on indigenous fish names; Chantelle Sawatzky, Darcy McNicholl, Margaret Treble, and Sheila Atchison for mapping, literature, database aid, and collation of photographs; Margaret Treble, Kevin Hedges, Tim Siferd, Andy Majewski, Shannon MacPhee, and Sheila Atchison for provision of occurrence data, specimens (such as Apristurus, Platytroctes apus, and Rouleina maderensis from Siferd), and related fieldwork

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in support; Claude Nozères for providing access and permission to use specimen photographs; David Orr, Newfoundland, for information on cruise data; Roberta Miller and Luci Bossé, Institut Maurice-Lamontagne, Mont-Joli, for distributional records; Doug Swain, Moncton, for information on the post-moratorium total allowable catch set for Atlantic Cod; and the staff of the DFO libraries. Chantelle Sawatzky spent considerable time and effort in the final stages of the manuscript, tying up loose ends, and thus contributed significantly to the overall outcome. P. Bartsch and C. Lamour, Museum für Naturkunde, Humboldt-Universität zu Berlin, for access to specimens; Barbara Brown and Scott Schaefer, American Museum of Natural History, New York, for distributional records; Dylan Radcliffe for his invaluable contribution to GIS programming that greatly facilitated map production; Patrick Campbell, Oliver Crimmen, and James Maclaine, Natural History Museum, London, for access to collections and distributional records; Sylvie Coad for assistance in libraries and for access to documents on native foods; Robert G. Fechhelm, LGL Ecological Research Associates Inc., Bryan, Texas, for maps of anadromous fish distributions; Jon Fong, California Academy of Sciences, San Francisco, for distributional records; Nora R. Foster, University of Alaska, Fairbanks, for checking holdings of Arctic fishes; John Gilhen, Nova Scotia Museum of Natural History, Halifax, for distributional records; Lou van Guelpen, Huntsman Marine Laboratory, St Andrews, New Brunswick, for access to records and specimens; Jack Hall, Ottawa, for photographing certain species to aid in illustrations; Dave Hardie and Jeff Hutchings, Dalhousie University, Halifax, for information on Arctic lake populations of Atlantic Cod; Stephen Hendrie, Communications Officer, Makivik Corporation, Kuujjuaq, for copies of publications; Erling Holm and Mary Burridge, Royal Ontario Museum, Toronto, for help in untangling collection localities and for distributional records and loans of materials; Haakon Hop, Norwegian Polar Institute, Tromsø, for assistance in contacting Peter Leopold for use of the Arctic Cod photograph; Christiane Hudon, Environment Canada, Montréal, for copies of papers and extensive databases that saved us much work; Tomio Iwamoto, California Academy of Sciences,



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San Francisco, for constructive comments on the macrourid key; Steen Knudsen for the Careproctus kidoi drawing; Sven Kullander, Swedish Museum of Natural History, Stockholm, for data on marine fish distributions from Tundra Northwest 99; Jasmine Lee, Ottawa, for bibliographic assistance; Harald Loeng, Institute of Marine Research, Bergen, for information on Russian work in the Arctic; J. Andrés López, University of Alaska Museum, for Beaufort Sea records; Catherine W. Mecklenburg, Point Stephens Research, Auke Bay, Alaska, for information exchanges on Arctic fishes, their systematics, and their distribution; W. Brian McKillop, Manitoba Museum, for distributional records; Suzanne Monette, formerly University of Ottawa, for providing technical support; J. Neilson, MacLaren Atlantic Limited, Dartmouth, Nova Scotia, for results of fisheries surveys; Jørgen Nielsen, Zoological Museum, University of Copenhagen, for access to collections; Ted Pietsch, University of Washington, Seattle, for confirming identification of Chaenophryne longiceps and for explaining the trivial name gladisfenae; Geoff Power, University of Waterloo, Ontario, for copies of articles; Wayne E. Roberts, University of Alberta Museum of Zoology, Edmonton, for distributional records; staff of the libraries of the Geological Survey of Canada (Ottawa), Environment Canada, and the Department of Indian Affairs and Northern Development (Gatineau); Duane Stevenson, National Oceanic and Atmospheric Administration (NOAA), for information on the taxonomy of Manefishes; Eric Taylor and Megan McCusker, University of British Columbia, Vancouver, for access to distributional records; Peter Troffe, Royal British Columbia Museum, Victoria, for checking holdings of Arctic fishes; Richard Verdon, Hydro-Québec, Montréal, for a database on collections from James Bay; Ken Vogt (deceased), University of Alaska, Anchorage, for information on the exploitation of Arctic Lamprey; Jim Whittome, University of Alberta, Edmonton, for fish distribution records; and Bruce Wing, Auke Bay Laboratory, Alaska, for checking holdings of Arctic fishes. We also thank the two scientific reviewers for their diligence in examining the entire manuscript, noting corrections needed, providing useful comments, and pointing out inconsistencies.

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Introduction

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PURPOSE OF THE BOOK Brian W. Coad

Marine Fishes of Arctic Canada provides a guide to the marine fishes found in Arctic Canadian waters. Accurate identifications are essential to any study. Some of the species described here are of actual or potential commercial value. Many are critical species in the ecology of northern waters. The book includes keys for identification; illustrations of the species and their key characters; distribution maps; notes on taxonomy; scientific and common names; descriptions of the species; habitat data; biology; importance; and traditional knowledge. The geographic area complements the areas covered by previous books on the fishes of adjacent waters, notably W.B. Scott and M.G. Scott (1988) on Atlantic Canada and Mecklenburg, Mecklenburg, and Thorsteinson (2002) on Alaskan fishes. The

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Alaska-Canada border is a political termination of the area of coverage and not a natural one for fishes. Various species that are now known only from northern Arctic Alaska may eventually be found in Canadian waters. The border between Arctic and Atlantic fishes is a literary one, as Scott and Scott (1988) cover fishes to the tip of Labrador. Arctic waters extend down the Labrador coast, but the inclusion of this area would add significant diversity in southern species at their northern limit, with which their book deals. Many species from these far northern waters are poorly known, both taxonomically and biologically, and much remains to be learned or elucidated. This book is the first to appear on such fishes in Canada, and it is hoped that it will lead to and facilitate further work.

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BIODIVERSITY Brian W. Coad and Noel R. Alfonso

The marine Arctic ichthyofauna of Canada comprises 58 families, 138 genera, and 221 species (see table below). This diversity is expected to increase as further surveys are carried out in deeper waters, as ice-free areas allow more access, and as climate change facilitates dispersal. For example, McAllister (1960b) listed 105 species for Arctic Canada, and Coad and Reist (2004) listed 189 species. The whole Canadian ichthyofauna, both marine and freshwater, comprises 233 families and 1,439 species, so the Arctic has about 25% of the families and about 15% of the species. Mecklenburg, Møller, and Steinke (2011), Christiansen, Reist, et al. (2013), and Lynghammar et al. (2013) review the taxonomy and zoogeography of all Arctic fishes. Fishes occurring in marine waters in the Arctic are either wholly marine (i.e., spend all of their life associated with marine systems, although some may venture into freshened estuaries) or exhibit anadromy (i.e., spend periods of their life history in marine systems, with the remaining periods spent in fresh waters). The vast majority of the 221 species here are wholly marine. However, species in six families – Petromyzontidae (1 species), Acipenseridae (1 species), Osmeridae (2 of 3 species), Salmonidae (17 species), Gadidae (1 of 8 species), Gasterosteidae (2 species) – exhibit anadromy (see details in the “Family and Species Accounts” section). The “Checklists of Species” section includes a list of extralimital species that may eventually be found in Canadian waters (summarized in the table below). There are 23 more families, 65 more genera, and 121 more species that may eventually be recorded, thereby increasing the species diversity by more than half (55%). The southern Davis Strait and Labrador Sea, opening onto the Atlantic Ocean, would be the source of 84% of these species, including many deepwater species and some expatriates. The remainder would be Alaskan species reaching the Beaufort Sea. The marine Arctic fauna consists of 20 jawless and cartilaginous fishes (Hagfishes and Lampreys, chimaeras, sharks, Skates, Sturgeons) and 202 Ray-finned Fishes (all others). The most speciose family is the Zoarcidae with 31 species, but 27 families are represented by only a single species. Most families have less than 10 species, but the Salmonidae have 17 species; Cottidae, 14 species; and Liparidae, 11 species.



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The commonest and most numerous species in terms of individuals is probably Boreogadus saida, a keystone species in the ecology of the Arctic on which many other fishes, birds, and marine mammals depend as a food source. One of the rarest fish is Bythites fuscus, first collected as a single specimen in 1834 off Greenland, and the second and third specimens known to science only being caught in the year 2000 off Baffin Island in Canadian waters. Many of the species are known only to scientists and come from the deeper waters of the Davis Strait, whereas others have been important food for Indigenous peoples for millennia. The northernmost species is Lycodes frigidus from northwest of Ellesmere Island at 85º48'27" N, 110º43'39" W. Other northern species include Salvelinus alpinus, Arctogadus glacialis, Boreogadus saida, Myoxocephalus quadricornis, and Reinhardtius hippoglossoides, all found at the northern tip of Ellesmere Island (83º N). Species found only in the eastern Arctic (east of the Boothia Peninsula) number 136 species (61.5% of the Canadian Arctic fauna), those only in the western Arctic (west of the Boothia Peninsula) number 30 species (13.6%), and those found in both areas number 54 species (24.4%). No species are endemic to Arctic Canada, although such species as Bythites fuscus and Lycodes mcallisteri have distributions restricted to the Arctic waters of Canada and immediately adjacent Greenland waters. The only non-indigenous species are Osmerus mordax in Hudson Bay, which spread from populations that were introduced to the Great Lakes and became established, and Oncorhynchus gorbuscha and O. keta that were introduced to Hudson Bay but are apparently not surviving. Thirty-seven new records for the Canadian Arctic that were not given in the annotated list by Coad and Reist (2004) are dealt with in this book: Harriotta raleighana, Rhinochimaera atlantica, Apristurus profundorum, Centroscymnus coelolepis, Somniosus pacificus, Rajella bathyphila, Acipenser fulvescens, Simenchelys parasitica, Holtbyrnia anomala, Maulisia mauli, Maulisia microlepis, Normichthys operosus, Platytroctes apus, Bathytroctes sp., Rouleina maderensis, Osmerus dentex, Salvelinus namaycush, Argyropelecus gigas, Myctophum punctatum, Gadomus longifilis, Enchelyopus

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cimbrius, Brosme brosme, Micromesistius poutassou, Lophius americanus, Himantolophus groenlandicus, Ceratias holboelli, Gigantactis vanhoeffeni, Scopeloberyx robustus, Hoplostethus atlanticus, Sebastes fasciatus, Psychrolutes phrictus, Cyclopteropsis mcalpini, Careproctus kidoi, Caristius fasciatus, Lycodes squamiventer, Ulvaria subbifurcata, and Chiasmodon harteli. One species, Lycodes rossi Malmgren, 1865, was deleted from the list. Three species are re-identified taxa (Holtbyrnia anomala was Holtbyrnia sp., Caristius fasciatus was Caristius sp., and Chiasmodon harteli was C. niger), one subspecies was recognized as a species (Osmerus dentex), one species was described as new subsequent to Coad and Reist (2004) (Careproctus kidoi), one species (Salvelinus namaycush) is a taxon that was previously thought to be confined to fresh waters, and one taxon that is mostly found in fresh water but enters marine areas (Acipenser fulvescens) was added. There has been no confirmation of Coregonus laurettae Bean, 1881, in Arctic marine waters of Canada, and it has been deleted from the list. Two records, Maulisia mauli and Gadomus longifilis, are new to Canada. Threatened and endangered species are listed according to the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) and on the Red List of Threatened Species issued by the International Union for Conservation of Nature (IUCN). These online sites were accessed in December 2013 and should be consulted for criteria and updated information. The status levels, from least to most endangered, identified by COSEWIC are “Not at Risk,” “Data Deficient,” “Special Concern,” “Threatened,” “Endangered,” “Extirpated,” and “Extinct.” On the IUCN Red List they are “Data Deficient,” “Lower Risk / Least Concern,” “Least Concern,” “Near Threatened,” “Vulnerable,” “Endangered,” “Critically Endangered,” “Extinct in the Wild,” and “Extinct.” COSEWIC assessments are for Canadian Arctic populations and are noted if they are for a specific Arctic population. These assessments also include species as a whole in Canada, and the Arctic distribution is at the margin of the main range, for example, Atlantic Ocean Brosme brosme. The IUCN Red List is for the species worldwide. COSEWIC (species and populations counted separately) lists “Not at Risk” (2), “Data Deficient” (6), “Special Concern” (6), “Threatened” (4), and “Endangered” (3), for a total of 20 species or populations assessed. IUCN indicates “Data Deficient” (4), “Lower Risk / Least Concern” (2), “Least Concern” (33), “Near Threatened” (3), “Vulnerable” (2), and “Endangered” (2), for a total of 46 species assessed. The remaining species await assessment.

COSEWIC has assessed the status levels of the following species: not at risk:Myoxocephalus quadricornis (saltwater form; Cottidae), Hippoglossus hippoglossus (Pleuronectidae). data deficient:Salmo salar (Nunavik population; Salmonidae), Gadus morhua (Arctic Marine population; Gadidae), Myoxocephalus quadricornis (freshwater form; Cottidae), Acantholumpenus mackayi

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(Stichaeidae), Anarhichas orientalis (Anarhichadidae), Hippoglossoides platessoides (Arctic population; Pleuronectidae).

special concern:Amblyraja radiata (Rajidae), Acipenser ful-

vescens (freshwater Southern Hudson Bay–James Bay populations; Acipenseridae), Salvelinus malma malma (freshwater Western Arctic populations; Salmonidae), Macrourus berglax (Atlantic Ocean; Macrouridae), Gadus morhua (Arctic lakes population; Gadidae), Anarhichas lupus (Anarhichadidae).

threatened: Sebastes fasciatus (Atlantic population), Sebastes

mentella (Northern population; Scorpaenidae), Anarhichas denticulatus, Anarhichas minor (Anarhichadidae).

endangered: Acipenser fulvescens (freshwater Western Hudson Bay populations; Acipenseridae), Coryphaenoides rupestris (Macrouridae), Brosme brosme (Atlantic Ocean; Gadidae).

IUCN has assessed the following species: data deficient:Apristurus profundorum (Scyliorhinidae), Somniosus pacificus (Somniosidae), Lepidion eques (Moridae), Paraliparis copei (Liparidae). lower risk / least concern:Coregonus artedi (Salmoni-

dae), Salmo salar (Sal­monidae).

least concern:Myxine glutinosa (Myxinidae), Lethenteron

camtschaticum (Petromyzontidae), Harriotta raleighana (Rhinochimaeridae), Rhinochimaera atlantica (Rhinochimaeridae), Hydrolagus affinis (Chimaeridae), Centroscyllium fabricii (Etmopteridae), Amblyraja hyperborea (Rajidae), Amblyraja jenseni (Rajidae), Malacoraja spinacidermis (Rajidae), Rajella bathyphila (Rajidae), Rajella fyllae (Rajidae), Rajella lintea (Rajidae), Acipenser fulvescens (Acipenseridae), Simenchelys parasitica (Synaphobranchidae), Synaphobranchus kaupii (Synaphobranchidae), Clupea harengus (Clupeidae), Osmerus dentex (Osmeridae), Osmerus mordax (Osmeridae), Coregonus autumnalis (Salmonidae), Coregonus nasus (Salmonidae), Coregonus sardinella (Salmonidae), Oncorhynchus nerka (Salmonidae), Salvelinus alpinus (Salmonidae), Stenodus leucichthys (Salmonidae), Lota lota (Gadidae), Chaenophryne longiceps (Oneirodidae), Gasterosteus aculeatus (Gasterosteidae), Pungitius pungitius (Gasterosteidae), Sebastes mentella (Scorpaenidae), Myoxocephalus quadricornis (Cottidae), Lycodes jugoricus (Zoarcidae), Lycodes paamiuti (Zoarcidae), Platichthys stellatus (Pleuronectidae).

near threatened:Centroscymnus coelolepis (Somniosidae), Somniosus microcephalus (Somniosidae), Bathyraja spinicauda (Rajidae). vulnerable:Amblyraja radiata (Rajidae), Gadus morhua (Gadidae).

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endangered: Sebastes fasciatus (Scorpaenidae), Hippoglossus hippoglossus (Pleuronectidae).

Rarity is based on mapping records that reflect accessibility and economic importance (very rare = 1–2 records; rare = 3–10 records; uncommon = 11–49 records; common 50–499 records; abundant = 500+ records; very abundant = 2,000+ records). Small, deep-water species, perhaps abundant under the ice pack, are seldom collected, while surveys for commercial species, such as Reinhardtius hippoglossoides, have numerous records. The majority of the very rare to rare species are deep-sea taxa or expatriates or are at their northern range limit, and the abundant to very abundant species are found in shallow water, are economically important, or are genuinely abundant. Very rare species number 43 (19.5%), rare 31 (14.0%), uncommon 32 (14.5%), common 77 (34.8%), abundant 32 (14.5%), and very abundant 6 (2.7%). The very abundant species are Salvelinus alpinus, Macrourus berglax, Boreogadus saida, Myoxocephalus quadricornis, Liparis fabricii, and Reinhardtius hippoglossoides. Species by provinces and territories may be summarized as follows. Some provinces have a limited Arctic coastline, for example, Yukon, Manitoba, and Ontario, whereas Nunavut extends across most of Arctic Canada. This disparity naturally affects the diversity for each province and territory; Manitoba and Ontario, for example, front on the Hudson and James Bays, which are species poor. Nunavut has 207 species (93.7% of the Canadian Arctic fauna); Québec, 100 species (45.2%); Northwest Territories, 80 species (36.2%); Yukon, 50 species (22.6%); Manitoba, 29 species (13.1%); and Ontario, 24 species (10.9%). Ninety-six species (43.4%) are found in one province or territory, 58 species (26.2%) in two, 30 species (13.6%) in three, 13 species (5.9%) in four, 10 species (4.5%) in five, and 14 species (6.3%) in six. Species found in all six provinces and territories are Mallotus villosus, Coregonus artedi, C. clupeaformis, Prosopium cylindraceum, Salvelinus namaycush, Boreogadus saida, Lota lota, Pungitius pungitius, Myoxocephalus quadricornis, M. scorpioides, M. scorpius, Aspidophoroides olrikii, Eumicrotremus derjugini, and Ammodytes hexapterus. These species inhabit shallow or estuarine waters and thus are easily caught or rank as very abundant to abundant. Ninety-one species (41.2%) are found only in Nunavut, and the majority are deep-sea species of Davis Strait and Baffin Bay. In addition to new records, it is evident that continuing fieldwork is expanding the knowledge on distribution both in terms of major water bodies and of ecoregions, as compared with Coad and Reist (2004). Distribution by area reflects major water bodies combined



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with accessibility and survey frequency. The High Arctic islands area is defined by being north of, or west of, the other areas. Some species only extend to the southern fringe of the High Arctic, for example, Coregonus clupeaformis, while others are widely distributed there, for example, Boreogadus saida. Baffin Bay and Davis Strait includes Nares Strait as well as Frobisher Bay and Cumberland Sound. Waters between the High Arctic islands have been poorly surveyed, but surveys associated with commercial fisheries in Davis Strait and recent deep-water surveys in the Beaufort Sea have increased knowledge of biodiversity. New records for Arctic Canada are still being made (see the following “New Information” section in the introduction, and the “Extralimital Species” section under “Checklists of Species”). Baffin Bay and Davis Strait have 175 species (79.2% of the Canadian Arctic fauna); Hudson Strait and Ungava Bay, 107 species (48.4%); Hudson and James Bays and the Foxe Basin, 55 species (24.9%); Queen Maud and Coronation gulfs, 60 species (27.1%); Beaufort Sea and Amundsen Gulf, 76 species (34.4%); and the High Arctic islands, 50 species (22.6%). Ninety-nine species (44.8%) are found in only one area, 56 species (25.3%) in two, 20 species (9.0%) in three, 6 species (2.7%) in four, 12 species (5.4%) in five, and 28 species (12.7%) in six. Eighty species are found only in Baffin Bay and Davis Strait, the majority being deep-sea species as noted above. Distribution by ecoregion reflects fish habitats and is based on Coad and Reist (2004) with updates (see map in the “Environment” section of the introduction). The James Bay–eastern Hudson Bay ecoregion has 42 species (19.0% of the Canadian Arctic fauna); Hudson Bay, 44 species (19.9%); Foxe Basin, 20 species (9.0%); Hudson Strait, 103 species (46.6%); Labrador Sea, 140 species (63.3%); Baffin Bay–Davis Strait Nearshore, 93 species (42.1%); Baffin Bay–Davis Strait Offshore, 119 species (53.8%); Lancaster Sound Region, 46 species (20.8%); High Arctic Archipelago, 25 species (11.3%); Arctic Basin, 11 species (5.0%); Viscount Melville Sound, 21 species (9.5%); Queen Maud Gulf, 60 species (27.1%); and Beaufort Sea–Amundsen Gulf, 76 species (34.4%). Sixty-one species (27.6%) are found in only one ecoregion, 53 species (24.0%) in two, 23 species (10.4%) in three, 27 species (12.2%) in four, 9 species (4.1%) in five, 11 species (5.0%) in six, 7 species (3.2%) in seven, 8 species (3.6%) in eight, 4 species (1.8%) in nine, 8 species (3.6%) in ten, 5 species (2.3%) in eleven, 3 species (1.4%) in twelve, and 2 species (0.9%) in thirteen. The three species in twelve ecoregions and the two species in thirteen ecoregions are Salvelinus alpinus, Gymnocanthus tricuspis, and Triglops pingelii; and Boreogadus saida and Liparis tunicatus, respectively.

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Biodiversity of Arctic Marine Fishes (Percentages for extralimital species are not given.) Family

Genera in the Canadian Arctic

Species in the Canadian Arctic

Extralimitals (Genera + Species)

Number of Species (and Arctic %) in Canadian Fauna (after Coad, 2014)

Number of Species (and Arctic %) in World Fauna (after Nelson, 2006)

Myxinidae Petromyzontidae Rhinochimaeridae Chimaeridae Cetorhinidae Lamnidae Scyliorhinidae Squalidae Etmopteridae Somniosidae Rajidae Acipenseridae Halosauridae Notacanthidae Anguillidae Synaphobranchidae Nemichthyidae Serrivomeridae Saccopharyngidae Eurypharyngidae Clupeidae Argentinidae Microstomatidae Platytroctidae Alepocephalidae Osmeridae Salmonidae Gonostomatidae Sternoptychidae Phosichthyidae Stomiidae Synodontidae Notosudidae Alepisauridae Paralepididae Myctophidae Lampridae Trachipteridae Macrouridae Moridae Phycidae Gadidae Bythitidae Lophiidae Caulophrynidae

1 1 2 1

1 1 2 1

1

1

1 2 4 1

1 3 9 1

0+1 1+1 0+1 0+1 1+1 1+1 0+1 1+1 1+1

2

2

2 1 1 1 1 1 1 1 4 5 2 6 2 1

2 1 1 1 1 2 1 1 5 6 3 17 2 1

3 (33.3%) 11 (9.1%) 3 (66.7%) 2 (50.0%) 1 5 8 (12.5%) 2 3 (33.3%) 3 (100.0%) 28 (32.1%) 5 (20.0%) 6 5 (40.0%) 2 4 (50.0%) 4 (25.0%) 3 (33.3%) 1 (100.0%) 1 (100.0%) 10 (20.0%) 3 (33.3%) 4 (25.0%) 13 (38.5%) 17 (35.3%) 11 (27.3%) 42 (40.5%) 17 (11.8%) 12 (8.3%)

6

6

1

1

26 (34.6%) 11 (27.3%) 4 (100.0%) 13 (61.5%) 4 (25.0%) 1 (100.0%) 1

70 (1.4%) 34 (2.9%) 8 (25.0%) 22 (4.5%) 1 5 138 (0.7%) 16 44 (2.3%) 17 (17.6%) 238 (3.8%) 25 (4.0%) 15 10 (20.0%) 15 32 (6.3%) 9 (11.1%) 10 (10.0%) 10 (10.0%) 1 (100%) 188 (1.1%) 23 (4.3%) 38 (2.6%) 37 (13.5%) 90 (6.7%) 31 (9.7%) 66 (25.8%) 23 (8.7%) 67 (1.5%) 20 273 (2.2%) 57 19 (5.3%) 3 56 (7.1%) 240 (3.3%) 2 10 350 (2.6%) 105 (2.9%) 25 (16.0%) 31 (25.8%) 107 (0.9%) 25 (4.0%) 5

2 3 (33.3%) 18 (16.7%)

5 18 (5.6%) 62 (4.8%)

Melanocetidae Himantolophidae Oneirodidae

6

0+1 1+1 1+1 1+1 1+1

2+4 2+3 3+4 0+2 0+1 3+7 1+1 2+2 1+1

64 (9.4%) 6 (16.7%)

1+2 4 6

4 8

5 3 3 7 1 1

9 3 4 8 1 1

1 3

1 3

2+2 1+1 1+1 0+2 1+3 5+8 0+1 1+1 1+2 4+8

16 (25.0%) 83 (9.6%)

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Family

Genera in the Canadian Arctic

Species in the Canadian Arctic

Extralimitals (Genera + Species)

Number of Species (and Arctic %) in Canadian Fauna (after Coad, 2014)

Number of Species (and Arctic %) in World Fauna (after Nelson, 2006)

Ceratiidae Gigantactinidae Linophrynidae Melamphaidae Rondeletiidae Barbourisiidae Cetomimidae Anoplogastridae Diretmidae Trachichthyidae Oreosomatidae Gasterosteidae Syngnathidae Scorpaenidae Hexagrammidae Cottidae Hemitripteridae Agonidae Psychrolutidae Cyclopteridae Liparidae Caristiidae Zoarcidae Stichaeidae Pholidae Anarhichadidae Nototheniidae Chiasmodontidae Ammodytidae Trichiuridae Stromateidae Scombridae Pleuronectidae Total 58 (23)

1 1

1 1

1+1

1

1

1

1

1

1

2

2

1

3

5

14

2 2 3 4 1 6 8 1 1

3 3 5 11 1 31 9 1 4

1 1 1 1

1 2 1 1

7 138

9 221

3 (33.3%) 3 (33.3%) 6 18 (5.6%) 1 1 3 1 (100.0%) 1 4 (25.0%) 4 5 (40.0%) 7 51 (5.9%) 8 61 (23.0%) 6 21 (14.3%) 9 (33.3%) 8 (62.5%) 51 (21.6%) 4 (25.0%) 62 (50.0%) 23 (39.1%) 8 (12.5%) 5 (80.0%) 0 5 (20.0%) 3 (66.7%) 10 (10.0%) 2 (50.0%) 15 31 (29.0%)

4 (25.0%) 22 (4.5%) 23 36 (2.8%) 2 1 35 2 (50.0%) 4 39 (2.6%) 10 8 (25.0%) 232 418 (0.7%) 12 275 (5.1%) 8 47 (6.4%) 35 (8.6%) 28 (17.9%) 334 (3.3%) 5 (20.0%) 230 (13.5%) 76 (11.8%) 15 (6.7%) 5 (80.0%) 50 15 (6.7%) 23 (8.7%) 39 (2.6%) 15 (6.7%) 51 60 (15.0%)

2+5 3+4 1+1 1+1 1+1 1+1 1+1 1+1 1+2 1+1 2+5 1+1 1+1 0+1 1+9 2+2 0+4 1+1 0+1 1+1 0+1



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NEW INFORMATION Brian W. Coad

Ongoing surveys in deeper waters continue to uncover species new to Arctic Canada and to extend distributions. In addition, taxonomic research remains active, and the understanding of species changes constantly. Some of these works and records arrived too late for incorporation into the main text of this book and are briefly summarized here. The principal areas of new collections are the deeper waters of the Beaufort Sea under the aegis of the Marine Fishes Project of the Beaufort Regional Environmental Assessment (BREA, 2011–15, www.beaufortrea.ca/), and the ongoing Baffin Bay and Davis Strait Multi-Species Stock Assessment Surveys in Northwest Atlantic Fisheries Organization Divisions 0A, 0B, and SFA3, all conducted by Fisheries and Oceans Canada, Central and Arctic Region. A presentation from BREA (www.beaufortrea.ca/wp-content/​ uploads/2013/03/8.1-A-Majewski-Offshore-Fish-Populations. pdf) mentions the Arrowtooth Flounder / plie à grande bouche – Atheresthes stomias (Jordan and Gilbert, 1880). Mecklenburg et al. (2002) do not record it north of the Bering Sea and consider Chukchi Sea records to be misidentifications. Confirmation of species’ identities from these recent efforts is underway. Specimens have been deposited in the University of Alaska Museum, Fairbanks, from the United States–Canada transboundary area but were not examined for this book. The records include species new to the Canadian Arctic in the Beaufort Sea, namely Liparis bathyarcticus Parr, 1931 (Nebulous Snailfish, limace nébuleuse; previously confused with L. gibbus and known from Arctic Alaska), Lycenchelys micropora Andriashev, 1955 (Manytoothed Eelpout; previously nearest known record from the northern Bering Sea), Lycodes raridens Taranetz and Andriashev, 1937 (Marbled Eelpout; known from Arctic Alaska), and Lycodes sagittarius (known from Arctic Alaska).

8

Orr, Kai, and Nakabo (2015) describe a new species of liparid, the Dusty Snailfish, Liparis lerikimae, from the Alaskan and Canadian Beaufort Sea at 175–500 m. Mecklenburg and Anderson (2015) synonymize Gymnelus knipowitschi Chernova, 1999, with G. hemifasciatus Andriashev, 1937, and G. barsukovi and G. bilabrus with G. viridis, based on fresh material, morphology, and DNA bar-codes. Mecklenburg and Steinke (2015) summarize the ichthyofauna in the Pacific Arctic region (which includes the Beaufort Sea) and include Arctic-wide maps and discussions on taxonomy relevant to Canadian waters. They review certain taxa from the Pacific Arctic and use DNA bar-coding to reveal hidden diversity, for example, Mallotus catervarius across Arctic Canada and Mallotus sp. in Hudson Bay and Atlantic Canada; generic changes, for example, Pleuronectes glacialis to be in the genus Liopsetta; and probable synonymies, for example, Hippoglossoides robustus in H. elassodon. Mecklenburg, Mecklenburg, Sheiko, and Steinke (2016) give an atlas and guide presenting the results of the Russian-American Long-Term Census of the Arctic (RUSALCA), primarily for the Arctic region north of Bering Strait. However, distribution maps, descriptions, taxonomy, and habitats cover the whole Arctic. About two-thirds of the species discussed are found in the Canadian Arctic. Nielsen, Hedeholm, et al. (2016) showed that the Greenland Shark (Somniosus microcephalus) is the longest-lived vertebrate, the largest fish examined at 502 cm being 392 years plus or minus 120 years old.

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ENVIRONMENT James D. Reist and Chantelle D. Sawatzky

The Global Context Canadian Arctic marine waters can be differentiated into those associated directly with the Arctic Ocean (e.g., Beaufort Sea, archipelago areas) and those further south that are sub-Arctic in nature and which tend to be “downstream” of the Arctic Ocean proper (i.e., the Hudson Bay complex including Hudson Strait, Ungava, Hudson and James Bays, and Foxe Basin; Baffin Bay; and Davis Strait). Both the geographical position and the latitudinal extent of these various areas of northern Canada also result in varying degrees of “arcticity” that are elaborated below. Important underlying aspects include connectivity both among these various Canadian marine systems and between them and the global marine context. Accordingly, a wider perspective is developed below. The Arctic Ocean and its adjacent seas are almost completely enclosed by land masses of the northern hemisphere (fig. 1). The Arctic Ocean is the smallest of the world’s oceans at about 14 million square kilometers in area and has a number of bordering seas along the coasts of the surrounding land masses. Also, it is among the shallowest of the world’s oceans (the average depth is 1,038 m; the deepest point is 5,450 m). Unlike other oceans of the world, the Arctic Ocean is dominated by marine inputs from the Atlantic and Pacific oceans, proportionately larger inputs of fresh water from numerous large Arctic rivers, cold atmospheric temperatures within a narrow range throughout most of the year, and ice that both reflects and absorbs great amounts of thermal and light energy. Most of the area is also characterized in winter by continuous darkness and in summer by continuous daylight. These features influence the nature and productivity of Arctic marine environments, particularly in the Canadian portion of the Arctic Ocean. This, in turn, influences the types, distributions, and abundance of Canadian Arctic marine fishes both here and in the areas downstream of the Arctic Ocean proper. The Arctic Ocean is generally characterized by wide continental shelves (0 m to about 200+ m depth), especially along the Eurasian coast, that rapidly drop off (200 m to about 500 m depth) to deep basins (about 500+ m depth). The epicontinental seas of the Arctic



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Ocean comprise about 37% of its area, compared with about 8% as the average for the remaining world oceans. Arctic Ocean deeps are separated into two large basins by the central Lomonsov Ridge: the Canada Basin in the west and the Eurasian Basin in the east. Each of these two basins in turn is divided into two relatively distinct basins. The Canada Basin extends close to the northern edge of the Canadian Arctic Archipelago and into the western edge of the Canadian portion of the Beaufort Sea. These abyssal areas comprise about 60% of the surface area of the Arctic Ocean. Marine inputs to the Arctic Ocean include the following: (a) warm surface waters originating from the Pacific Ocean via the Bering Strait (about 0.8 million cubic meters per second, or 0.8 Sverdrups; note that one Sverdrup is a flow of 1 million cubic meters per second); and (b) cold mid-depth waters originating from the Atlantic Ocean and entering the area via Fram Strait (about 3.9 Sverdrups). The large, mostly north-flowing rivers that drain large portions of the northern hemisphere’s land masses contribute about 0.11 Sverdrups. Annual precipitation over the Arctic Ocean is low, adding only about 0.03 Sverdrups. The major outputs of water from the Arctic Ocean occur via the Canadian Arctic Archipelago (about 2.0 Sverdrups, primarily as surface waters), Fram Strait (about 1.8 Sverdrups of both mid- and deep-waters), export of sea-ice (0.10 Sverdrups on the surface), and a small amount via evaporation (0.02 Sverdrups). Water inputs to Hudson Bay originate from the Atlantic Ocean via Hudson Strait (about 0.5 Sverdrups as surface waters due to the shallow depths), Arctic Ocean water entering Foxe Basin via Fury and Hecla Strait (about 0.05 Sverdrups), and similar amounts via river flow and precipitation (about 0.05 Sverdrups). All output is via Hudson Strait (about 0.6 Sverdrups as surface waters due to the shallow depths). These sources of water differ with respect to temperature and salinity, and thus density. As a result of the input volume, density differences, and the relatively narrow areas through which both inflow and outflow occur, the Arctic Ocean is characterized by a highly structured marine environment. This structure can be observed as relatively stable currents in the area as well as stable profiles of water masses.

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Siberia

Barents Sea

Lomonosov Ridge

Chukotka Bering Sea

Chukchi Sea

Eu

Arctic Ocean

ra

sia

n

Ba

sin

Fram

Stra it

Bering Strait

Alaska Yukon

Arctic Archipelago

Baffin Bay

it

Foxe Basin

ra

Nunavut

St

Northwest Territories

s vi

Fury & Hecla Strait

Atlantic Ocean

Da

Pacific Ocean

Beaufort Sea

Nares Strait

Canada Basin

Labrador Sea North Pole

Hudson Bay

3500 m 2000 m 1000 m 200 m Provincial boundaries, limit of Canadian Territorial Waters & Exclusive Economic Zone

Fig. 1. Polar projection of the Arctic Ocean with depth zones indicated. Also shown are the political boundaries, Canadian territorial waters (12 nm), and Exclusive Economic Zone (200 nm) bordering the area of interest.

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INTRO DUCTION

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Water Masses  Both the temperature and the salinity of the different input waters vary significantly. Typical temperature and salinity profiles for two Canadian areas of the Arctic Ocean are shown in figure 2.

Fig. 2. Typical temperature and salinity profiles in Baffin Bay and Beaufort Sea (modified from Wadhams, 2000, after Aagaard and Carmack, 1989). Salinity and temperature differences alter water densities, which maintain the integrity of the water masses indicated.

Thus, in combination, temperature and salinity as well as other characteristics of the water can be used to distinguish water masses in specific areas. Furthermore, both temperature and salinity affect the density of sea water. As a result of the density differences present between the various water inputs to the Arctic Ocean, and influenced to a large degree by ice presence, little mixing occurs between Arctic water masses (fig. 3). This results in a depth-structured marine

environment as well as a spatially structured environment, both of which influence the fish fauna. Several distinct layers, most prominently seen in the Canadian Beaufort Sea, can be defined for the Arctic marine environment. This layering occurs across the geographical extent of the Arctic Ocean proper, albeit with some local modifications. Less complicated but similar situations also occur in the sub-Arctic seas of the Canadian Arctic (i.e., Baffin Bay–Davis Strait, and Hudson Bay complex). The surface is dominated by an ice layer consisting in part of the polar pack and in part of seasonal summer ice (see below). Beneath the ice, the uppermost layer of water, the Arctic Surface Layer, extends to a depth of about 200 meters. The upper 50 meters of this layer is referred to as the Polar Mixed Layer, which in winter becomes hypersaline as the ice forms and salt is extruded; in summer it becomes hyposaline due to ice melt and freshwater inflow from terrestrial areas. Below this layer, occupying the next 150 meters or so, is the Halocline, a complex zone with waters of varying densities. In the western Canadian Arctic the Halocline is dominated by waters originating from the Pacific Ocean and flowing through the Bering Strait; hence it is referred to as the Pacific Halocline. In the eastern Canadian Arctic the Halocline is dominated by waters originating from the Atlantic Ocean and is referred to as the Atlantic Halocline. In the summer, in nearshore areas with large freshwater inputs, the Polar Mixed Layer may be very freshened and extend well offshore. Fish species that are less saline tolerant such as some Arctic anadromous salmonids, as well as some primarily freshwater species, may occur in these freshened surface waters. In these areas at this time, but below the surface layer and in the Halocline Layer, fish with a wholly marine ecology may also be present. These differing associations with depth and water density (salinity and temperature) explain the occasional spatial co-occurrences of freshwater, anadromous, and marine fish species that have been reported in some areas. Depending upon the adjacent river discharge and the season, such freshened areas may extend well offshore. Wind and local water currents eventually disrupt such freshened zones, especially as these weaken further offshore. Offshore in the western Canadian Arctic, the Polar Mixed Layer is more saline, but

Fig. 3. Water masses present across the Arctic Ocean from the Bering Strait to Fram Strait. Approximate residence times of the waters are shown in bold type as years.



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Fig. 4. Surface currents and ice movements of the Arctic Ocean (to approximately 200 m depth); these interact with surface waters of Pacific origin.

due to its Pacific source it is typically less saline in the summer than are the deeper waters. In winter freshwater inputs decrease substantially or cease, and ice forms with accompanying extrusions of salt. This results in hypersaline surface waters. Such waters are often very cold with temperatures several degrees below zero. This combination of hypersalinity and negative temperatures physiologically excludes many species of fish from such areas; thus, most Arctic anadromous fishes leave the Arctic marine environment during winter. Similarly, in winter many marine fishes likely leave the hypersaline, cold surface layers and move to deeper, less-saline, and warmer waters. Below the Halocline, extending from about 200 meters to 800– 900 meters depth, the Atlantic Layer is present. This water primarily originates from the Atlantic Ocean and extends north and west around the central Arctic Ocean basin, eventually turning to northern Greenland and the Canadian Arctic Archipelago. In Canadian areas it is found in the deeper portions of the eastern edge of the Beaufort Sea. In such waters, marine fishes with faunal associations to the Atlantic Ocean can be found. Thus, as an example, Greenland Halibut (Reinhardtius hippoglossoides) are found in the Atlantic Layer in the western Canadian Arctic as well as in eastern Canadian Arctic areas such as Davis Strait and Baffin Bay. Below the Atlantic Layer, from 900 meters to the bottom, is the Arctic Deep-Water Layer, sometimes referred to as the Arctic bottom water. These deep waters are divided by submarine ridges into the various basins noted above. Little exchange of water from the various basins occurs, and due to a dearth of work little is known

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about the fish fauna of these waters. There is the potential for some degree of endemism among the fish in such waters. The layering noted above effectively isolates the various water masses from each other, and little or no mixing occurs except in the upper surface waters. This factor, the various currents present in the area, and the semi-isolation of various portions of the Arctic Ocean by submarine ridges and continental shelves result in varying residency times for the water masses. Thus, the average residency time for Pacific and Atlantic surface layer waters in the Arctic Ocean is about ten years, that for the Atlantic Layer is about thirty years, and that for the Arctic Deep-Water Layer varies from seventy-five to three hundred years depending upon the basin, for example, water in the Canada Basin has the longest residency time of any in the Arctic Ocean.

Currents  As can be expected from the above discussion of water masses, the entry of various water masses results in the presence of a complex series of currents in the Arctic Ocean (fig. 4). Similarly to the water masses, these currents are structured by depth. Surface currents consist of two major entities: Transpolar Drift and Beaufort Gyre. These affect both the polar pack and the seasonal ice and determine to a great degree the way in which ice moves in the Arctic Ocean as well as the way in which the surface layer of water moves both during the open water season in the summer and under the ice in winter. The Transpolar Drift generally moves surface water (i.e., to 50 m depth, consisting of Pacific waters, entrained

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Fig. 5. Current flows affecting Pacific waters in the Arctic, primarily between 50 m and 200 m depth (i.e., Pacific Halocline Layer); these interact with other surface waters.

fresh water, and ice) across the Asian side of the geographic North Pole from the area of the Bering Strait to Fram Strait and the Barents Sea. The Beaufort Gyre is a clockwise rotation of surface water and ice centred upon the North American side of the pole over the Canada Basin. Ice and water tend to be mixed to a limited degree between both the Transpolar Drift and the Beaufort Gyre. Minor surface currents originating from these major currents are also responsible for the export of ice and surface waters out of the Arctic Ocean into marginal Arctic seas and ultimately into the North Atlantic Ocean. Much of this material is exported through the various channels of the Canadian Arctic Archipelago as well as through Nares Strait between Canada and Greenland. Depending upon the nature of the ice being exported (i.e., first year vs. multiyear) and the prevailing environmental conditions (i.e., warm vs. cool summer), such ice can choke these channels throughout the year. Ice and surface water is also exported out of the Arctic Ocean by surface currents through Fram Strait and, to a lesser degree, through the Barents Sea. The total annual export of ice from the Arctic Ocean varies between 0.8 million and 1.3 million cubic kilometers. To place this in context, the major rivers draining into the Arctic Ocean provide about 2,555 cubic kilometers of water to the Arctic Ocean per year. Pacific Ocean waters enter the Arctic Ocean via the shallow Bering Strait (with a sill depth of about 50 m) and occupy the 50–200 meter depths of the Halocline Layer (fig. 5). A portion of this water is incorporated into the Polar Mixed Layer, thus joining both the Transpolar Drift and the Beaufort Gyre. A portion of Pacific water



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is incorporated into a longshore eastward flow along the northern coast of Alaska into the Canadian portion of the Beaufort Sea, subsequently entering the southern portion of the Canadian Arctic Archipelago, and then exiting into Hudson Bay, and flowing out of the Arctic via Hudson Strait. Another portion flows eastward along the northern edge of the archipelago but below the Beaufort Gyre and is incorporated into flows entering the various channels of the northern Canadian Arctic Archipelago. Some of these waters flow south through Nares Strait between Canada and Greenland, and a further portion flows out through Fram Strait. Additional surface currents affect the shallow waters of Davis Strait and Hudson Bay (0–200 m depth) (fig. 6). The flow of Atlantic waters into Davis Strait and Baffin Bay occurs via the west Greenland Current along the nearshore of Greenland. This current flows north until it encounters the outflow from the Arctic Ocean through Nares Strait. These waters mix and form the southward-flowing Labrador Current along the eastern nearshore of Baffin Island, eventually bordering the western edge of the Labrador Sea, then entering the North Atlantic. Various smaller surface currents gyre westward across Davis Strait and join the Labrador Current. At the south end of Baffin Island a westward current splits off and enters the northern side of Hudson Strait, providing inflow of Atlantic waters into Hudson Bay. This current generally moves northward, following the coast of Foxe Basin in a counter-clockwise fashion. In western Foxe Basin it is joined by the outflow of Arctic Surface Layer that occurs through Fury and Hecla Strait. Together these waters flow south along the western edge of Hudson and James Bays, then turn

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Fig. 6. Current flows affecting the Atlantic Layer in the Arctic, primarily between 200 m and 900 m depth. Some mixing occurs with deeper water to ca. 1,700 m.

north to flow along the eastern coasts. Outflow occurs via a surface current along the southern portion of Hudson Strait, and this water eventually links with the southward-flowing Labrador Current, then enters the North Atlantic Ocean. The shallow depths in Hudson Strait prevent the entry of separate deep-water flows from the Atlantic Ocean into Hudson Bay. Beneath the Halocline Layer, waters of 200–900 meters in depth form the Atlantic Layer and enter the Arctic Ocean from the Greenland and Norwegian Seas via Fram Strait. These waters form complex mid-depth circulation patterns that move in the following ways. First, an eastern portion circles the Arctic Ocean in a counter-clockwise direction, eventually moving eastwards along the Eurasian shelf drop-off, then south along the Canada Basin, thus entering the western portions of the Canadian Beaufort Sea. This water mass then moves east along the shelf drop-off bordering the Canadian Arctic Archipelago. Second, portions of this deeper water form several gyres in the deeper basins and sub-basins of the Arctic Ocean. These generally tend to gyre in a counter-clockwise fashion across the polar region from the Asian to the North American sides. Most of these waters exit the Arctic Ocean still at depth via Fram Strait. Atlantic deep water also flows northwards along the western side of Greenland through Davis Strait to Baffin Bay. This water occupies the deeper areas of the eastern Canadian Arctic and crosses Baffin Bay, then flows south along the eastern coast of Baffin Island. Such water eventually exits to the North Atlantic via the Labrador Sea. As noted above, the Arctic Deep-Water Layer has a long residency time. This water originates in part from the deep waters of

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the Norwegian and Greenland Seas. Limited mixing with the Atlantic Layer likely occurs. Relatively poor understanding of the deep-water movements exists, but, as noted earlier, they are complicated by the various submarine ridges present in the Arctic Ocean. The various water masses and currents described above enter and exit the Arctic Ocean and the marginal Arctic seas differently. They also tend to maintain their integrity to a large degree. A biological consequence is differential drift and movements of larval as well as older marine fish in the Arctic Ocean. Thus, the variable water masses and currents, combined with the differing geographical regions they influence, can be expected to determine to a large degree the zoogeography of Arctic marine fishes. Detailed analysis of this has not yet occurred, but initial examples are known, for example, Greenland Halibut (Reinhardtius hippoglossoides) of the Canadian Beaufort Sea, and the dominance of the fauna of Hudson Bay by Atlantic forms of marine fishes.

The Arctic Environment Seasonality of the Arctic The environment of the Arctic is characterized by extreme seasonality. This seasonality is typically recognized where it affects terrestrial and freshwater environments, but it is also highly relevant in many parts of the marine environment. The daily duration of incident light that drives primary production ranges from none throughout much of winter to twenty-four hours during summer, and the

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Canada Basin

Lincoln Sea

Greenland

Beaufort Sea

Alaska

Kane Basin

Baffin Bay

Foxe Basin

it

Hud son Str

ait Ungava Bay

Hudson Bay James Bay

Canada

Legend Canadian Exclusive Economic Zone Record minimum sea ice extent, September 2012 median Polar pack minimum ice extent, 1969–99 median Hudson Bay Drainage Basin Arctic Seaboard Drainage Basin Mackenzie River Drainage Basin

diel duration, angle of incidence, and intensity all vary with latitude. Climatic parameters of air temperature, wind, and precipitation, for example, reach extreme values and seasonality in the Arctic. These affect the marine environment in differing ways but exhibit their greatest influence in nearshore zones. For example, wind drives surface currents and mixes fresh and saline waters; thus, it may be extremely important in the marine life histories of anadromous fishes (see Family Salmonidae). Similarly, precipitation, both as runoff from terrestrial areas and as deposition directly on ice, though typically lower in the Arctic than further south, greatly influences the amount of fresh water present in nearshore zones or surface layers of the marine environment. This in turn affects salinity gradients and so determines to some degree the composition of nearshore and upper-pelagic marine fish communities. Air temperatures, though exhibiting extremely low values in the winter, have little direct effect on marine fish; rather, the effect is indirect in that it is mediated through the formation of ice, which in turn is relevant to marine fish. In contrast to the air, marine water temperatures are generally quite stable, especially within a water mass, and from a biological perspective are relatively benign at an average of about 2°C. This is especially true in offshore areas and in deeper waters under ice. However, water temperatures in shallow nearshore areas are greatly influenced by warm freshwater inputs from nearby rivers as well as by air temperatures. Warm, freshened areas are also zones in which the influence of wind is high, and these result in a mixing of fresh and marine waters. Such mixing supports high local productivity, and these areas are physiologically preferred by many fish species.



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tra

Fig. 7. Canadian Arctic marine waters with contributing freshwater drainage basins. Until recently, multi-year pack ice dominated much of the northwestern marine areas and some archipelago channels during the open-water season. Climate change has reduced the extent of the sea ice from the longerterm median, thus allowing greater human access to marine areas.

S vis Da

Mackenzie River

These effects are especially relevant during the open-water season. For rivers that continue to flow during the winter, the development of freshened zones under the land-fast ice may offer lower salinity habitats that are relevant to some species. Canadian Arctic marine waters represent a diversity of environments united by the common presence of ice. Ice dominates most of the area in all seasons and greatly affects physical, chemical, and biological processes. These in turn influence Arctic marine fishes. In winter about thirteen million square kilometers of the Arctic Ocean are covered by ice, including the Canadian portion of the Beaufort Sea, the waters of the archipelago, Hudson and James Bays, Baffin Bay, and much of Davis Strait as well as northern portions of the Labrador Sea.

Ice Coverage and Type In summer the ice cover over the Arctic Ocean is reduced to about nine million square kilometers as a long-term average – to even less in recent years – and many of the Canadian Arctic marine areas become ice free for a few months of the year. However, polar pack ice remains present throughout the summer in the northern portion of the Canadian Beaufort Sea and along the northwestern archipelago islands. It should also be noted that in most cases “ice-free” areas may have drifting remnants of winter ice, and there is high interannual variability in the presence, coverage, age, and thickness of the sea ice that persists in the summer. In summers that approach long-term normal air temperatures, almost complete ice coverage also occurs in many of the channels of the archipelago as a result

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Fig. 8. Polynyas and shore leads of the Canadian Arctic (after Hannah, Dupont, and Dunphy, 2009). (1) Cape Bathurst (2) Lambert Channel (3) Roes Welcome Sound (4) Committee Bay (5) Foxe Basin (multiple) (6) Frobisher Bay (7) Cumberland Sound (8) Fury and Hecla Strait (9) Franklin Strait (10) Bellot Strait (11) Prince Regent Inlet (12) Lancaster Sound (13) Viscount Melville Sound (14) Karluk Brooman (15) Queens Channel and Penny Strait (16) Dundas Island (17) Hell Gate-Cardigan Strait (18) Lady Ann Strait (19) Bylot Island (20) Coburg Island (21) North Water (NOW) (22) Flagler Bay (23) Lincoln Sea

of the clogging of the narrow passages by multi-year ice that has drifted off the polar pack or formed locally. During summer other areas generally become ice free or are covered to a limited degree by loose, unconsolidated ice floes. These conditions typified the area until the mid-1990s; since then substantive changes in climate and associated sea-ice dynamics have become evident (e.g., the summer sea-ice areal extent is now about 4.5 million square kilometers, see fig. 7, with the record minimum, in 2012, being lower and the southern margin of polar pack ice being displaced well to the north in our area; see also the “Climate” section in the introduction). Ice in the Arctic can be differentiated into two types depending upon its age: seasonal ice and multi-year ice. Seasonal ice forms and melts more or less completely on an annual cycle. It is typically formed in the west in coastal zones along the mainland and the southern fringes of the Canadian Arctic Archipelago, and in the east throughout Hudson and James Bays, Hudson and Davis Straits, and Baffin Bay. Seasonal ice is relatively thin (up to 2.5 m) and is present in most areas from about October to July. If, for whatever reason, it does not melt completely in the first summer, it is incorporated into multi-year ice. This, now second-year, ice characteristically persists over the continental shelf portion of the marine environment and can be termed summer ice. Successive annual accumulations of summer ice form multi-year sea ice that is typically present as fringing ice on the edge of the polar pack and throughout the various channels of the Canadian Arctic Archipelago, Foxe Basin, and Kane Basin north of Baffin Bay. Typically such ice is three to eight meters thick and is dynamic in the sense that it undergoes a seasonal melt and re-formation cycle as well as generally moving south and

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east through the archipelago. Outside of the continental shelf to the north and west of Canadian land areas, polar pack ice is present over the abyssal zones of the Arctic Ocean. This is multi-year ice of relatively great ages and can be very thick (to 50 m) in some areas as a result of pressure ridging or the incorporation of ice shelves that have calved from northern Ellesmere Island, although overall thickness has been diminishing in recent years. The Arctic Ocean polar pack is typically about six million square kilometers in area but has recently been reduced in overall extent. In nearshore areas seasonal ice forms as land-fast ice affixed to the shore and is relatively immobile. In offshore areas seasonal ice forms and, together with multi-year ice, fringes both the polar pack and the land-fast ice. Offshore seasonal and multi-year ice, as well as the polar pack, are constantly in motion, driven by currents, tides, Coriolis forcings, and wind. This dynamical nature of the ice results in a number of features that are extremely important to the ecology of Arctic marine fishes. First, ice of varying types forms a physical, relatively productive habitat (see the “Habitats” section in the introduction). Stable, seasonally present, land-fast ice also allows for human activities, which may include travel to and from fishing sites and fishing through the ice (see the “Fisheries” section in the introduction). Ice also offers a refuge from predators (i.e., marine mammals and birds) for some species of marine fish. Second, pack-ice movement creates pressure ridges in the ice that manifest as deep keels of ice below the surface; these are often very deep and can physically scour the bottom in shelf areas and directly affect habitats for marine fishes as well as other organisms. Third, shear zones between the moving pack ice and immobile land-fast

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ice typically form regular openings in the ice cover that are termed shore leads (see fig. 8). In many areas these leads recur every year and are regularly formed, but locally variable, features of the environment; they typically open wider early in the polar spring, exhibit high local productivity, and are of great significance to the marine biota in the areas. Fourth, during the winter the shear zones in the vicinity of large rivers form deep, keel-like areas of broken ice offshore of the deltas, referred to as stamukhi. In areas where freshwater flows continue under the ice during winter (e.g., Mackenzie River), stamukhi can trap large volumes of fresh water either on top of heavier marine waters or completely displacing marine waters. Due to ice cover there is little mixing of these waters, and the areas provide freshened overwintering habitat critical to fish species that prefer low salinities (e.g., anadromous coregonines of the western Canadian Arctic). Fifth, open-water areas surrounded by ice, called polynyas, occur in many locations (see fig. 8). Polynyas may either remain open throughout the year or freeze over in the coldest months. They are associated with recurring leads and result from a combination of upwelling (especially warm) water from deeper sources, mixing currents, wind, and tides. Polynyas are usually areas of very high local productivity. Thus, despite occupying only a small area of the Arctic, polynyas and shore leads are extremely important in the ecology of the Arctic marine biota including fishes. Both the open water and the higher productivity associated with polynyas and shore leads act as focal points to concentrate biota in Arctic marine waters; this includes not only the fishes but also their prey and predators.

Tides The tides present in the Canadian portion of the Arctic generally are relatively small, varying from 0.5 meters in the north and west to 1–5 meters in the east and southern areas, and rarely to 7–12 meters in places. Ice affects the importance of tides in Arctic waters. Tidal movement of ice results in physical scouring of the intertidal areas of the nearshore in addition to any scouring due to general ice motion. As a result, these areas tend to be relatively depauperate with respect to benthos and fixed biotic components such as kelp. Two consequences are pertinent to Arctic fish ecology: first, the intertidal community is displaced to deeper waters below this scour zone; and, second, due to reduced light penetration in such deeper areas, this combined intertidal and subtidal community typically occupies a smaller area of the nearshore zone than do either of its components in ice-free zones. These two factors result in a comparatively depauperate fauna in the Arctic of fishes and other organisms adapted to intertidal habitats (see also the “Habitats” section in the introduction).

Arctic Marine Biota Age and Derivation of the Arctic Marine Ecosystem Geological evidence indicates that the Arctic Ocean has existed since Paleozoic times (i.e., about 81 million years before the present). However, the area has undergone considerable modification since



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then due to plate tectonics, shifts in the geographical position of the poles, repeated sea-level changes, Pleistocene glaciations, and related changes in climate. Arctic sea ice was first formed in the middle Pliocene epoch apparently as a result of climatic deterioration in high latitudes. The thickness of the ice at that time was likely much greater than it is today as suggested by a lack of diatoms in bottom sediments, assumed to have resulted from low light penetration. The appearance of diatoms about 100,000 years ago indicates generally increasing temperatures in the area and concomitant decreasing ice thicknesses. The modern characteristics of the Arctic sea-ice system were acquired at about this same time, and, similarly, it can be assumed, ice-associated biota also became more widespread. The most significant pervasive recent influence on the Arctic Ocean has been the Pleistocene glaciations that occurred over the last million or so years, with the latest Wisconsinan advance ending about 25,000 years ago. The effects of the last glaciation have been well studied with regard to land areas and the related biota but are poorly known for marine areas. At the glacial maximum the majority of the Arctic Ocean was covered by permanent polar pack ice, which likely had characteristics similar to those observed today but was much more extensive in area and thicker, possibly with reduced seasonal melt. This would have limited light penetration and hence biological productivity. This polar pack ice was likely not consolidated with glaciers except in coastal areas where glaciers grounded in the sea. In the western Canadian Arctic (i.e., offshore Beringia) and similar areas throughout the Arctic Ocean, seasonal ice formation and melting, and related shelf productivity, probably occurred similarly to that of today. This would be especially widespread where rivers entered the Arctic Ocean, such as Beringia and the northern coast of Siberia. Thus, much of the marine and anadromous fauna of the western Canadian Arctic would have been displaced westwards during the glacial advance but likely would have been similar to the fauna present in the area today. In the eastern Canadian Arctic the situation appears to have been quite different. Environmental reconstructions based upon foraminifera present in sea sediments suggest a steep thermal gradient extending across the glacial North Atlantic Ocean, centred upon 42° N latitude. South of this was a relatively stable southern zone in the glacial Atlantic Ocean. North of this was a dynamic zone that was polar in character and which likely underwent seasonal cycles similar to those observed today in the polar pack. In the northern zone, year-round polar-pack ice appears to have been extensive over all of present-day Baffin Bay, Davis Strait, and the Labrador Sea. Similarly, the Greenland, Iceland, and Barents Seas were covered by polar pack ice. In the Pleistocene North Atlantic, south of the polar pack, a large zone of seasonal ice was present that extended south to the present-day Grand Banks area of Newfoundland. Unlike today, much of the productivity of this area of the Arctic Ocean was likely considerably lower and displaced southwards into the present-day North Atlantic Ocean. Thus, polar marine fish faunas of the eastern Canadian Arctic were likely similarly displaced southwards for the most part. It is also probable, however, that some marine fishes

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persisted in low numbers under much of the pack ice throughout the entire Arctic Ocean, similar to today. Most of the Canadian Arctic Archipelago and marginal seas were covered by continental glaciers during the Pleistocene. The exceptions were northwestern Banks Island, nearby portions of the Beaufort Sea, and perhaps extreme northern Ellesmere Island. These continental ice sheets effectively isolated marine areas of the western Canadian Arctic from eastern Arctic areas. Interior Arctic seas (Hudson and James Bays and adjoining areas) were also covered by ice sheets. Due to the incorporation of water into glacial ice, Pleistocene sea level was between 100 and 200 meters lower than it is today, which exposed areas of land that are present-day continental shelf areas. Thus, Beringia (unglaciated Alaska, land masses in the Bering Sea region, and Chukotka) formed a land barrier between the Pacific and Arctic oceans. The latest glacial advance and retreat occurred over a period of at least 125,000 years, but perhaps as long as 250,000 years; thus, the separation was for a substantial period of time. The separation of the Arctic Ocean and the Pacific Ocean by the Beringian land mass occurred several times during the Pleistocene. Presumably during each phase, marine fishes from the North Pacific Ocean had access through the Bering Strait and moved into the Arctic marine waters of western North America. Successive waves of colonization, isolation, and differentiation thus likely affected the fish faunas in this area.

Sources of Arctic Marine Fishes  Fishes in Arctic marine waters can be differentiated as wholly marine (i.e., spend their entire lives in the sea) or as anadromous (i.e., move between marine and fresh waters during life). These types of fish populated Arctic marine waters by different pathways and likely also at different times; thus, they must be considered separately. Furthermore, Canadian Arctic marine areas consist of continental areas that were glaciated and offshore areas that were not. The timing and sources for populating these areas therefore differed as well.

Marine fishes in unglaciated areas Wholly marine fishes present today in the Arctic Ocean appear to be derived from three separate components: pre-Pleistocene relictual fish present in the Arctic Ocean, recent Pacific fish, and recent Atlantic fish. The first faunal element (Arctic relictual) appears to represent a small proportion of the present-day marine fish fauna and to be derived from temperate and boreal marine areas that bordered the Arctic Ocean during preglacial times. A few species may also be considered to be Arctic endemics (e.g., Polar Cod, Arctogadus glacialis), having evolved in situ. All these species were mostly derived from the Pliocene to recent times. In the Canadian portion of the Arctic Ocean many of these fishes likely originated as a result of the repeated early connections with the Pacific Ocean. However, a significant part of this relictual element is also represented by Atlantic taxa that probably colonized the Arctic long before the Pleistocene period. The second faunal element (recent Pacific) represents postglacial elements derived from the Pacific Ocean during the successive inundations of Beringia. These species are well represented especially in the western Canadian Arctic.

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Many are nearshore species that likely reflect the shallow nature of the pathway (i.e., the present sill depth in the Bering Strait is about 50 m), as well as being perhaps nearshore marine fishes that survived along the northern fringe of Beringia during glacial times. The third faunal element (recent Atlantic) comprises the greatest proportion of the Arctic marine fish fauna. As noted earlier, large portions of the Arctic Ocean and adjacent seas were not glaciated but were likely covered by polar pack ice for much of the Pleistocene. From the perspective of wholly marine fish, it appears that for long periods during the late Pleistocene the only significant link between the Arctic Ocean and southern oceans was to the Atlantic Ocean via Fram Strait and the Norwegian Sea. Also, fish in this group were likely associated with seasonal ice ecosystems that were present in the North Atlantic Ocean at the glacial maximum, and they followed the retreating edge of this area as it moved northwards in recent times. Therefore, Atlantic marine fishes more or less had continuous access to the Arctic Ocean throughout the Pleistocene, whereas the connections for Pacific fish were limited to the times between glacial advances (i.e., interglacial periods during which the Bering Strait was inundated).

Marine fishes in glaciated areas Deglaciation of continental ice sheets in the Canadian Arctic occurred in a northeasterly direction beginning about 18,000 years ago. The last remnants of the ice masses centred over the northeast channels of the archipelago disappeared about 6,000 years ago; however, present-day remnants of the Pleistocene ice remain as ice caps and glaciers on land. The sequential disappearance of the ice and removal of intervening barriers resulted in a number of corridors being open to fish colonization of newly exposed areas. First, a marine connection through Hudson Strait between the western Atlantic and present-day Hudson Bay opened about 8,000 years ago, allowing marine fishes to penetrate into present-day James Bay and southern Hudson Bay while the northern areas of Hudson Bay and Foxe Basin were still covered by glacial ice. Subsequent deglaciation of Foxe Basin allowed Atlantic marine fishes to colonize this area. Further north, the eastern portions of the archipelago became ice free about 7,000 years ago. This allowed limited penetration of Atlantic faunal elements westward into the deeper portions of the eastern archipelago. The westward penetration of these species would have been limited by the development of modern current flows from the west through the archipelago, coupled with rising land masses as isostatic rebound occurred. Although detailed analyses are not available, it appears that the area of the Boothia Peninsula and Somerset Island in the central Arctic forms a zoogeographic discontinuity (e.g., faunal differences are present east and west of here, and there is evidence for genetic discontinuities in some taxa) separating eastern and western Arctic faunas. This possibility requires further investigation. Second, as sea levels rose during deglaciation, the central portion of the Beringian land bridge became inundated and the Bering Strait opened about 11,000 years ago. This allowed some species of Pacific Ocean fish to colonize the western Arctic. However, as noted earlier, shallow sill depths in the Bering Strait then and continuing

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to the present time would have restricted the types of fish that did so (i.e., shallow-water species for at least part of their life history), although larval drift in surface waters would have remained possible. These new species, along with any taxa originally from the Pacific Ocean that had survived in coastal areas of north Beringia and perhaps eastern Siberia, then formed the core of species that colonized the western Canadian Arctic marine areas. Deglaciation of the marine channels of the western archipelago was complete by about 10,000 years ago, and modern water masses and currents were likely set up at about the same time. The generally eastward and northeastward flows would have facilitated marine fish colonization of the western archipelago beginning at this time. Similar to the Bering Strait, however, shallow sill depths at the entrances of the Canadian Arctic Archipelago, combined with the water mass characteristics and circulation patterns discussed earlier, likely restricted the types of faunal elements that had access to this area. The colonizing faunal elements thus were likely primarily shallow-water, Pacific species, or western Arctic species that predated glaciations. Some of these species also appear to have colonized northern Hudson Bay once Fury and Hecla Strait had opened. Furthermore, entrainment of large volumes of fresh, glacial meltwater in this area likely further restricted the types of marine fishes that entered the region, as well as the distance into the archipelago that they penetrated; that is, both fresh water and generally eastward currents would have been a barrier to any marine fish species that were stenohaline. From the perspective of anadromous Arctic fish, several socalled refugia appear to have been present in North America in which both anadromous and freshwater forms existed. Arctic species that presently exhibit anadromy could have survived in these refugia as anadromous forms (A), freshwater forms (F), or both in some cases. Freshwater forms could also have become anadromous secondarily following re-invasion of deglaciated areas (i.e., Fa). Taxa that exhibit this possibility include derivatives of Lake Whitefish (Coregonus clupeaformis) forms, Lake Trout (Salvelinus namaycush), and perhaps also Burbot (Lota lota) (see the “Family and Species Accounts” section) originating from southern continental refugia. The potential refugia, and their possibility as sources for anadromous or freshwater, secondarily anadromous colonizers (Fa), include the following: south coast Beringia (A, Fa); north coast Beringia including the unglaciated portions of the northwestern archipelago (A, Fa); possibly northern Ellesmere Island (Fa only?); Missourian/Mississippian refugia south of the continental ice masses (F, Fa); and Atlantic coastal refugium south of the ice sheet (A, Fa). The latter could also have included similar coastal refugia in Greenland and perhaps areas further east towards Europe. Anadromous species in the western Arctic (osmerids, coregonines, chars, and salmons) originated primarily from Beringian refugia, likely with a strong bias for species present in north Beringia during the glacial maximum. Additionally, there appear to have been contributions of freshwater, secondarily anadromous taxa from the Mississippian and the southern Beringian refugia that entered the western Arctic via glacial lakes along the Mackenzie River basin and via drainage reversals of the Peel River, respectively. The southwestern Canadian archipelago was likely recolonized by



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anadromous forms from northern Beringia and by freshwater forms from the Mississippian refugium. The central, northern, and eastern archipelago was likely recolonized by a combination of anadromous forms from Atlantic refugia and of freshwater forms from the northern Ellesmere refugium (if this supported fishes), with some contribution potentially coming from the Mississippian refugium via Hudson Bay and from northern Beringia. Recolonization of James and Hudson Bays, Foxe Basin, and Hudson Strait likely resulted primarily from Mississippian and Atlantic refugia south of the main continental glacial ice masses. Ice and shallow sill depths combine to form barriers not only to fish but also to water masses. Thus, large amounts of fresh water from rivers that began draining north to the Arctic Ocean, as well as glacial meltwater during deglaciation, likely resulted in highly freshened bodies of water entrained under sea ice. These freshened areas dominated parts of the Canadian Arctic especially in the nearshore shallow zones of the west. Continued presence of glaciers over the eastern archipelago for much longer would have restricted the exit of such water from the area; thus, these freshened zones likely persisted throughout the year and for long periods of time. This would have facilitated colonization of deglaciated areas by both freshwater and anadromous fishes but would have restricted colonization by stenohaline marine fishes. Such events likely account for the significant eastward distribution of coregonines in the central Arctic, for the northern penetration of Arctic Char (Salvelinus alpinus) to the limits of land, and for the colonization of western Hudson Bay by various anadromous taxa.

Summary Generally the present-day Arctic marine ecosystem appears to be a relatively young ecosystem that has yet to equilibrate from the recent glaciations. The evidence to support this consists of the following: little endemism in the biota, reduced numbers of genera in comparison with both the Atlantic and the Pacific oceans, dominance of few species in terms of both numbers and biomass especially in shelf areas, general reduction in species’ complements and standing stocks in comparison to more southerly marine ecosystems, and high levels of biologically plastic organisms. Plastic species tend to be generalists and so exhibit wide tolerances to variable biological and environmental conditions (e.g., they are euryhaline and eurythermal) and may be seen as having high levels of morphism or variable forms. Although it is difficult to assess, given the poor coverage for some areas, distributional changes also appear to be presently occurring in parts of the Arctic. This further suggests that the present-day Arctic marine ecosystem is ecologically young and in flux.

Polar Productivity Arctic marine waters generally exhibit low and highly pulsed productivity due to the seasonality of incident radiation. Most areas appear also to be nutrient limited; thus, a general average figure for primary productivity is less than fifty grams of carbon per square meter per year. This is about 20% or less of the productivity that is

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typical of the more southerly oceans. Locally important and highly productive areas occur throughout the Arctic. These are typically associated with shelf areas that are influenced by nutrient input from large rivers, local offshore upwelling, or both. Similarly, perennial polynyas and shore leads often form along the shelf drop-off and thus are near upwelling or mixing zones that result in high local productivity. As can be expected in areas of increased productivity, the diversity of all organisms including marine fishes, as well as the numbers and biomass within species, is much increased. Ice creates two additional zones of productivity. The first is the under surface of the ice, which forms a complex physical and chemical surface for primary production by ice-associated diatoms. This, in turn, results in a relatively complex community of secondary producers (e.g., gammarid crustaceans) that then provide food for cryophilic predators such as Arctic Cod (Boreogadus saida). Fish such as Arctic Cod, in turn, are pivotal components of the sea-ice ecosystem that are being preyed upon by marine mammals and sea birds. Production associated with the under-ice community can account for about 15% or more of the total local Arctic marine productivity. The second area of high productivity associated with ice is the ever-changing ice edge along leads, and the melt and fracture zones. As expected, this is a highly dynamic habitat, but it shares many of the characteristics of polynyas (e.g., local nutrient input), thus acting as a focal point for the biota. Surface, early-season, meltwater ponds on sea ice may also contribute to local productivity; however, the significance of this is uncertain. Thus, despite the common perception of overall low productivity, areas within the Arctic Ocean can be very productive locally. Given that there are large numbers of such areas present throughout the Arctic, the absolute amount of production is relatively high despite the pulsed nature of energy input from the sun. One aspect of this productivity is associated with the export of ice and nutrient-laden waters from the Arctic Ocean. As noted, ice and water export is substantial in terms of the volumes moved by Transpolar Drift to the North Atlantic Ocean via Fram Strait and through the Canadian Arctic Archipelago. The exported ice and water is nutrient laden and contains large amounts of ice-associated organisms such as mats of algae and invertebrates. This material is deposited into the local seas and is believed to contribute substantially to the productivity of the Labrador Sea in the western North Atlantic and the marginal seas of the eastern North Atlantic (e.g., the Greenland, Iceland, and Norwegian Seas). Similarly, nutrient-laden outflows from the archipelago into Foxe Basin likely contribute to locally high productivity there, and outflows from both the archipelago and Hudson Bay contribute significantly to marine productivity in Hudson Strait and southern Davis Strait and along the Labrador coast.

Adaptations of Arctic Marine Fishes There are fewer species of Arctic marine fishes present in comparison to those of other oceans. Within species generally there is also less specialization, although local specialization may be exhibited as increased morphism in Arctic fishes. As described earlier, the Arctic

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Ocean marine environment is vastly different and in many areas is much more rigorous, especially with respect to variability, than are temperate marine environments. It is a logical extension then to note that many Arctic marine fish species exhibit special adaptations that allow them to occupy such environments. As has been generally noted, Arctic fish appear to have relatively wide tolerances to many environmental parameters. Most Arctic marine fishes, particularly coastal and shelf species, are eurythermal and are especially tolerant of the high summer temperatures that regularly occur in some habitats such as the nearshore. Similarly, Arctic marine fishes, especially those of the nearshore areas, appear to be euryhaline and therefore tolerant of wide ranges of salinity. Whether such tolerance is biased towards low salinities, or whether it also includes tolerances of hypersalinities such as those occurring under early winter ice, is unknown. The low overall and highly seasonally pulsed productivity results in a short but intense feeding period for many fish, with the consequence that energy must be stored for long periods to be used throughout the year. As a result many Arctic fish appear to be well adapted for long periods of starvation. This may be especially true for shallow-water marine (e.g., Arctic Cod) and some anadromous fishes such as Arctic Char that may not have annual access to the adjacent seas. Extreme and narrow seasonality of plankton production has resulted in the eggs of Arctic fish generally being larger than those of southern fish. The consequence is that the fish hatch at larger sizes and can begin immediately feeding on a wider range of organisms, which presumably is an advantage in short productivity seasons. Overall less light, highly seasonal light, and relatively more red wavelengths in the light generally mean that Arctic fish have larger eyes than do their southern counterparts. This, of course, is only applicable to taxa living at roughly the same depths. Reduced efficacy of eyes, at least during some periods of the year, has also increased reliance on other sensory systems such as olfaction and touch. These are seen as longer or larger tactile appendages such as barbels and pelvic fins, and as exposed cephalic lateral-line systems rather than ones enclosed in canals. Lower temperatures present in Arctic seas decrease metabolic reaction rates. Thus, various adaptations to this appear to be present in Arctic fishes. Basic compensation with respect to metabolic rates occurs; that is, physiological optima are at lower temperatures in Arctic fish compared to more temperate taxa or forms. Additional adaptations appear to include increased numbers of organs associated with metabolism (e.g., pyloric caeca); increased numbers of locomotory body parts (e.g., vertebrae and myomeres), which presumably impart greater locomotory ability in colder, more viscous waters; increased sedentary lifestyle; and likely also physiological changes such as decreased thermal optima for enzymes. Normal salinity sea water freezes at about −1.8°C. Increased salinity lowers the freezing point even more. Thus, in addition to cold tolerance, Arctic marine fishes must either emigrate from very cold hypersaline waters or exhibit anti-freeze properties. Anadromous fishes exhibit behavioural adaptations in that they leave Arctic marine waters in winter, thereby avoiding the problem. Such fish may also seek out waters of lower salinity (e.g., freshened surface waters)

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during their summers at sea to reduce osmoregulatory demands. Although not known for certain, it is likely that even widely tolerant Arctic marine fishes move out of inhospitable habitats during parts of the year. Some evidence also exists for the presence of anti-freeze solutes in some Arctic fish, but limited research has been conducted. Anti-freeze solutes in Arctic fish, unlike in their Antarctic counterparts, appear to be protein based and only developed seasonally. The presence of ice creates distinct advantages for Arctic fish in that it reduces predation from marine mammals and birds by confining these to openings in the ice. Ice also provides a habitat with associated food for some Arctic fishes. Adaptation to the under-ice habitat likely accounts for the superior and oblique mouth position of ice-associated species such as Arctic Cod in comparison to that of temperate members of the family. Similarly, there are likely buoyancy adaptations present that optimize the association of some fish with the under surface of the ice. Such adaptations may only be developed during some stages of life (e.g., relatively larger swim bladders in young fish would enhance their association with ice as a refuge from predators). High lipid content in flesh and livers also is an adaptation that enhances buoyancy and energy metabolism in many species. The above adaptations, as well as others, enable Arctic marine fishes to optimize life in highly variable and, to us, extremely rigorous environmental conditions.

begins to melt, an intense phytoplankton bloom lasting two to three weeks is triggered. Herbivorous zooplankton graze on the phytoplankton and, depending on the type of habitat over which the ice edge lies, attract predators including Arctic Cod and herring (marine habitat) or Arctic Cisco (Coregonus autumnalis), Least Cisco (Coregonus sardinella), and Rainbow Smelt (Osmerus mordax) (estuarine habitat). The fishes, in turn, attract marine mammal predators.

Estuarine Food Web Freshwater and marine food webs overlap in estuarine areas (fig. 12). The ocean front often contains high concentrations of mysids, a keystone species in estuarine food webs. They are preyed upon by Ringed Seal, Bowhead Whale, and forage fish such as Arctic Cod, Arctic Cisco, Least Cisco, Saffron Cod (Eleginus gracilis), Rainbow Smelt, Arctic Char, and Dolly Varden (Salvelinus malma). Estuarine regions are also receptacles of terrestrial organic carbon, which has been found in food webs of the Mackenzie and Alaskan Beaufort Sea shelves (fig. 12). A significant amount of this organic carbon is transported to the nearshore estuarine environment by large coastal rivers (e.g., Mackenzie River); it also results from coastal erosion.

Benthic Food Web

Arctic marine food webs can be characterized by several features: the inclusion of fewer species than in temperate marine food webs (but not necessarily fewer trophic levels); a high degree of seasonality in environmental conditions, including light, sea ice, snow cover, temperature, and nutrients; and a food-web structure that differs with distance from the coast and with depth. Sea ice also forms a distinct habitat and, along with polynyas, is a focal point for food web interactions (see figs. 9–14). The general roles of species are the same geographically across the Arctic, but the species complement influences the position in the food web, with the exception of Arctic Cod, which occurs everywhere and plays a role in all types of food webs. This single species (Arctic Cod) may effect up to 75% of the energy transfer between zooplankton and vertebrates.

Benthic food webs play a significant role in overall productivity, turnover rates, and remineralization of organic matter, particularly on shallow Arctic shelves. Primary production does not occur during the winter in the Arctic; thus the input of food into benthic food webs is seasonally pulsed. Furthermore, as depth increases, food reaching the benthic community becomes limited because the amount and quality of the sinking particulate organic matter is reduced by zooplankton-mediated processes in pelagic food webs. As a result, many benthic invertebrates are omnivorous and rely on many types of food sources such as settling sea-ice algae, phytoplankton, detritus, benthic diatoms, terrestrial input, and advected organic matter. Particulate organic matter reaching the sediment may be consumed immediately or may persist for an extended period of time, thereby providing a “food bank” for members of the benthic community. Winter survival of benthic deposit and suspension feeders provides a food source for predators and scavengers, thereby maintaining food-web stability throughout the year.

Sea-Ice Food Web

Polynyas

Ice algae inhabit the lower few centimeters of sea ice in pockets and brine channels. Ice algae typically bloom in April or May, prior to the start of the pelagic phytoplankton bloom, which usually occurs in June. A community of grazers, such as pelagic amphipods (e.g., Themisto libellula), feeds on the ice algae and, in turn, is preyed upon by forage fish (e.g., Arctic Cod). The forage fish are preyed upon by marine mammals such as Ringed Seal and Beluga. Ice algae, ice-associated zooplankton, and pelagic amphipods play unique roles in the production and transfer of energy from sea ice to forage fish and are thus considered keystone species. At the ice edge the food web has an open water component with a period of high localized productivity (fig. 13). When first-year ice

A polynya is a geographically recurrent area of open water or limited sea ice isolated within thicker annual or winter pack ice. Polynyas are ecologically significant in the Arctic because they are generally highly productive and therefore attract upper-trophic-level populations. Phytoplankton production is typically limited in the Arctic to a short period in June, July, or August when diatom blooms occur in response to the ice melt. In Arctic polynyas (e.g., North Water and Northeast Water Polynyas) diatoms bloom in April or May, soon after the end of the polar night, resulting in a significant extension to the phytoplankton growing season. This increased productivity allows zooplankton to thrive, providing prey for higher trophic levels including larval and juvenile Arctic Cod.

Arctic Marine Food Webs



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Humans Sharks e.g., Somniosus microcephalus

Marine Mammals

Reinhardtius hippoglossoides Macrourus berglax

Gaidropsarus sp. Lycodes sp.

Cottunculus microps

Sebastes mentella Boreogadus saida

Antimora rostrata

Triglops nybelini

Bathylagus euryops

Lampanyctus macdonaldi

Liparidae sp.

Hippoglossoides platessoides

Benthosema glaciale Cephalopoda

Copepoda Decapoda

Isopoda

Bivalvia

Amphipoda Euphasiacea

Mysidacea Echinodermata

Polychaeta

Fig. 9. Eastern Arctic marine food web – humans. Example of a marine food web from the eastern Arctic (NAFO Area 0B, Davis Strait), separated by top predator: humans (fig. 9), marine mammals (fig. 10), and Centroscyllium (fig. 11).

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Marine Mammals Ex. Delphinapterus leucas Balaenoptera acutorostrata Phoca groenlandica Cystophora cristata

Reinhardtius hippoglossoides Gaidropsarus sp. Lycodes sp.

Cottunculus microps

Sebastes mentella

Boreogadus saida Antimora rostrata Fig. 10. Eastern Arctic marine food web – marine mammals. Example of a marine food web from the eastern Arctic (NAFO Area 0B, Davis Strait), separated by top predator: humans (fig. 9), marine mammals (fig. 10), and Centroscyllium (fig. 11).

Liparidae sp. Lampanyctus macdonaldi

Benthosema glaciale Cephalopoda

Copepoda Decapoda

Isopoda

Amphipoda Bivalvia

Euphasiacea

Mysidacea Polychaeta

Echinodermata

Centroscyllium fabricii

Artediellus atlanticus

Cottunculus microps

Sebastes mentella

Liparidae sp. Fig. 11. Eastern Arctic marine food web – Centroscyllium. Example of a marine food web from the eastern Arctic (NAFO Area 0B, Davis Strait), separated by top predator: humans (fig. 9), marine mammals (fig. 10), and Centroscyllium (fig. 11).

Benthosema glaciale Cephalopoda Copepoda

Decapoda

Isopoda



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Amphipoda

Bivalvia

Euphasiacea

Mysidacea

Echinodermata

Polychaeta

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Glaucous Gull Beluga

Thick-billed Murre Arctic Char Capelin

Sculpin Squid

Black Guillemot

Arctic Cod

Forage Fish Herring

Least Cisco

Saffron Cod

Arctic Cisco

Whitefish Rainbow Smelt

Red-necked Phalarope

Arctic Tern

Red-throated Loon

Flounder

Amphipods

Copepods

Mysids, Amphipods, Isopods

Mysids

Benthic Amphipods Isopods Polychaetes Molluscs Bacteria Phytoplankton

OCEAN

Terrestrial Organic Matter

RIVER

KEYSTONE SPECIES Keystone species that are important for food-web integrity. Keystone species that, in addition, are important for Inuvialuit subsistence. Indicate predominant energy pathways.

Fig. 12. Western Arctic – estuarine food web. Predator-prey relationships are based on published reports. Arctic Char refers to both Arctic Char and Dolly Varden.

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Polar Bear

Glaucous Gull Beluga Whale

Mergansers

Ringed Seal

King Eiders Scoters Common Eiders

Arctic Cisco

Arctic Cod

Herring

Long-tailed Ducks

Pelagic Ice Amphipods

Scaup

Ice-associated Zooplankton Ice Algae Benthos (Amphipods) Ice Algae (Detritus)

to sea floor KEYSTONE SPECIES Fig. 13. Western Arctic – ice-edge food web. Predatorprey relationships are based on published reports.

Keystone species that are important for food-web integrity. Keystone species that, in addition, are important for Inuvialuit subsistence. Indicate predominant energy pathways.

Polar Bear

Beluga Whale Black Guillemots Arctic Charr

Squid

Gulls

Murres

Capelin

Herring

Jaegers

Loons Ringed Seal

Arctic Cod

Forage Fish

Bearded Seal

Long-tailed Duck

Bowhead Whale Hyperiid Amphipods

Terns

Eider Ducks

Euphausids, Mysids, Hyperiid Amphipods

Fish Larvae, Chaetognaths, Pteropods Sculpins

Arctic Flounder Polychaete Worms

Copepods Shrimps Phytoplankton

Benthic Amphipods

clams

Phytoplankton Detritus

KEYSTONE SPECIES Fig. 14. Western Arctic – marine food web. Predatorprey relationships are based on published reports.

Keystone species that are important for food-web integrity. Keystone species that, in addition, are important for Inuvialuit subsistence. Indicate predominant energy pathways.



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to sea floor

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Fig. 15. Northern Canada.

The Canadian Arctic Ocean The Area The Canadian portion of the Arctic Ocean and adjacent seas, the area considered in this book, encompasses the region from 141° W longitude at the border with Alaska to the international border with Greenland running through the centre of Nares Strait and Baffin Bay south to Davis Strait. Technically Canadian jurisdiction over marine waters extends to 200 nautical miles offshore (i.e., the Exclusive Economic Zone, EEZ), although herein we include western waters outside this limit west to the 141° W longitude north to the pole, as well as eastern waters north of the 200 nautical mile limit east to the 60° W longitude. In practice such distinction is presently

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irrelevant in that virtually no work has been conducted on marine fishes in such offshore areas; thus, few records are known. The adjacent Arctic seas of Canada include the following areas arranged from west to east and then southwards: Beaufort Sea, Lincoln Sea, Kane Basin, Baffin Bay, Davis Strait, northern portions of the Labrador Sea, Hudson Strait, Ungava Bay, Foxe Basin, Hudson Bay, and James Bay. All related connecting waterways such as the channels throughout the Canadian Arctic Archipelago are also included. With the exception of the Beaufort Sea and the archipelago waterways, most of these seas can be considered to be sub-Arctic in nature; that is, they are heavily influenced by temperate environments. The total area of Canadian Arctic seas is about three million square kilometers, that is, about two to three times that of Canada’s

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portions of the 200 nautical mile Exclusive Economic Zone of the Pacific and Atlantic oceans combined. The Arctic coastline is about 173,000 kilometers long, or 71% of Canada’s total coastline, the majority resulting from the numerous islands that make up the Canadian Arctic Archipelago. This area makes up much of the continental shelf present in the Canadian Arctic, which otherwise is relatively narrow (i.e., 100–200 km). Hudson Bay is generally regarded as a shallow inland sea (< 200 m depth) primarily dominated by continental shelf habitat. Northward-flowing rivers drain about 74% of Canada’s land area as well as 148,000 square kilometers of the United States north into the Canadian Arctic marine areas (see fig. 15). About 3.6 million square kilometers (36% of Canada) drain directly into the Arctic Ocean, primarily by the Mackenzie River in the west, and 3.9 million square kilometers (39% of Canada) drain into Hudson and James Bays by the rivers surrounding these areas. High inputs of sediment-laden fresh water (e.g., the Mackenzie River discharges about 333 cubic kilometers of water annually) into the Arctic marine ecosystems have significant effects on the nearshore marine environment. These relatively warm waters create significant opportunities for anadromous and euryhaline marine fishes throughout much of the Canadian Arctic, and such species make up a substantial portion of the Canadian Arctic marine fish fauna. Canadian Arctic marine waters are not homogeneous, due to the following factors: size and geographical scope (particularly latitudinal heterogeneity in driving factors), discontinuous associations between water bodies that are separated by intervening land masses and relatively shallow channels, distinct climatic regimes, and distinct oceanographic situations. In addition, in Canada the geographical distribution of large rivers that drain to Arctic marine areas is heterogeneous. Thus, the greatest influence of these is similarly heterogeneous and is most significant in the Beaufort Sea due to the Mackenzie River, and in Hudson and James Bays due to the following rivers (arranged in order of decreasing mean discharge): Koksoak, Nelson, La Grande, Moose, Churchill, Nottaway, and numerous smaller rivers. Fresh waters have considerable influence on nearshore marine environments, and in Canada much of this results directly from the size of the area drained (and hence water volumes).

Ecoregions in Canadian Arctic Marine Waters Using the physical criteria discussed, as well as general ice and productivity characteristics, the following ecological regions (ecoregions) can be delineated for Canadian Arctic marine waters (fig. 16). In the northwest, outside 200 m depths, the Arctic Basin ecoregion includes the abyssal depths beyond the shelf drop-off. This is an area that likely has low productivity, but it has been little studied ecologically due to the presence of large amounts of polar-pack and multi-year ice. Fishes typical of abyssal and continental slope habitats are likely present here. Onshore from the Arctic Basin in the south is the Beaufort Sea– Amundsen Gulf ecoregion; in the centre of the Canadian Arctic Archipelago is the Viscount Melville ecoregion; and in the north is the High Arctic Archipelago ecoregion. These three differ substantially from each other with respect to overall marine productivity



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and ice characteristics. Limited knowledge regarding marine fishes in these ecoregions results from persistent ice presence and limited research in the area. Productivity is highest in the Beaufort Sea area due to input from the Mackenzie River, longshore currents bringing Pacific water to the area, and upwelling Atlantic water from the Arctic Basin. Marine fish faunas here are dominated by elements with affinities to those of the Pacific Ocean and by anadromous salmonids. Atlantic faunal elements in deeper areas appear to be present based upon recent work. Productivity is likely at intermediate levels in the Viscount Melville ecoregion due to inputs from the Arctic Basin. Multi-year ice cover in much of this area has precluded significant work on marine fishes. Arctic gadids are present, and surrounding land masses likely support anadromous populations of Arctic Char. Productivity is lowest in the High Arctic Archipelago due to a combination of low temperatures, the extensive presence of ice all year, and low inputs from the Arctic Ocean. Little or no work on Arctic marine fishes has been conducted in such areas, although ubiquitous species such as Arctic Cod and sculpins are likely present. Also, although populations of Arctic Char occur throughout the islands of this area of the archipelago, it is unknown whether anadromy is regularly exhibited. The gradation in the productivity of these three ecoregions also parallels air temperatures and the increased duration and surface coverage of both multi-year and seasonal ice further north and east. The two southern ecoregions support substantial populations of anadromous coregonines and chars, whereas it appears that low productivity in the north precludes any advantage of anadromy for chars over much of the area. The Queen Maud Gulf ecoregion is situated along the central southern portion of the archipelago and adjoining mainland. A shallow, shelf-like area (< 100 m depth), it appears to exhibit moderate productivity, especially of anadromous populations of Arctic Char and to a lesser extent anadromous coregonines. A variety of marine fishes are known to occur in this area, but extensive research has not been conducted; thus, information is limited. Eastern areas may remain choked with sea ice during summer. North and east of this ecoregion is the Lancaster Sound ecoregion that occupies the central and eastern portions of the archipelago. It is an area of moderate to high productivity although much of it is heavily influenced by multi-year ice flowing through the channels from the northwest. The eastern end of the region is characterized by the highly productive Lancaster Sound polynya, recurrent leads in the seasonal ice, and local currents from the Labrador Current. As expected, the marine fish species’ composition in this area shares similarities with that in the North Atlantic Ocean. Anadromous populations of char are common in this area. Along the eastern coastline of the archipelago including southern Ellesmere, Devon, and Baffin islands, the Baffin Bay–Davis Strait Nearshore ecoregion can be delineated from the Baffin Bay– Davis Strait Offshore ecoregion. In the north these ecoregions are delineated by recurrent leads in the ice as well as by depth (i.e., usually 200 m). The nearshore zone is characterized by many fiords along the coastline that are separated by shallow water sills from the deeper offshore waters. Southern fiord areas appear to be moderately productive for anadromous populations of Arctic Char as well

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Greenland

Alaska

Baffin Bay

Da vis Str ait

Foxe Basin Hudson Strait Ungava Bay

Ecozone Arctic Basin Atlantic EEZ Baffin Bay - Davis Strait Nearshore

Hudson Bay

Baffin Bay - Davis Strait Offshore Beaufort Sea - Amundsen Gulf Foxe Basin High Arctic Archipelago

Canada

James Bay

Hudson Bay Hudson Strait James Bay - Eastern Hudson Bay Labrador Sea Lancaster Sound Region Pacific EEZ Queen Maud Gulf Viscount Mellville Sound

Fig. 16. Ecoregions of Canadian marine waters. Arctic marine waters are separated into 12 distinct ecoregions.

as populations of marine species such as Greenland Halibut that may be semi-isolated from offshore populations. Fiords in northern areas in this ecoregion likely have lower overall productivity, hence lower species’ diversity. Marine species of fish are likely present but poorly documented. Due to constraints associated with access and potential duration at sea, it is likely that most populations of Arctic Char in northern areas are non-anadromous or exhibit anadromy episodically when summer conditions allow access to the sea. The deep offshore waters of the Baffin Bay–Davis Strait Offshore ecoregion appear to be moderately to highly productive for Arctic waters, and they support populations of economically important marine fishes such as Greenland Halibut. At the southern end of this ecoregion the south-flowing Labrador Current is deflected into the eastern end of Hudson Strait, which likely contributes to very high local marine productivity in this area. As expected, the marine fishes here show great affinities to those of the North Atlantic Ocean. Offshore, especially in eastern Hudson

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Strait, anadromous species such as Atlantic Salmon (Salmo salar) are likely present in surface waters. The Foxe Basin ecoregion is a relatively enclosed area north of Hudson Bay proper. It receives input from the Canadian Arctic Archipelago via Fury and Hecla Strait, and from the Atlantic Ocean indirectly via Hudson Strait. This area is relatively highly productive as evidenced by extensive marine bird populations, but little work has been conducted on the marine fish populations. Populations of Arctic Char in this area likely exhibit anadromy. The Hudson Strait ecoregion is primarily delineated by currents and bathymetric features. Atlantic water inflow occurs into Hudson Bay along the north side of the strait, and outflow occurs primarily along the south side. Likely as a result of these flows, the area appears to be relatively highly productive, supporting anadromous fishes such as Arctic Char in the north and Atlantic Salmon in the south, as well as numerous marine species primarily of Atlantic affinity. Recurrent leads in the area likely result from current action.

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The Hudson Bay ecoregion, occupying the western portion of the bay, is primarily an inland sea, most of which is relatively shallow (< 200 m depth). It is a moderately productive Arctic region that supports anadromous populations of Arctic Char in the northwest as well as whitefishes in the south. Atlantic Salmon populations occur in the northeast portion of this ecoregion along the Québec coast. Numerous marine species are present in this ecoregion but are poorly researched. They are primarily of Atlantic origin, but some Arctic marine and anadromous species are also present. Some anadromous species are also likely secondarily so and derived from freshwater forms from the Mississippian refugium. The eastern and southern portions of this area are distinct and recognized as the James Bay–Eastern Hudson Bay ecoregion. This area is relatively shallow with numerous chains of onshore and offshore islands. Large rivers draining Québec and northeastern Ontario ameliorate the marine conditions to a great degree and likely provide the basis for high local productivity especially in estuarine and nearshore areas. This productivity is particuarly relevant for anadromous fishes such as whitefishes and Brook Trout (Salvelinus fontinalis). Extensive shore leads occur along the coast in several areas. Marine fishes in this area are similar to those of the Hudson Bay ecoregion generally. The northern portion of the Labrador Sea is Arctic in nature and is considered herein. The area is characterized by warmer surface waters than those of the previous Arctic ecoregions and thus has relatively high inshore and offshore productivity. As a result, this region supports substantial populations of anadromous fishes such as Atlantic Salmon, as well as both nearshore and offshore marine fishes that are characteristic of the North Atlantic Ocean. As noted, substantial research has not been conducted on many of the ecoregions defined here. This is especially true for their fish faunas. With the exception of a few species that are important in fisheries or are locally abundant, generally marine fish faunas throughout the Canadian Arctic are poorly understood; thus, the above conclusions must be considered to be generalities only.

Summary Despite the environmental extremes present in the Arctic, much of the marine environment is hospitable to a number of wholly marine and anadromous fish species. Although spatially highly



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heterogeneous, the productivity of many areas of the Canadian Arctic marine environment is moderately high for a polar environment. Consequently the diversity and abundance of marine and anadromous fish populations is high in some areas. Many of these fish populations, especially those based on anadromous fish, support important local fisheries (see the “Fisheries” section in the introduction). Other species, such as Arctic Cod, are pivotal in Arctic food webs and are therefore central to the marine ecology of the area. As noted in several places in the foregoing, the Arctic is characterized both by high variability associated with high seasonality and by relatively constant but slow rates of change (at least until the last few decades). Relative stability has yet to be achieved in the present postglacial period. Thus, the physical, chemical, and to some degree biotic settings described above represent conditions that characterized the twentieth century up to about 1990. Since this time the effects of human-induced change have become apparent, manifesting primarily as climate change that has had profound effects on Arctic marine systems (see the “Climate” section in the introduction). Anthropogenic changes are superimposed upon the ongoing postglacial ecological equilibriations. The cascading consequences of change in the physical and chemical components of the ecosystems will affect the marine fish biota. Moreover, secondary consequences of climate change that generally allow for increased human activities in the Arctic will further affect Arctic marine fishes and their ecosystems.

sources:Dunbar (1968); McIntyre et al. (1976); DeVries (1977);

McAllister (1977); Menzies (1977); Stirling & Cleator (1981); Hocutt & Wiley (1986); Martini (1986); Aagaard & Carmack (1989); Parsons et al. (1989); Smith (1990a, 1990b); Harwood & Stirling (1992); Hsiao (1992); H.E. Welch et al. (1992); Clarke (1993); Michaud, Fortier, Rowe, & Ramseier (1996); Melnikov (1997); Kosobokova, Hanssen, Hirche, & Knickmeier (1998); Wadhams (2000); Tremblay, Gratton, Fauchot, & Price (2002); Arrigo & van Dijken (2004); Dick & Chambers (2005); Iken, Bluhm, & Gradinger (2005); Mincks, Smith, & DeMaster (2005); Carmack & Wassman (2006); Dunton, Weingartner, & Carmack (2006); Vander Zanden & Fetzer (2007); Cobb et al. (2008); Renaud, Morata, Carroll, Denisenko, & Reigstad (2008); Hannah, Dupont, & Dunphy (2009); Loseto, Stern, & Ferguson (2009); Darnis et al. (2012); Kędra, Kuliński, Walkusz, & Legeżyńska (2012); Boetius et al. (2013); Christiansen & Reist (2013); Ji, Jin, & Varpe (2013); Mathias (2013); Jakobsson et al. (2015); V. Roy et al. (2015).

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HISTORY OF RESEARCH James D. Reist

Arctic ichthyology as a distinct subdiscipline can be traced to the Danish priest Otto Fabricius who was the first to publish a scientific account of forty-five fish species from west Greenland in Fauna Groenlandica (1780), although some descriptions had been published earlier by Otto F. Müller in Zoologiae Danicae Prodomus (1776). With the exception of western Greenland (of relevance to the Baffin Bay–Davis Strait area), early focal areas for Arctic ichthyology from the eighteenth century to the first half of the twentieth century were mostly on European and Siberian waters. Pennant, however, in 1785 included an account of some fishes from Hudson Bay in his Arctic Zoology. Some species described during this period are distributed widely in Arctic marine waters, thus have relevance in our discussion (see the authors credited with the scientific nomenclature of many individual species in the “Family and Species Accounts” section). Earlier, as well as during this period, the European focus with respect to the North American Arctic generally, and Canada in particular, was on exploration of the north principally for navigation routes to eastern Asia. Accordingly, little other than casual mention of local fishes is documented in the many voyage accounts. Authors of eighteenth- and nineteenth-century works that mention fishes in Arctic Canada are listed in the bibliography and include Forster and Pennant from the eighteenth century, and Bean, Fielden, Gordon, Gray, Günther, Halkett, Holmqvist, Low, Prince, Richardson (see figs. 17 and 18), Ross, Sabine, Wakeham, and Walker from the nineteenth century. In addition, synopses of northern explorations from early history to 1962 can be seen online (www.nauticapedia.ca). European exploration of the Canadian Arctic began in 1576 with Martin Frobisher’s first expedition to Baffin Island. The ensuing 441 years have resulted in the development of a knowledge base regarding marine fishes, their habitats and ecology, and the ecosystems supporting them, one that is biased towards the southern Arctic waters and coastal areas that typically become free of sea ice in summer. Thus, significant gaps remain, particularly in ice-covered and high Arctic areas; knowledge development to address these is an ongoing activity, albeit one executed slowly due to logistical and cost constraints. Throughout the period, several themes have

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Fig. 17. Stenodus leucichthys after Richardson (1836a).

Fig. 18. Gymnelus viridis after Richardson (1855)

dominated knowledge acquisition for the Canadian Arctic generally and marine fishes particularly. Although these themes are not mutually exclusive and overlap each other, particular themes have dominated at one time or another over the centuries. This dominance, in turn, has affected the focus, coverage, and delivery of knowledge regarding the taxonomy and distribution of Arctic marine fishes. In rough chronological order of dominance the themes are (1) the exploration, discovery, and development of a potential trade route connecting Europe to eastern Asia (1576 to 1880s, and renewed in recent years due to sea-ice loss); (2) whaling and the fur

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trade (1800 to ca. 1940); (3) geographical exploration, sovereignty, and military security (1900 to 2000s); (4) renewable resource-based economic development including fisheries (1950s to the present) and relevant to both the sustenance of Indigenous peoples and commercial possibilities; (5) non-renewable, resource-based economic development (i.e., hydroelectric development of northern rivers, mining, shipping, oil and gas development; 1970s to the present); and (6) ecosystem integrity, biodiversity, and conservation (late 1990s to the present). Overarching the last three topics, particularly in the past two decades, have been the notable changes in the Arctic driven by climate change and associated changes in the physical and biotic marine environments. European vessels (e.g., the voyages of Martin Frobisher in 1576– 1578, John Davis in 1585, 1586, and 1587, and Robert Bylot in 1616) chartered by English economic associations explored Davis Strait and Baffin Bay (north to 77o45') and the outer edges of the adjacent sounds to assess the potential for mines and to explore the route to Cathay (China). Small vessel size and heavy ice thwarted much exploration other than that of adjacent coasts. Moreover, the vessels were not equipped for fishing and did not carry able naturalists (these were to come much later); thus, the knowledge developed regarding fishes and their ecosystems was limited and restricted to that necessary for navigation and survival. Limited incidental harvesting of coastal and anadromous fishes likely occurred to replenish supplies; however, little note of this exists in journals. Whaling in Baffin Bay and Davis Strait was developed over the seventeenth and eighteenth centuries, but little further exploration occurred in the area during this period. It was not until the end of the Napoleonic wars (1815) that the attention of the British Admiralty returned to the northern New World. A large naval force with professional officers, combined with a period of local climate warming (hence diminished summer sea ice), saw the initiation of a sixty-year period of government-sanctioned explorations (e.g., John Ross, 1817–18, to Melville Bay and the northern polynya; Edward Parry, 1819–20, to Lancaster Sound west to Melville Island; John Ross, 1829–30, to Somerset Island). The middle of the century (about 1845) saw the John Franklin voyage go missing, followed by numerous search and relief voyages from both the east and the west (Beaufort Sea), which, together with previous overland expeditions to the Arctic coast, documented the geography. Exploration and mapping of the Northwest Passage were primary objectives; however, this period also saw the addition of naturalists to ships’ complements. Physical oceanography (e.g., tides, depth soundings, currents, ice conditions), along with biological oceanography such as tows and capture of marine invertebrates, dominated the science conducted. Thus, in addition to obtaining fishes for provisions and overwintering (e.g., Inuit provided fish to John Ross during his 1830 overwintering), scientific advances were primarily in the physical sciences, although some journals document fishes. Most of these studies focused on coastal fishes and anadromous species in both the sea and the adjoining fresh waters. Fishes were collected by some thirteen Arctic expeditions between 1819 and 1855, and many of these collections were summarized through the works of J. Richardson (Fauna Boreali-Americana, part 3, “The Fish”), J.C. Ross,



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and others. British vessels dominated the exploration during the early part of this period; however, the ships of other countries explored much of the area later (e.g., several American expeditions began in about 1855, and Norwegians such as Sverdrup and Amundsen conducted several expeditions in the archipelago from 1893 to 1925 aboard the Fram, Gjoa, and Maud). These were primarily explorations to map the archipelago, but significant scientific work was also conducted, especially during overwinter periods; fishing for provisions likely also occurred but is little noted. Heavy sea ice in the middle and latter parts of the nineteenth century impeded wider activity in the southern archipelago. (Marine transit of the Northwest Passage finally occurred in 1906 when Roald Amundsen completed the journey in the Gjoa.) In the west, exploration, whaling, and the search for the missing Franklin expedition occurred in the Beaufort Sea and western archipelago throughout much of this same period. In 1880, Britain ceded sovereignty over the Canadian Arctic Archipelago to Canada, and the new nation of Canada was not to be outdone. Sovereignty activities were initiated in the Arctic through several expeditions by CGS Arctic commanded by Captain J.-E. Bernier (1906 to early 1920s), the DGS Neptune expedition in the eastern Arctic (1903–4), voyages by V. Stefansson, and the Canadian Arctic Expedition (1913–16), many under the auspices of the Geological Survey (later to become Geological and Natural History Survey of Canada). Much of this effort, however, was focused on freshwater fishes. In the south, exploration of Hudson Bay was initiated in 1610 by Henry Hudson seeking a southern route westwards to Cathay. Several unsuccessful expeditions ensued, and further efforts were abandoned until the Nonsuch returned a load of furs to Europe (1668–9). The formation of the Hudson’s Bay Company in 1670 established the fur trade in the area, the development of coastal fortified trading posts, and annual visits by vessels. Coincident with the marine explorations of the New World and the Arctic expeditions noted above, land exploration and the development of the inland fur trade were occurring. These included overland expeditions to the northern coast (e.g., Samuel Hearne to the Coppermine River area in 1769–72, and Alexander Mackenzie to the mouth of the Mackenzie River in 1789). Mapping and claims to territory were the primary objectives, intertwined with economic reasons; however, scientific activities such as describing local fauna also occurred. Fishing contributed to local provisioning, and anadromous fishes undoubtedly were being captured in Arctic rivers and coastal areas. To facilitate the transport of furs to European markets, both the Hudson’s Bay Company and its rival, the North West Company, established a wide range of forts and trading posts throughout the Arctic over the seventeenth and eighteenth centuries; many of these were near the mouths of Arctic rivers and were serviced by annual re-supply voyages. The exploitation of fishes, particularly autumn runs of anadromous species, was widespread for local sustenance. The addition of naturalists to either the expeditions or the complements of the trading posts facilitated work on biota including fishes. For example, in 1773 one Mr John Reinhold Forster provided to Th. Pennant an account of fishes from Hudson Bay, the descriptions of which he received from a “naturalist” at York Fort on western

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Fig. 19. Myoxocephalus scorpius, CMNFI 1958-0258. Watercolour drawn by J. Dewey Soper.

Fig. 20. Boreogadus saida. Watercolour drawn by J. Dewey Soper.

Hudson Bay. This work contains a clear account of Lake Sturgeon, an anadromous species in the area, among accounts of freshwater fishes. Similarly, Pennant’s account in 1792 of Monkfish (Lophius) from this area is based upon a manuscript by Th. Hutchins, a Hudson Bay factor, written in 1782. Given the exploratory focus of both land-based and sea-based expeditions and the rigours of survival, the accounts of marine fishes during this period focused heavily upon the coastal and anadromous species relevant to provisioning, were sporadic, and depended upon the interest and availability of an appropriate naturalist. Unique individuals with a natural history interest in the north combined with opportunity have sporadically contributed to the knowledge of marine fishes. For example, the U.S. Army Signal Service established L.M. Turner in Fort Chimo (now Kuujjuaq, northern Québec) to take meteorological and geophysical observations during the 1882–4 first International Polar Year. Mr Turner’s natural history interests contributed to collections of biota including fishes from Ungava Bay; his specimens and unpublished notes are deposited in the Smithsonian Institution (Washington, DC). He documents the first occurrence of Capelin in that area and, interestingly, also combines scientific observation with local knowledge of the fishermen and traditional knowledge of the Indigenous peoples regarding the distribution and dynamics of coastal marine fishes in the area. These specimens aided the later development of the first compendium of marine fishes for this region (see references in Vladykov, 1933a). Knowledge of new fishes was often incorporated into the Eurocentric taxonomy of the day; thus, where similarities existed, North American species were equated with their European counterparts during this period. As knowledge of North American taxonomy grew and specific studies were completed, the early synonymies were resolved, and descriptions of new species progressed. For example, The Fishes of North and Middle America: A Descriptive Catalogue of the Species of Fish-Like Vertebrates in the Waters of North America, North of the Isthmus of Panama by D.S. Jordan and B.W. Evermann (Bulletin 47, U.S. National Museum, 1896–1900) is a comprehensive

compendium and realignment of fish taxonomy. The focus of much of this early work was on freshwater and anadromous species, but some research on Arctic marine fishes occurred during the period. From early 1900 onwards, fur trading waned, but northern discovery and development continued through a variety of means. The Northwest Mounted Police (later the Royal Canadian Mounted Police, RCMP) and missionaries were established in trading centres throughout the north. Overland and marine explorations by the RCMP, and the occasional naturalist travelling the area, contributed to the development of knowledge regarding marine fishes and other biota. For example, RCMP and missionary records from the western Arctic (Aklavik) provide documentation of anadromous fish catches of Pacific salmon, whitefishes, and chars in the Mackenzie Delta area beginning about 1912. This same period also marks the initiation of expeditions with distinct natural history themes organized by the Canadian government. The Canadian Arctic Expedition (1913–18) had three geographic focal areas (west, inland, and east) with a major component focused on natural history and biota. Extensive marine work was done, specimens collected, and scientific publications developed. Fishes were observed and collected, but summaries of that research were not published, although incidental observations exist in other literature of the period. The fish specimens, however, were deposited in the national museum collections of the day and form the basis for much of this book. Additional collections were made in the 1920s, and, in 1924–6 for example, the National Museum of Canada hired J. Dewey Soper for an expedition to Baffin Island. Some of the fish observed were collected and deposited in what is now the Canadian Museum of Nature (see the “Illustration Credits” section). In addition to extensive mapping and zoological investigations on land and in fresh water, Soper collected marine fishes (e.g., Arctic Cod, Sticklebacks, Sculpins, Arctic Char; figs. 19 and 20). During the early to mid-1900s the initial delineation of three groups of researchers took place, each with slightly differing mandates, which persist today (and among which the research directions have widened somewhat). These groups consist of the

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following: university researchers who are concerned most with developing basic knowledge of the systems; government researchers who are concerned with developing and applying knowledge regarding fisheries, ecosystem conservation, and economic development; and museum researchers who are concerned primarily with taxonomy and with understanding and documenting diversity of biota. Although now these groups are mostly distinct and reasonably delineated by discipline and organizational association, in the early days their roles and interests overlapped. Moreover, the rigours, costs, and threats regarding research in the Arctic continue. Therefore, cross-disciplinary work among these groups remains the norm with regard to the knowledge development of Arctic marine fishes and their ecosystems. Recognition of the essentially unknown state of knowledge regarding fishery issues and potential prompted the government of the day to establish a board of management in 1898, which evolved into the Biological Board of Canada in 1912, and thence to the Fisheries Research Board of Canada (FRBC) in 1937. Two themes governed the operation of the board: (1) original research on aquatic biota, and (2) the problems of management, processing, and marketing faced by governments and industry. Early staffing of these arm’s-length government organizations (1898–1934) incorporated primarily university professors. A growing dichotomy between basic and applied research resulted in formal staffing of the research boards by government scientists from around 1934 onwards. The Fisheries Research Board persisted as a distinct entity until its integration within the Department of Environment in 1971 as a research component, Fisheries and Marine Services; the latter moved to a separate Department of Fisheries in 1973 and became the Department of Fisheries and Oceans (DFO) in 1979. Thus, depending upon the time frame, since 1900 the research regarding Arctic marine fishes has been conducted variously by university professors, scientists within the FRBC or its antecedents, groups within the DFO, and individuals associated with museums of natural history. In the late 1960s FRBC scientists conceived a series of regional works to describe the national fish fauna of Canada, extending and complementing existing regional works. With the exception of the fauna of the Arctic marine waters, these were variously delivered over the next two decades (e.g., A.H. Liem and W.B. Scott, 1966, Fishes of the Atlantic Coast of Canada, updated as W.B. Scott and M.G. Scott, 1988, Atlantic Fishes of Canada; J.L. Hart, 1973, Pacific Fishes of Canada; J.D. McPhail and C.C. Lindsey, 1970, Freshwater Fishes of Northwestern Canada and Alaska; and W.B. Scott and E.J. Crossman, 1973, Freshwater Fishes of Canada). Of course, many of the northern species described in each of these compendia occur in our area of interest and should be consulted for additional information. A companion volume regarding the marine fishes of Arctic Canada became the labour of love for D.E. McAllister of the Canadian Museum of Nature, to whom this book is dedicated. McAllister’s efforts were constrained by academic issues such as limited databases and the absence of comprehensive keys to Arctic fishes and by taxonomic issues associated with some groups – areas in which he was able to contribute significantly. Additionally, the perennial problems of limited sampling and geographic coverage, high costs of field



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research, sea-ice presence over extensive areas, and focal themes for research other than fish taxonomy and distribution all limited knowledge development. Although ameliorated recently to some extent, these constraints continue to limit the knowledge development for Arctic marine fishes. The large geographical areas, complex and highly variable marine systems, inhospitable climate, and presence of extensive sea ice throughout much of the year have all influenced the development of knowledge regarding Arctic marine fishes. Four general areas can be delineated, each with its separate history of work: (1) eastern Arctic, (2) Hudson Bay and associated areas, (3) Canadian Arctic Archipelago, and (4) western Arctic. Exploration and knowledge development have generally progressed from east to west and south to north, albeit with geographical overlap during various periods. In any event, both environmental conditions (particularly the persistence of summer sea ice) and the nature of the regional issues requiring research have affected the nature and focus of research on Arctic marine fishes since the end of the Second World War. It should be emphasized that, in the period from the beginning of the twentieth century to the end of the First World War, official domestic interest in Canada’s Arctic was mostly from the perspective of establishing sovereignty through geographical explorations. Following the Second World War and into the Cold War period to the late 1970s, the strategic importance of the Arctic for continental defence was the overarching interest, which focused mostly on oceanographic expeditions to determine the navigability of Arctic waters (and occasionally the collection of the marine biota encountered). Generally, natural history themes, particularly those profiling fishes, were very limited in scope and “official” interest. Some notable exceptions, however, occurred particularly where biology intersected defence or local development themes.

Eastern Arctic and Hudson Bay areas Based upon material in various collections (1919–30 from coastal work and including marine sampling from several expeditions, e.g., the 1930 expedition aboard the SS Loubyrne to Hudson Bay that was sponsored by the government) and the availability of a reasonable literature, J.R. Dymond (University of Toronto) and V.D. Vladykov (Biological Board of Canada) provided two synopses of the fishes of Hudson Bay in 1933. In 1947 the FRBC developed a research program that was delivered mainly by university staff, particularly Max Dunbar, a professor at McGill University, who focused upon eastern Arctic investigations until 1955 when this evolved to the Arctic Unit. In 1964 the unit was renamed the Arctic Biological Station of the FRBC (subsequently of the DFO) and housed in Sainte-Annede-Bellevue near Montreal. Two mandates initially drove the work: the physical and biological oceanography of the eastern Arctic, and the development of marine resources in the interests of Indigenous peoples. Both the scope and the geographical mandate evolved to include all fishes, fishery resources, and biological oceanography in the Canadian north, and marine mammals on all coasts. Accordingly, multiple research expeditions investigated both anadromous

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Fig. 21. MV Calanus contending with ice en route to the Belcher Islands, 12 July 1959.

and nearshore marine fishes across the Arctic (e.g., Mackenzie Delta and western coast, Coppermine River, lakes on the Back River, Rowley River). The oceanographic theme was (and mostly continues to be) generally delivered through ice-capable vessels that are not specifically fitted to conduct fisheries research. Recognizing the need for a vessel specific to conducting marine fish sampling as well as oceanography, Dunbar directed the building of the MV Calanus in Nova Scotia in 1948, and it became the base for work in the Hudson Bay area and the eastern Arctic in the 1950s to the early 1970s (fig. 21). The work included physical and biological oceanography, fisheries research, and research regarding other Arctic renewable resources (e.g., shellfish, marine mammals), particularly in northern and western Hudson Bay and Foxe Basin, Hudson Strait, and Ungava Bay. As the MV Calanus, like all before her, was an ice-strengthened rather than an ice-breaking vessel, the continued heavy presence of summer sea ice still precluded access to many areas and severely limited seasonal work. Fish research included Arctic Char, Arctic (Polar) Cod, Greenland Shark, Greenland Halibut, and, to some extent, Atlantic Cod (Gadus morhua). The origins of ecosystem research in the Canadian Arctic can be traced to these efforts. There was significant research regarding pelagic (e.g., Calanus spp., copepods) and benthic (e.g., annelids, echinoderms) invertebrates and fishes, as well as marine mammals. Moreover, in the resulting scientific literature can be seen initial understanding of Arctic fish fauna

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and marine systems in comparison to sub-Arctic or boreal systems, of the origins and associations of the Arctic fishes with temperate faunas, and of marine fishes as integral parts of the larger ecosystem. Work in Greenlandic marine waters by Danish researchers during the early-twentieth century also provided information about the fauna adjacent to that of the Canadian eastern Arctic, including A. Jensen (1926, List of the Fishes of Greenland) and J.R. Pfaff (1940, “Fishes,” in Report of the Fifth Thule Expedition, 1921–1924).

Western Arctic Although much of the work in the west from 1944 onwards was concentrated on freshwater fishes and ecology, Arctic Unit staff, FRBC staff, and university associates continued research here on marine fishes and oceanography throughout the 1950s (e.g., CGMV Cancolim II, a Defence Research Board ship, worked in the Beaufort Sea in 1951 and enabled specific research on nearshore fishes in the Herschel, Holman, and Tuktoyaktuk areas). Samples, expedition reports, and data collected by these initiatives formed the basis for follow-on research, particularly with faunal collections deposited in museums. The data points noted from literature and deposited specimens in natural history museums form the basis for the mapped distributions shown herein. These show the importance of

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Fig. 22. MV Salvelinus, icebound in Starvation Cove on the south shore of Victoria Island, 23 August 1968.

such collections and archival efforts with regard to newly explored areas and poorly known faunas (e.g., the unrecognized diversity present in the form of species new to science), in particular. The evolution of research activities from earlier times to this period of interest involving the Defence Research Board (formed in 1947 as an offshoot of the National Research Council of Canada) primarily focused upon physical, chemical, and biological oceanography with limited (but at times focused) sampling and research concerning marine fishes specifically. Ultimately, government oceanographic research in the western Arctic became centred at research laboratories on the Pacific coast (i.e., Institute of Ocean Sciences) and that in the eastern Arctic centred on the Atlantic coast (i.e., Bedford Institute of Oceanography). Within the DFO, however, biological research themes, particularly those regarding fishes, have expanded and are concentrated in Winnipeg (see below). From 1959 to about 1975 the FRBC’s Arctic Biological Station operated a second vessel, MV Salvelinus, in the western Arctic to develop understanding of the Beaufort Sea and adjacent areas east to Cambridge Bay (fig. 22). Work continued upon focal species such as Arctic Char, coastal coregonines, Arctic Cod, and Greenland Cod (Gadus ogac). Again, sea ice and logistics were the major limiting factors in the research that was confined mostly to the southern Canadian Beaufort Sea and the seasonally ice-free nearshore corridors in the southern archipelago.



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Consolidation and Centralization of Government Activities The FRBC essentially controlled the nature and direction of research with regard to Arctic fishes and fisheries. Separate government units were responsible for the management of fishery resources – such as Fisheries and Marine Service of the Department of Environment – which, for the north, relied mostly upon research produced by FRBC university staff. However, in 1973 the FRBC was integrated as a research component, in an advisory capacity, into the federal government’s Department of Fisheries. In the 1980s the decision was made to close the Arctic Biological Station and consolidate both fishery management and research activities for the Arctic at the Freshwater Institute in Winnipeg (freshwater fish and fishery research and management initiatives had had a checkered history but were consolidated in this institute in the early 1970s). Thus, the primary location for government-based research on fishes in the Arctic (defined as including areas north of 60o N latitude, i.e., anadromous, marine, and freshwater fishes and fisheries in the then Northwest Territories and later coastal Yukon, the Northwest Territories, and Nunavut) became the Winnipeg laboratory, and stock assessment, environmental impacts, and fishery management responsibilities were separated into a unit distinct from that of research activities. The staff assessing stocks and those

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working on environmental fishery-management activities derived originally from government units within the Fisheries and Marine Services, whereas scientific staff investigating northern fishes came from the closure of the Sainte-Anne-de-Bellevue laboratory and during the 1980s were supplemented with new research staff and focused research themes. In the western Arctic, management and research themes focused initially upon freshwater and coastal anadromous fishes; this was overlaid by significant interest, continuing into the late 1980s, in the development of coastal oil reserves in Alaska (Prudhoe Bay) and gas reserves in the nearshore of the southern Canadian Beaufort Sea. Accordingly, in the west, research into coastal marine fishes and anadromous fishes in inland waters was conducted under three large sequential government programs (Arctic Gas Studies in the 1970s, the Beaufort Sea Project from 1973 to 1975, and the Northern Oil and Gas Action Program in the 1980s). These programs established marine distributions and ecology for key species as baselines for assessing the potential impacts of oil and gas development (which at that time included nearshore artificial islands and caissons as drilling platforms, and plans for both landbased and sub-surface sea pipelines). For example, reports from the Beaufort Sea Project note 300 fish collection lots in the three project years, compared to 450 collection lots from seventeen studies during the previous twenty years. Bottom and mid-water trawling along the Mackenzie shelf by RV Pandora and MV Theta occurred in 1974 and 1975. Coverage, however, was limited by gear (including ship size) and sea-ice conditions. These studies also marked the move from the primarily strategic drivers that had dominated overall research agendas prior to 1970 towards environmental perturbations resulting from industrial development. As noted, during the initial period of government activities with regard to fisheries, research and fishery development were two parts of the FRBC work. The third part, that of fishery management, was performed separately within government, which simultaneously conducted stock-assessment activities. The 1973 consolidation of FRBC with the Department of Fisheries ended that separation. Within the department, however, relative to the Arctic, the separation of stock-assessment activities from fish and fishery research activities continued for some years; it came to an end in the mid1980s with the formal de-linking of fishery management sectors from the science research and stock-assessment sector. During this period, research activities in the east continued and focused primarily upon anadromous fishes, especially Arctic Char. They also included basic work regarding marine species of potential fishery importance (i.e., Greenland Halibut). In both the western and the eastern Arctic, though, there was limited interest from university-based researchers in Arctic fishes, with most of the effort being conducted by government research groups in association with large-scale northern development programs. This was undoubtedly driven by funding limitations that affected the scope of university research. Towards the end of this time frame, in 1987, a new DFO facility, the Maurice Lamontagne Institute, was built in Mont-Joli, Québec, and elements of Fishery and Marine Services staff that were oriented towards issues in Québec, and some from Sainte-Anne-de-Bellevue,

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moved there. From the late 1970s to the 1980s, marine fishery research activities had focused upon Hudson and James Bays. A driving factor for fish research in marine systems was the hydro-electric development on the rivers of northern Québec (e.g., La Grande, Great Whale) with a focus upon anadromous species in the area (i.e., coregonines), nearshore marine ecology of Arctic Cod, and the potential effects of altered freshwater inputs upon the marine system. Unlike in the situations further north, university researchers and graduate students were more directly involved in the programs, and a number of publications resulted. Similar work was conducted on the estuaries of western rivers (e.g., Nelson, Churchill) affected by hydro-electric development in Manitoba, primarily through consultants to the industry. Access to the Arctic – especially that associated with suitably equipped marine fishery research vessels – funding (or lack thereof), and demand for knowledge, particularly with respect to the environmental impacts of large-scale developments, have consistently guided the nature and scope of government-based research regarding fishes in the Arctic. The limiting factor has always been the costs associated with such work, especially those involving vessels. In the 1990s, interest and funding waned due to global economics, particularly those associated with oil and gas development in the west. With respect to marine fishes, this hiatus lasted most of the decade at least in the western Arctic and Hudson and James Bays. The exception was the continuation of a low level of activity with regard to anadromous fishes, primarily in support of stock assessment and management decisions. In the east, activities for both marine and anadromous fishes continued to support management needs. Concerning marine fishes, however, a developing interest in commercial exploitation, particularly of Greenland Halibut, stimulated interest in stock-assessment activities. In the 2000s, research generally into Arctic marine systems experienced an upsurge, driven in part by fishery-development initiatives from land-claim organizations; climate-change effects on marine systems, which allowed greater access to remote areas; the International Polar Year (2007–8); and increased northern economic development. This has involved loose coalitions of researchers from government, universities, and occasionally museums, focusing primarily on systems research, as follows.

Canadian Museum of Nature and Fisheries Research Board of Canada (1970s to 1990s) In the late 1960s the FRBC had intended to produce comprehensive guides to the fishes present in all regions of Canada by enlisting expert contributions from university staff, museum ichthyologists, and government scientists. The intended guide for Arctic marine fishes proved to be an impossible undertaking due to the dearth of samples, the lack of comprehensive geographical coverage, and logistical considerations. Nevertheless, efforts to develop such a compendium were made by Don McAllister, newly hired as curator of fishes at the National Museum of Natural Sciences, and Jerry Hunter, a scientist at the Arctic Biological Station. Throughout the period from 1958 to his death in 2001, Don vigorously acquired specimens, developed keys and distributional treatises, described some species, and conducted primary research on various Arctic marine fishes. As an example of

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these efforts, in 1990 some 137 species were documented as occurring in Canadian Arctic marine waters, of which 95 were represented in museum collections (58,336 specimens from 1,737 stations). These were biased towards coastal sampling (e.g., summer ice-free zones), primarily towards southerly locations in the Arctic and towards the more accessible areas of Davis Strait and Baffin Bay, biases that remain to this day. At that time Don highlighted several high-priority issues: (1) collecting specimens of the unrepresented species, (2) collecting deep-water and mid-water fishes, especially in the northwestern Arctic, (3) collecting in the south-central and northern archipelago waters, and (4) making winter collections. The work of Don and his contemporary colleagues laid the foundation for the present synopsis and in effect has guided some aspects of research on marine fishes. Interestingly, the second, third, and fourth topics remain as issues; however, work to fill those gaps in the intervening time to the present constitutes the final period of consideration. Two other considerations are relevant in the context of the role of museums and their scientists (i.e., taxonomists and systematists). First, it should be noted that fish specimens were collected by a number of expeditions throughout much of the time period involved, and these and related capture data found their way to natural history collections and museums, including several in North America (e.g., Royal Ontario Museum, Smithsonian Institution). In the case of the eastern Arctic investigations noted earlier, many of those specimens were initially housed at the Sainte-Anne-de-Bellevue laboratory, and the majority were transferred later to the Canadian Museum of Nature. The second contextual consideration is associated with a general orientation of the nature of natural history science and its evolution over time. Cataloguing the diversity of life and organizing it systematically were dominant themes of natural history science from the seventeenth to the early-twentieth century. Accordingly, the exploration of new lands incorporated these activities. In the south, as biota and their distributions became better known, taxonomy and distributional studies gave way to studies emphasizing ecology and systems processes. In the context of fishes, this meant an increasing emphasis on the application of knowledge to exploitable populations and fisheries management. With the publication of comprehensive guides to fishes being available in most areas of the country by the late 1980s, the perception was that most of the inventorying of fish diversity had been completed. This, and shifts in the popularity of taxonomy as a discipline, contributed to a move away from comprehensive documentation of diversity. In the north, however, due to the perennial and continuing limitations on access and sampling (i.e., funding and costs, institutional will, field logistical limitations such as ice-capable fishing vessels, persistent sea-ice presence in many areas), the full development of inventorying and documentation of fish diversity remains to be completed. Recently, as a result of the awareness and concern over biodiversity loss, interest has increased in the basic pursuits of biodiversity science such as taxonomic and distributional studies. A perennial issue that continues to this day is the under-appreciation by field workers of the importance of preserving voucher specimens and having identities verified by qualified experts (e.g., see many of the comments in the “Family and Species Accounts” section).



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Land Claims, Changing Focal Roles of Government, Climate Change, and Northern Development The period from about 1970 to the present has been one of great change in northern Canada, the Arctic in particular, and especially the ecosystems involved. This has involved the settlement of comprehensive land claims with Indigenous peoples that now cover much of the Arctic marine waters (i.e., only the coastal waters of western and southern Hudson and James Bays that are adjacent to the provinces of Manitoba, Ontario, and Québec are excluded). An outcome of this process has been shifts in managerial control of harvestable renewable resources, development of co-management regimes, and increasing demand for scientific information to support management decisions. Throughout the same period the focal roles of government have also shifted (e.g., the dissolution of FRBC, and the shift of research and management roles primarily towards government needs that are explicitly associated with stock assessment, ecosystems management, and the impacts of large-scale developments). Superimposed upon these changes, the effects of climate change, which are amplified in the north, have taken hold and resulted in major environmental changes, particularly since the late 1990s. In the marine environment these include changes in sea ice (e.g., multi-year dominance giving way to younger ice, diminishment of areal coverage especially in late summer, earlier break-up and later freeze-up); changes in the nature and delivery of fresh water to the marine system, with subsequent cascading effects in the sea; alteration of productivity regimes in many areas from an ice-dominated benthic system to more of an open-water-dominated pelagic system; and warming and acidification of marine waters. At the ecosystem level these changes appear to be factors limiting the present distributions of many sub-Arctic fishes; thus, shifts in species composition and distributions may be a significant future consequence. Arguably the greatest consequence of climate change in the context of humans and their activities is the altered access in the Arctic caused by its effects on the various components of the cryosphere (i.e., the frozen system). For example, with the diminished presence of summer sea ice, more areas of the marine system are now accessible both spatially and seasonally. Over the past few decades this has increased interest in northern development, especially that associated with renewable resources (i.e., fisheries), non-renewable resources (e.g., mining and associated marine transport, oil and gas and hydro-electric development), and marine system usage (e.g., destinational and transpolar shipping, development of protected areas). All these factors have contributed over the last few decades to a general stimulus of national interest and activity in the Canadian Arctic. Both university and government scientists have been involved in this activity, much of which has concerned physical, chemical, and biological oceanography, hydrography, and an understanding of the fundamental changes of the ecosystems and their overall consequences to regional biota. Recent efforts in some regions, however, have focused specifically upon anadromous and marine fishes and are summarized below. With few exceptions, most of the programs noted were designed to understand the biology of the fish species, rather than to study fish taxonomy, biodiversity,

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or distribution. With that bias in mind, once the accuracy of taxonomic identification has been verified, the outputs from these and similar studies do provide occurrence information that is useful for determining distributions. Distributional points from many of these studies have been included in a separate database that forms the basis for point distributions on the species maps. In some cases, specimens (e.g., marine collections of fishes including those new to regional faunas or to Canadian waters) have also been deposited in relevant museum collections and contribute directly to this present work.

Hydro-electric work in eastern Hudson and James Bays (1970s to 1980s) and western Hudson Bay (1980s to 1990s) Anadromous and nearshore estuarine and marine fishes were the focus of this work, and several publications in the primary literature resulted; additional reports are in the grey literature. Nearshore oil and gas development in the western Arctic (1970s to early 1990s) Most of the findings from the Beaufort Sea Project and the Northern Oil and Gas Action Program work on anadromous and coastal marine fishes on the Beaufort shelf were documented in a number of government, DFO, and industry reports. In the 2000s, renewed interest regarding the Mackenzie gas pipeline along the Mackenzie River valley focused primarily on freshwater fishes although some work was conducted on anadromous species in coastal areas. During the same period, potential development of coastal and nearshore gas reserves stimulated the Northern Coastal Marine Studies research on the Beaufort Sea shelf executed by DFO aboard the CCGS Nahidik (2003–9), a significant component of which focused upon marine fishes. See Journal of Marine Systems, volume 127 (November 2013), for some initial publications from this work. Marine systems research by the Department of Fisheries and Oceans (1980s to late 1990s) Scientists from the Winnipeg laboratory conducted research on marine fishes as key components of marine ecosystems, with focal sites on the western Hudson Bay coast at Saqvaqjuac (Greenland Cod, Arctic Cod) and in the central archipelago at Resolute Bay (Arctic Cod). In the 2000s this work transitioned to activities associated with oceans management particularly linked to the delineation and development of marine protected areas in Arctic waters, and is ongoing at present. Fisheries science Information needs for the development and management of sustainable commercial fisheries have traditionally driven the need for activities in fisheries science. In the Arctic there has been a long history of attempts to develop anadromous fisheries commercially, primarily those associated with chars throughout the entire area, with whitefishes in the western Arctic and James Bay, and with Atlantic Salmon along southern Hudson Strait. Research associated with these fisheries documents a wide range of knowledge, of which

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distributional information is perhaps most relevant in the context of this book. The occasional study of the marine diets of anadromous fishes has also contributed information regarding marine fishes as prey. Similarly, several limited surveys of the commercial potential of marine fishes have been conducted in coastal areas of the western Arctic and the Hudson Bay areas; however, only small-scale, local, ongoing fisheries have been further developed, primarily due to economic factors and limited resource base. Reports resulting from these various efforts provide basic knowledge, albeit biased, regarding local marine fish communities. For Baffin Bay and Davis Strait, however, the situation is different. A multispecies survey of demersal species has been conducted for many years on the Atlantic coast of Canada. Effort associated with this activity has typically been low towards the northern margin but periodically has included the southern margins of Davis Strait in support of occasional exploitation of stocks in this area. In the 1990s, interest increased in exploitation of Greenland Halibut, a demersal fish distributed throughout eastern Canadian waters north into Baffin Bay and Davis Strait, leading to the development of an inshore long-line fishery conducted through the sea ice on Cumberland Sound (a fiord on the east coast of Baffin Island) and an offshore demersal fishery. To support the needs of fishery management in the offshore, multi-species bottom trawl surveys have been conducted in the region since 1999. The survey has annually focused upon particular areas within the larger region (e.g., Baffin Bay, Davis Strait, Hudson Strait) and has been conducted aboard the FV Paamiut (a Greenlandic trawler), using a standardized depth-stratified design. The focus has been on shrimp and fish species such as Greenland Halibut, redfishes, and Grenadiers. However, the identification and enumeration of all fish and most invertebrate species caught during these surveys has provided locations of occurrence that are used in the species distribution maps herein. Voucher specimens were collected in the early years of the survey (1999–2001) and occasionally since then as new or rare specimens are encountered. In 2011 the government of Nunavut commissioned a research vessel, MV Nuliajuk, which is suitable for conducting inshore scientific surveys. During 2012 and 2013 inshore marine research was conducted in association with the DFO and university researchers from various institutions. Surveys of potential inshore commercial resources are developing the knowledge regarding marine fishes along the vast coastline of eastern Nunavut, including an annual long-line survey in Cumberland Sound (Greenland Halibut, Greenland Shark, Arctic Skate, Grenadiers, and Eelpouts) and exploratory small-scale gill-net, long-line, and trawl surveys in other coastal areas (e.g., Merchants Bay, 2011; Scott Inlet, 2011–13 and continuing) that have captured a suite of species similar to that in offshore waters.

Climate change research in the Western Arctic Increasing recognition in the 1990s of climate change as a significant driver of Arctic change in the marine system, and awareness of large gaps in understanding the physical processes associated with climate variability, led to a number of intersecting oceanographic programs in the ensuing decades. They included the following,

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many of which focused in the Beaufort Sea, but some also conducted work further east: Surface Heat Budget of the Arctic Ocean (SHEBA, 1997–8), Shelf-Basin Interactions (SBI, 1998–2008), Canadian Arctic Shelf Exchange Studies (CASES, 2001–7), Circumpolar Flaw Lead Study (CFL, 2007–8, International Polar Year), and ArcticNet activities (2004–present). These programs were mostly delivered through funding to university consortia but have also involved government and museum scientists. Most projects within the programs focused on topics other than marine fishes, and all were executed from ice-breaker platforms (and sea ice); thus, the sampling of fishes was limited in most cases to upper-water-column or near-surface work, or remote sensing (e.g., hydro-acoustics). Several publications concentrating mostly on Arctic Cod have resulted. Some project components of the ArcticNet programs also focused on anadromous fishes (primarily Arctic Char in the east). Outputs include various web-based materials, annual reports for multi-year programs, primary publications, and regional syntheses (i.e., IRIS, Integrated Regional Impact Studies).

Offshore oil development in the western Arctic (offshore marine) The suspected presence of oil-bearing strata in the deeper shelf and slope waters of the southern Canadian Beaufort Sea, combined with climate change effects allowing greater access, have heightened concerns by the Inuvialuit that such development may have an impact on subsistence food resources and the area’s marine systems. In 2006 this led to the development of the Beaufort Regional Environmental Assessment (BREA), a government program explicitly designed to address gaps and to integrate the existing knowledge of the coastal and marine ecosystems. In the preliminary “gaps” analysis, the knowledge base regarding the offshore marine fishes was highlighted by its absence, and a marine fishes project was developed. Led by James Reist through the DFO, the project involves government, museum, and university researchers in surveying the fishes and their ecological relationships in shelf and offshore deeper waters (targeting depths of 2,000 m for benthic sampling), with a focus on the southern Canadian Beaufort Sea; the work is conducted from a leased ice-capable fisheries trawler, FV Frosti. Preliminary results (for the 2012 to 2014 field years) increased distributional knowledge of the species known in the area, incremented the species complement in this area by around 14 (including the addition of one or two species new to the Canadian fauna), and provided samples across trophic levels for follow-on analyses of pathways and energetic studies. The hydro-acoustics assessment of mid-water fishes is also resulting in biomass estimates and habitat-use information, primarily for Arctic Cod, the main pelagic fish species in the area. The presence of seasonal ice in the northern Canadian Beaufort Sea continues to limit sampling; however, the work planned for 2014 and beyond attempts to address this gap. Publications in the immediate future will outline findings, and species newly added to the regional or Canadian faunas as a result of this research are included in the present work. Similar research has been recently conducted in the Alaskan Beaufort Sea (e.g., Logerwell et al., 2015).



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General Knowledge remains extremely limited regarding marine fishes of the Canadian Exclusive Economic Zone waters northwest and north of the Arctic Archipelago, those within the archipelago (especially the northern areas), and those in the northern portion of Baffin Bay including Nares Strait, especially in waters deeper than 1,000  meters.

Regional Works Linked to the Arctic Area As noted in the introduction, Marine Fishes of Arctic Canada is the first comprehensive regional work focused on the marine fishes of Canadian Arctic waters. As such, its specific antecedent is Coad and Reist’s (2004) Annotated List of the Arctic Marine Fishes of Canada. Both that contribution and the present book are based upon initial work by D.E. McAllister. General ichthyological treatises exist for other areas of the Arctic marine environment. Many of these have a Eurasian focus and do not directly overlap here. However, as some species are widely distributed in Arctic waters, treatments in other regional compendia may provide relevant information, in particular: Andriyashev, Fishes of the Northern Seas of the USSR (1954, 1967); and Andriyashev and Chernova, Annotated List of Fishlike Vertebrates and Fish of the Arctic Seas and Adjacent Waters (1995). Russian freshwater compendia also contain relevant information. In addition to these works of general relevance, the following regional works link specifically to fishes found in the Canadian marine Arctic in the contexts indicated. There is also literature available with regard to individual species, with that for widespread taxa (e.g., Arctic Cod) being more comprehensive. • Christiansen, Reist, et al. (2013), “Fishes,” in Arctic Biodiversity Assessment, which provides a synoptic overview of the fishes present in Arctic fresh and marine waters, in the context of Canadian marine fishes. Most Arctic marine fishes are benthic in habit and distributed primarily upon coastal shelf seas and adjacent slopes. Regional examinations are not provided, but appendix materials are available and give some details. • Coad and Reist (2004), Annotated List of the Arctic Marine Fishes of Canada, which provides a preliminary listing of the fishes present, local names, and brief notes on their ecology and distribution. • Dymond (1933), “Biological and Oceanographic Conditions in Hudson Bay,” part 8, “The Coregonine Fishes of Hudson and James Bays.” • Galbraith and Hunter (1976), “Fishes of Offshore Waters and Tuktoyaktuk Peninsula.” • Hunter, Leach, McAllister, and Steigerwald (1984), “A Distributional Atlas of Records of the Marine Fishes of Arctic Canada in the National Museum of Natural Sciences of Canada and Arctic Biological Station.” • Kendel, Johnston, Lobsiger, and Kozak (1975), Fishes of the Yukon Coast.

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• Legendre, Hunter, and McAllister (1975), “French, English and Scientific Names of Marine Fishes of Arctic Canada / Noms français, anglais et scientifique des poissons marins de l’Arctique canadien.” • Logerwell et al. (2015), “Fish Communities across a Spectrum of Habitats in the Western Beaufort Sea and Chukchi Sea.” • Majewski, Lynn, Lowdon, Williams, and Reist (2013), “Community Composition of Demersal Marine Fishes on the Canadian Beaufort Shelf and at Herschel Island, Yukon Territory” (also see additional papers regarding larval fishes in the same, special issue of Journal of Marine Systems, as well as the references therein for additional publications regarding the Beaufort Sea). • McAllister (1960), “List of the Marine Fishes of Canada.” • McAllister, Legendre, and Hunter (1987), “Liste des noms inuktitut (esquimaux), français, anglais et scientifiques des poissons marins du Canada arctique / List of Inuktitut (Eskimo), French, English and Scientific Names of Marine Fishes of Arctic Canada.” • McPhail and Lindsey (1970), Freshwater Fishes of Northwestern Canada and Alaska, which contains summaries of freshwater and anadromous species found in Beringia and northwestern Arctic Canada east to northern Foxe Basin, including taxonomic and zoogeographic information. • Mecklenburg, Mecklenburg, and Thorsteinson (2002), Fishes of Alaska, a summary of all of the fishes found in Alaskan waters. Of particular interest are the descriptions of fishes from the Alaskan Beaufort, Chukchi, and northern Bering Seas that are also found in Canadian waters. Earlier works on Alaskan fishes that are cited are also relevant. • Møller et al. (2010), “A Checklist of the Fish Fauna of Greenland Waters,” an overview of the marine fishes in Greenlandic waters, the western and southern portions of which exhibit species in common with the eastern Arctic portion of our area. Also see some of the literature cited in reference to eastern Arctic Canadian species. • Mueter et al. (2013), “Arctic Marine Fishes,” which provides a general overview of Arctic marine fishes and devotes one section to those in Canadian waters. • Okamura et al. (1995), Fishes Collected by the R/V Shinkai Maru around Greenland, with Japanese text; photographs of most species are provided. • Percy (1975), “Fishes of the Outer Mackenzie Delta.” • Scott and Crossman (1973), Freshwater Fishes of Canada, which contains summaries of freshwater and anadromous species found in Canadian freshwaters, including those throughout the Arctic. • Scott and Scott (1988), Atlantic Fishes of Canada, a compendium (updated) of northwestern Atlantic marine species north to the

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Labrador Sea, including many that are distributed further north into the eastern Arctic portion of our area. • Vladykov (1933a), “Biological and Oceanographic Conditions in Hudson Bay,” part 9, “Fishes from the Hudson Bay Region (Except the Coregonidae).” • Walters (1955), “Fishes of Western Arctic America and Eastern Arctic Siberia,” including nearshore marine and freshwater and anadromous species.

Summary The geographic scope, remoteness, and environmental conditions of Canada’s Arctic marine ecosystems have hampered, and continue to hamper, the development of knowledge regarding the taxa and diversity of fishes present in the area. The faunas range from sub-Arctic to Arctic in ecological nature and occupy seas that are wholly ice covered for at least some portion of the year. Most of the present knowledge results from the last seventy years of effort despite more than four hundred years of investigatory activity in the region. The coverage by surveys is biased towards coastal areas, southern areas, and areas that were (and are increasingly) ice free in summer. Thus, large areas remain either unsurveyed or poorly sampled. Where reasonable coverage has occurred, the combination of sporadic exploratory surveys, ad hoc observations, and fishery-management activities accounts for the majority of the knowledge base. Accordingly, it is highly likely that some taxa originally present in the region remain undocumented and that the known distributions of many others are substantively under-estimated. Some areas have seen no sampling or, at best, isolated sporadic sampling. Thus, the present fauna overall is poorly known, a situation that is all the more disturbing because Arctic change, primarily that associated with climate change, appears to be altering faunal complements and species distributions before they are properly documented. Generally, diversity is likely to be higher than that documented herein, and present understanding does not allow for adequate consideration of local differentiation and variation within taxa across the vast geographic area involved. Although this compendium has been in the works for over fifty years, it is quite possible that updates will be required in the near future as distributions change and ecosystems equilibrate.

sources: Forster (1773); Pennant (1784–5); Anderson (1917b);

Dunbar & Hildebrand (1952); Hattersley-Smith (1952); Dunbar (1953, 1956, 1983); Fisher (1958b); Collin (1960); Dymond (1964); Ricker (1975); Johnstone (1977); Scudder (1987); McAllister (1990a); Grainger (1995); Tunnicliffe (2012, 2013); Doel, Friedman, Lajus, Sörlin, & Wråkberg (2014).

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HABITATS James D. Reist

The habitats available to Arctic marine fishes are many and varied and share similarities with those found in southern oceans. However, great differences exist as well, particularly in habitats near to the surface and land. As noted in the “Environment” and “Climate” sections of the introduction, the Arctic environment is characterized by extremes of seasonality and variability. These factors in turn greatly affect the habitats within which Arctic marine fishes must live. Arctic marine habitats are significantly influenced by atmospheric climate variables. This influence is greatest near the air-water

interface, but the effects are transmitted to a greater or lesser extent to less proximate marine habitats. The most significant physical parameters are the cold atmospheric temperatures and the extreme seasonality of incident radiation, including thermal energy – for example, there is almost none for up to six months of the year. These result in the formation and persistence of large amounts of ice, which in turn distinguishes the Arctic Ocean and greatly influences the marine habitats available to fish. Ice is also a habitat in and of itself, as discussed below. (See figs. 23–35.)

Fig. 23. Beaufort Sea ice, August 2014.



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Fig. 24. Banks and Victoria Islands.

Fig. 25. Sea ice off Baffin Island.

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Fig. 26. Spring in the Canadian Arctic.

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Fig. 27. Northeast Baffin Island.

Variability in Arctic Habitats and of Associated Fish The extreme seasonality and variability of the Arctic environment also creates a highly dynamic situation in many Arctic marine habitats, to which the fish have adapted. General adaptations to cold environments are discussed in the “Environment” section of the introduction, and specific adaptations may be found in this section. The seasonality of habitat characteristics results in a high degree of seasonality of fish association with such dynamic habitats. Thus, superimposed on all the variability is the temporally structured association of many fish species with different specific habitats as they follow their seasonal cycle throughout the year. Perhaps the best example of this for the Arctic is the extreme seasonal habitat association shown by anadromous fishes. Habitats used by anadromous taxa range from wholly freshwater ones such as rivers and lakes, through estuarine zones of varying degrees of salinity, to wholly marine habitats. Use of a particular habitat is highly seasonal and is closely tied to the species’ biology. For example, the anadromous components of all Arctic anadromous



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species use accessible rivers as migratory corridors, typically downstream in the spring and upstream in the autumn. Such corridors link the freshwater habitats that are used for spawning and overwintering (lakes, rivers, or both, depending upon the species) with the estuarine and marine habitats that are primarily used for summer feeding. So far as is known, no species of anadromous fish, especially within the Salmonidae, remains in the marine system during winter. Some, however, may use the freshened areas of the estuaries of rivers that flow continuously during winter (e.g., Arctic Cisco are found in the estuary of the Colville River, northern Alaska). Limited winter work to investigate this issue has been conducted in Canada. Additionally, some anadromous species (e.g., Pacific Salmon) have a life history that dictates remaining in marine areas for several years, including overwintering; it is unclear where these Arctic populations overwinter. In addition to temporal variability, spatial variability is evident within particular habitats. Estuaries represent an excellent example of this variability, grading from low-salinity, wholly freshwater zones, through variously mixed zones, to zones of high-salinity marine waters. In the Arctic this gradation is almost always paralleled by a high thermal gradient in summer water temperature

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Fig. 28. Cliffs near Arctic Bay.

Fig. 29. Sachs Harbour (Ikahuak).

Fig. 30. Kugluktuk.

Fig. 31. Davis Strait.

from warm in fresh water to relatively cold in marine waters. The size, duration, and temporal stability of such spatial zones within particular estuaries is directly associated with the size of the river. Furthermore, freshwater plumes from larger rivers often extend far offshore and persist seasonally due to density differences, thus in essence extending the estuarine habitat well into the marine environment. Fish respond to such spatial structuring within habitats, and this response may be highly organized throughout the life history of individuals with particular life-history stages (or sizes or ages) sequentially associating with very specific conditions within the habitat. Thus, for example, the increased salinity tolerance of Arctic anadromous species appears to depend in part on the size of the individuals, with smaller individuals being less tolerant. Consequently, larger individuals that have made the transition to sea (i.e., smoltified) in previous years are associated with higher-salinity waters that are typically found in outer estuarine and nearshore habitats, whereas smaller individuals are found in less saline, inner

estuarine waters. Similar partitioning of the habitat can also occur on small spatial scales. For example, Arctic Cod use the underside of sea ice in part to avoid predation and in part to feed. Small cod can enter small interstitial spaces in the ice, but larger individuals cannot and so remain in larger cracks and openings or associated with the underside of the ice. Both temporal and spatial variability within habitats appear to be greatest for habitats that are close to the air-water interface (surface and epipelagic zones) and to terrestrial areas (estuaries, nearshore zones, continental shelves). Marine habitats that are more distant from such areas exhibit greater stability. Thus, mesoand bathypelagic habitats and associated abyssal and benthic areas in the Arctic Ocean appear to be comparatively stable both spatially and temporally. This may also be true for epipelagic habitats covered by polar pack ice. However, little appropriate work has been conducted in all these areas in the Arctic, and understanding is limited.

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Fig. 32. Pauline Cove, Herschel Island.

Fig. 33. East coast of Baffin Island.



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Fig. 34. HMCS Labrador in ice in Baffin Bay, 1955. Fig. 35. Below, James Bay in summer and winter.

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Fig. 36. Darnley Bay, August 2015.

Factors That Define Arctic Marine Fish Habitats Habitats available to fish can and have been described according to numerous schemes. Those that seem to work the best are defined in terms of the fish themselves and in part represent the adaptation of fish to the specific conditions found in particular habitats. Using the general background outlined above, this approach will be followed here. Habitats are defined from the perspective of the organisms using them and are structured by the physical and to a lesser extent the chemical properties of the environment in which they occur. Thus, the major factors that structure Arctic marine habitats include proximity to land; proximity to freshwater inputs, and the volume and seasonality of those; characteristics of marine water mass including temperature, salinity, and flow; depth; type and size of substrate (i.e., physical surface features such as boulder content); type and coverage of any vegetation; ice type and coverage; exposure to wind, currents, and tides; and bottom topography. These factors combine variously to create unique habitats that can be grouped together in



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any number of ways. It is important to note that no one variable defines a habitat; rather habitats are distinguished by unique combinations of many variables.

General Marine Habitats Most general schemes that define marine habitats group them primarily upon the basis of proximity to coasts and secondarily upon the basis of depth and association with the bottom. Thus, marine environments of the water column can be differentiated into neritic or shelf-zone habitats, and those present farther offshore as oceanic habitats. Similarly marine environments associated with the bottom (i.e., benthic habitats) can be separated into those on continental shelves and those occupying abyssal areas. (See figs. 37–43.)

Neritic Water-Column Habitats Waters over the continental shelves tend to be shallow (about 200 m or less) and are greatly influenced by nearshore and surface

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Fig. 37. Baffin Bay at 600 m, September 2012, Lycodonus mirabilis.

Fig. 38. Baffin Bay at 600 m, September 2012, Lyodes cf. eudipleurostictus.

Fig. 39. Baffin Bay, September 2012, bottom invertebrates and possibly a Boreogadus saida (centre).

Fig. 40. Davis Strait, bottom habitat, October 2012.

processes. Habitats in the water column that are available to Arctic fish include the following. First, ice consisting primarily of seasonal and land-fast types creates several surface (epipelagic) habitats that are highly relevant to many Arctic marine fishes. Multi-year ice may also be present in shelf areas but is more typical of oceanic areas in the Arctic (i.e., the polar pack). When present in association with land-fast or seasonal first-year ice, multi-year ice provides additional habitat diversity in some nearshore areas, particularly those found in the channels of the Canadian Arctic Archipelago that receive the input of multi-year ice from the polar pack of the Arctic Ocean. Productivity in shelfzone waters tends to be higher than in offshore waters, and much of this is directly associated with ice communities. Some Arctic fish species such as Arctic Cod have adapted to this unique habitat and are found associated with ice throughout the Arctic Ocean. Shear zones at the edge of land-fast ice and near upwelling areas create leads and polynyas. Such openings, as well as ice edges generally, constitute a special type of epipelagic habitat and tend to be focal points for many species. True pelagic Arctic fish are generally few in number, although Arctic Cod may at times exhibit pelagic tendencies. Epipelagic species congregate near ice edges to take advantage

of high productivity occurring locally and in the nearby polynya and also to use the ice as a refuge from predators such as marine birds and mammals. Second, below the ice, the Polar Mixed Layer (the upper part, about 50 m, of the water column) provides an epipelagic habitat that experiences wide-ranging seasonality in salinity and temperature. Although this layer has been poorly studied from the perspective of marine fishes, it is likely that some life-history stages of several species occupy this habitat at least seasonally, for example, young Arctic Cod in offshore areas in the summer. Nearshore epipelagic waters are generally hyposaline and warm in the summer due to freshwater inputs. Arctic anadromous fishes such as Arctic Char likely occupy this habitat primarily for summer feeding, especially areas close to shore, but they may also venture into offshore epipelagic waters. Third, over the continental shelves the deeper portions of the epipelagic water column (about 50–200 m) are generally more marine in nature, though still greatly influenced by mixing processes. Nearshore areas at these depths appear to be dominated by benthic fishes (e.g., sculpins), whereas offshore areas may include mixtures of both benthic (deeper-water sculpins) and more pelagic types (such as adult Arctic Cod).

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Fig. 41. Kelp in Browns Harbour, Northwest Territories, 22 July 2015.

Oceanic Water-Column Habitats Open water or pelagic oceanic environments are progressively differentiated by depth from the surface to the bottom into epi-, meso-, bathy-, and abysso-pelagic habitats. As for the neritic areas, oceanic epipelagic habitats occupy surface waters typically to about 200 meters depth. In the Arctic Ocean this habitat corresponds almost exactly with the water mass of the Arctic Surface Layer and is dominated by ice. Oceanic ice habitats consist primarily of multi-year ice and therefore differ significantly from the ice habitats of nearshore areas. Multi-year ice tends to be much thicker and more broken and so provides larger interstitial spaces and surface relief features than does seasonal, land-fast ice. Consequently larger fish such as adult Arctic Cod appear to associate with this habitat in oceanic areas. However, productivity is likely low; thus density and abundance are very low. As for most offshore Arctic areas, this habitat has been very poorly studied. Mesopelagic habitats occur at a depth of roughly 200 to 1,000 meters, over the edges of the continental shelves and oceanic areas, which more or less correspond to the water mass of the Atlantic Layer present in the Arctic Ocean. Bathypelagic habitats are confined to offshore oceanic areas from depths of about 1,000 to 3,000



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meters, and in southern oceans are usually delineated by the 10°C (upper) and 4°C (lower) isotherms. Given the cold environment and unique situation of the Arctic Ocean, such thermal demarcation is generally not applicable here. Rather, bathypelagic habitats are delineated by water masses of differing temperature and salinity. Abyssopelagic habitats are defined by depth (> 3,000 m) and temperature (≤ 4°C). Extreme depths present in oceanic trenches are referred to as hadal habitats. As noted in the “Environment” section, the Arctic Ocean is generally characterized by wide continental shelves, and Arctic marine areas within Canada are almost completely dominated by such shelf habitats, hence by epipelagic waters. Only in the extreme northwestern portion of the Canadian Beaufort Sea, along the northwestern margin of the Arctic Archipelago, and in Baffin Bay and Davis Strait do depths generally exceed 500 meters and are usually less than 3,000 meters. Thus, oceanic habitats in the Canadian Arctic can generally be limited to epipelagic, mesopelagic, and bathypelagic types. Most species of Arctic fish are benthic rather than pelagic in nature; therefore, very few taxa are associated with specific oceanic pelagic habitats as defined here. The exception likely occurs in the eastern Canadian

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Fig. 42. Scott Inlet, 2013, epibenthic habitat.

Arctic where some pelagic species with affinities to the Atlantic Ocean occur.

Bottom Habitats Habitats that are associated with the substrate or bottom are termed benthic and are similarly structured primarily by depth and proximity to shore. Epibenthic habitats are bottom habitats in shallower shelf areas (to about 200 m). These are greatly influenced by oceanic processes of the shelf that in the Arctic include significant freshwater inflows and ice actions such as physical scouring and periodic disturbance. Continental-shelf edges usually drop off sharply to continental-slope habitats. Habitats in this zone are referred to as mesobenthic and correspond to the shoreward margin of the mesopelagic habitat of the oceanic area. Deeper benthic habitats correspond by depth to similar pelagic habitats; archibenthic to bathypelagic; and abyssobenthic to abyssopelagic. Epibenthic habitats dominate in the Canadian Arctic marine environment because of the extensive continental shelves. This is especially true for most of the Beaufort Sea, the Hudson and James Bays, the Hudson Strait, and channels throughout the archipelago. In the eastern Arctic the continental shelf is narrow and rapidly gives way to a steep continental slope. This area is dominated by the deeper meso- and abysso-benthic habitats. In addition to the above views of benthic marine habitats, further subdivision is generally done based upon proximity to shore

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and also the specific nature of the substrate in the habitat. Thus, nearshore habitats can be differentiated into those that are subject to tidal influence (i.e., intertidal) and those that are subtidal. Nearshore areas can also be defined according to freshwater inputs, that is, estuarine habitats. Surrounding land forms and exposure tend to partially isolate nearshore water bodies. Therefore, coastal habitats can include the following: lagoons separated from offshore areas by barrier islands such as those found in the Beaufort Sea and southeastern Hudson Bay; and fiords that dominate the coastline of the eastern Arctic and which, though quite deep, are usually partially cut off from deeper offshore areas by shallow sills. Fiords and similar embayments in the Canadian Arctic are often fairly large in area, exhibit a wide range of specific habitats, and may be somewhat isolated by shallow sills from the adjacent seas. Accordingly, these areas may have an ichthyofauna that is relatively distinct from that in the adjacent sea and elsewhere. Most fiords and embayments in the Canadian Arctic are poorly characterized with respect to their biodiversity (including fishes) and oceanographic processes. Benthic habitats are also subdivided in terms of the substrate that dominates an area. The substrate type (e.g., mud, sand, silt, rock, boulder) determines to a large degree the ecological community structure. For example, soft substrates provide surface habitat with which organisms can associate in various ways: epibenthic habitats support organisms associated with but usually above the bottom; benthic habitats support organisms that live on the substrate; and

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Fig. 43. Grounded iceberg, 15 ft. depth, Resolute Bay.

sub-benthic habitats provide space into which organisms can burrow (faunal assemblages, especially invertebrates associated with the latter, are referred to as infauna, a relevant food source for many benthic and demersal fishes). Rock substrates provide epibenthic and benthic habitats, but not sub-benthic areas. Stable substrates result in other differences; for example, kelp can attach to rock and boulder substrates, thereby adding physical complexity to such areas, and the kelp in turn provides additional habitat for organisms such as fish.

Habitat Associations of Fishes in the Canadian Beaufort Sea The majority of Arctic fishes as adults are benthic in habitat association. This likely results from the greater overall availability of prey and productivity at this level (i.e., as in situ lower trophic groups relying upon detrital fall) and the seasonal dynamics of pelagic production. Shelf and slope habitats especially benefit from detritus filtering to the bottom and tend to be influenced by water mass characteristics, climate drivers, and sea ice. That is, these habitats are dynamic both in space and in time. The association of water masses differing in salinity and temperature with particular benthic habitat zones is perhaps best understood for the Beaufort Sea in the Canadian Arctic. Density-driven layers



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form distinct water masses in the Arctic Ocean, which intersect the bottom at different depths in the Canadian Beaufort Sea. The combination of water mass characteristics, substrate type (e.g., grain size increasing with depth), and species’ preferences (e.g., foraging mode, and preferences for temperature, salinity, and depth), and perhaps availability of prey, all appear to influence the composition of the benthic fish communities at specific depths. Mixing zones between water masses and associated physical dynamics also likely influence habitat occupancy. Generalities that are apparent include (1) greater benthic fish diversity in the offshore-shelf and upper-slope zones in this area; (2) shifts in composition across depth zones; and (3) dominance of Arctic Cod across all zones shallower than around 500 meters. Replacement by different species within fish families (e.g., Cottidae, Zoarcidae) and the occurrence of some species only at specific depths indicate fairly tight habitat associations of many species in this area (fig. 44). Many of the benthic species of marine fishes in the Canadian Beaufort Sea associate with distinct habitats. Depth associations of sculpin species, Zoarcid species, Greenland Halibut, and Arctic Skate are particularly noteworthy. Arctic Cod, however, is found associated with all habitats, although occurrence decreases beyond 900 meters depth. This perspective focuses upon approximately sixty species of the benthic fish community. Limited numbers of pelagic species occur in this area (i.e., about five to seven marine species), and those

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appear to be distributed patchily and to be highly variable in abundance interannually). Detailed analyses, similar to those given here for benthic fishes, are not available. Like elsewhere, in this area Arctic Cod appears to occupy most habitats including ice (if it is present), surface mixed waters (especially as young), and shelf and upper-slope waters. Numerically this species is the most abundant in these habitats, although it occurs sporadically in deeper zones to at least 1,000 meters. Both individuals and individual species may be variable in their habitat association. For example, although epibenthic association is implied by its being a flatfish (Pleuronectidae), the Greenland Halibut in this area exhibits diet and trophic markers that are indicative of both benthic and pelagic feeding, despite its being found only at depths greater than 500 meters. Whether this results from active pelagic feeding or incorporation of detrital-fall pelagic items, or both, is presently unknown. Anadromous species in this area provide another general exception with respect to habitat associations. Most anadromous fishes are closely associated with coastal habitats in the Arctic, and the vast majority are adapted to upper pelagic habitats. Benthic feeding, however, appears to be frequent and is the norm for particular

Offshore Shelf

Upper Slope

Lower Slope

900 m +

300– 900 m

200– 300 m

60– 200 m

0– 60 m

Nearshore Shelf

species (e.g., whitefishes). The use of offshore pelagic habitats by anadromous species is poorly understood in the Arctic. These descriptions pertain generally to the southern portion of the Canadian Beaufort Sea at depths of between about 40 and 1,500 meters. The circumstance in deeper waters is generally unknown for this area. Similarly, in the eastern Arctic, fish association with specific habitat zones is likely, but information is limited.

0–10 km

100 km

150 km

200 km +

Fig. 44. Habitat associations of selected Arctic marine fishes.

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CLIMATE James D. Reist

The climate of the Arctic is intimately associated with the axial tilt of the Earth, which, depending upon the annual orbital position relative to the sun, allows for seasonal daily periods of full incident radiation (summer) or those of no solar radiation (winter). North of the Arctic Circle (66°30' N latitude) these periods occupy the full day length for extended periods. Moreover, the angle of incidence of the sun’s radiation is relatively low over much of the Arctic, also varying with latitude (i.e., about 100 watts per square meter north of the Arctic Circle or about half of that in southern locales). Incident radiation (or lack thereof) in the thermal and visible ranges is an essential element of climate and of production relationships throughout the Arctic. Accordingly, both the high seasonal periodicity (i.e., the majority of incoming radiation between the vernal and autumnal equinoxes) and the angle of incidence of incoming radiation affect local climates that are heavily pulsed, and amounts are overall lower in magnitude than those in southerly latitudes. Annually the seasons are characterized by long periods of relative warmth and cold. Temperatures below 0°C induce freezing, and a dominant aspect of the Arctic is ice present in various forms (i.e., sea ice, freshwater ice, snow, frozen landscapes, and permafrost). Ice further modifies both the annual climate and the productivity cycles (e.g., sea ice affects wind fetch, evaporation, atmospheric heating, and the radiant energy available to the water column; heating and melting of sea ice also reduces energy available for production). Ocean circulation patterns and associated large-scale heat transport within the water column further affect local weather and Arctic climates. Canadian Arctic marine waters are spatially extensive and occupy a wide range of latitudes (i.e., about 54° N in southern James Bay to about 87° N at the northern margin of the Canadian Exclusive Economic Zone, 200 nautical miles north of Ellesmere Island) and longitudes (i.e., 141° W at the border with Alaska to 57° W in central Baffin Bay). These waters border the central Arctic Ocean, encompass the Arctic Archipelago, and border the continental mainland along three extensive coastlines (i.e., about 173,000 km, or 71% of the total Canadian marine coastline, or about four times the circumference of the Earth at the equator). The areal extent of



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Arctic marine waters in the Canadian sector, including north to the pole, is about 3.2 million square kilometers. Extensive sea ice and almost complete coverage occur on these waters over the winter; in summer, seasonal melting occurs along coastlines and extends over many of the water bodies (e.g., all of Hudson Bay, most of Baffin Bay and Davis Strait, and much of the southern Beaufort Sea). Persistent pack ice dominates the central Arctic Ocean and many of the channels of the archipelago, although in recent decades cumulative degradation of this has occurred. Ice coverage, duration and seasonal extent, presence of open-water areas during winter (i.e., polynyas and shore leads), and heat content and transport associated with marine water masses all interact with atmospheric conditions (e.g., Arctic low-pressure systems) to generate a highly variable climate both temporally and spatially. Temporal variation occurs seasonally, interannually, and over longer time frames; thus, trying to characterize a “typical” climate is difficult and is further complicated by recent apparent changes in the climate. Spatial variation can be generalized regionally, albeit with the presence of high variability. Climate affects fishes and their supporting ecosystems through production pathways, ocean heating, and mixing of waters among habitats, and by providing key cues for life-history activities. Such linkages are species specific and more directly affect fish species that occur close to shore and ice edges and in upper levels of the water column than they do those that are offshore or in deeper waters. In addition to direct effects (e.g., thermal regimes, wind mixing), climate indirectly affects fishes through seasonal or perennial ice presence and through seasonal flows of fresh water (i.e., from both ice melt and rivers). Ice also physically influences fishes through its scouring of shallow areas and as a habitat, as well as through salt extrusion (and hypersalinities) during its winter formation. Climate regions delineated for the terrestrial Canadian Arctic correspond roughly with major marine regions, the climate of which will therefore be described separately. These regions are relatively large, and thus extensive spatial variation within them is to be expected. The marine regions are (1) the Beaufort Sea, (2) the Arctic Archipelago and Foxe Basin, (3) the northwestern Arctic

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Archipelago, (4) the south-central Arctic Archipelago, (5) Baffin Island and the northeastern Arctic Archipelago, (6) Hudson and James Bays and Hudson Strait, and (7) Baffin Bay and Davis Strait. Adjacent terrestrial landscapes and topography also affect regional climates especially from west to east due to the Coriolis effect. Thus terrestrial climate zones that respectively influence the above marine regions include Western Arctic Tundra, Central Arctic Tundra, Arctic Mountains and Fiords, and Arctic Tundra in the north, and Northeastern Forest in the south. Finally, the climate of these marine regions is also affected seasonally, annually, and over decadal scales by large-scale hemispheric patterns of climate in lower latitudes in relation to those of the Arctic. These include those of the North Pacific (e.g., Aleutian Low) as well as those of the North Atlantic (e.g., North Atlantic Oscillation). The historical (i.e., pre-1990) mean annual air temperature range was −20°C to −15°C in the northern portion of Arctic Canada (i.e., from the southern archipelago to the pole); −10°C to −15°C throughout the northern mainland, Beaufort Sea, southern archipelago, Foxe Basin, and southern Baffin Island; −5°C to −10°C throughout Hudson Bay, northern Québec, and southern Davis Strait; and 0°C to −5°C in James Bay. For these same areas respectively, January mean temperatures ranged from −35°C to −30°C, −25°C to −30°C, and −20°C to −25°C (inclusive of James Bay). The southern margin of Davis Strait in our area is ameliorated by Atlantic water inflows, and its temperature ranged from −15°C to −20°C. July mean temperatures ranged from 0°C to 5°C over the northern Beaufort Sea, the northern archipelago and much of Baffin Bay and Davis Strait. They ranged from 5°C to 10°C over the southern Beaufort Sea, southern archipelago, Foxe Basin, northern Hudson Bay, and Hudson Strait, and generally averaged from 10°C to 15°C over southern Hudson and James Bays, with the extreme southern area of James Bay being marginally warmer. Average sea-surface temperatures by contrast are much warmer in winter (beneath the ice) and colder in summer; e.g., the annual means for the northern Beaufort Sea were −0.5°C to 0°C, and 3°C to 4°C in southern shelf areas. Recent climate changes have generally increased mean temperatures throughout the Arctic; however, high variabilities remain. Ice cover is the dominant attribute of the Arctic marine system, particularly in the shallow waters found in much of the Canadian Arctic. Its formation and duration over time (seasonally and interannually) is governed primarily by atmospheric climate. Wind, atmospheric-pressure gradients, and freshwater inputs (and salinity densities) influence both the large-scale patterns of summer seaice distribution and the large amounts of annual sea-water export from west to east (44,000 cu km annual outflow through to Baffin Bay). Arctic marine waters also feed back and influence the climate system both locally and globally. The complex interaction among climate drivers (insolation, pressure differentials) and the marine system (ice, nutrients, production, currents, bathymetry) influences both the diversity and the productivity of marine fishes.

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Beaufort Sea The Beaufort Sea marine ecosystem is a true Arctic system closely associated with the Arctic Ocean; accordingly, climate is driven by processes originating in the geographically central polar area (e.g., semi-stable polar vortex in winter and Arctic high in summer). This is mediated to some degree by the North Pacific influences transmitted over the western Arctic land mass. Sea ice (polar multi-year pack ice, seasonal sea ice, land-fast ice) heavily modifies local climates, thereby affecting the marine ecosystem. Winter ice coverage is more or less complete, broken only by shore leads along the shear zone between land-fast and pack ice, and by a few polynyas towards the east. The mean annual temperature range of thirty-six Celsius degrees, and the cold winters (−28°C to −30°C) extending from the end of August through to the end of the following May have historically dominated. A short, three-month summer with mean temperatures of 5°C to 8°C occurs. Generally it is dry and cold with annual precipitation of about 250 millimeters. Irradiance is moderate between 20 (south) and 10 (north) kilo-Langleys annually (where 1 kLy equals 11.622 kilowatt hours per square meter). Pressure gradients from west to east and a long open-water fetch paralleling the continental coast result in high storm frequency with high winds of long duration in late summer and winters. Summer open-water areas about a hundred kilometers offshore of coasts typified the area to the mid-1990s, and accompanying extensive pack ice tended to ameliorate storms and cool the air masses. Recent summer sea-ice changes (i.e., melt, export, thinning) have resulted in much more open water during summer (though still highly variable across years). This has two consequences for the marine system: greater wind mixing in upper layers, and increased heat transport into the ocean. The effects of these consequences on the structure and function of the ecosystem are poorly understood at present. The combination of sea-ice melt, freshwater inputs, eastwards movement of surface waters of Pacific origin, and the deeper waters of Atlantic origin all result in a heavily layered system of water masses structured by density (i.e., salinity) and temperature (see the “Environment” section in the introduction). The top 50 meters (Polar Mixed Layer) is freshened and, together with Pacific-origin waters (about 150–200 m depths), tends to be cold (about −1.4°C) in summer and slightly colder (−1.8°C) in winter, with salinities of 30–33 practical salinity units. A semi-stable halocline between about 200 meters and 300 meters depth is present, below which resides water that is primarily of Atlantic origin (300–900+ m depth, 0.4°C to 1.0°C, and slightly higher salinities of 34–35 psu). Deeper western layers, over 900 meters, are characterized by the cold saline waters of the Canada Basin (i.e., Arctic bottom water, −1.4°C, > 34 psu). Seasonal storms, ice dynamics, and continual winter freshwater flows from the larger rivers accumulate nutrients on the shelf and in upper water layers, particularly over winter. These facilitate early-season, under-ice primary and secondary production in the following polar spring when sunlight returns. The high latitude means that the sun is above the horizon continuously from late May to mid-July and below it from early December to early January. Although the tides

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are relatively low in amplitude, ice scour of bottom habitats is typically significant in shelf areas, particularly in the shear zone (i.e., 17–25 m depths) where pack ice impinging upon land-fast ice creates large pressure ridges.

Arctic Archipelago and Foxe Basin The topography (low in the west, high in the east), cyclonic activity (dominantly from the Baffin Bay–Davis Strait area), sea-ice presence and persistence, and variation in net annual radiation delimit three sub-areas in this region. They overlap and are indistinct from each other, as well as from the areas to the west and north (Beaufort Sea and Arctic Ocean), those to the east (Baffin Bay–Davis Strait), and those to the south (Hudson Bay complex).

Northwestern Arctic Archipelago The northwestern portion of the Arctic Archipelago encompasses the islands north of Lancaster Sound, Parry Channel, and McClure Strait, and west of the Devon and Axel Heiberg islands. This area is characterized by low annual irradiance (i.e., 5–10 kLy); annual mean temperature ranges of 37–40 Celsius degrees; January mean daily temperatures of −33°C to −35°C; June daily means of 5°C to 3°C, south to north; winter beginning around 20 August and extending to about 15 June; and less than 100–150 millimeters of annual precipitation. To the late 1990s, land-fast ice consisting of second-year or multi-year ice typically dominated the western channels, with four-tenths to six-tenths coverage persisting in late summer. In eastern areas complete ice melt and export typically occurred, resulting in open waters. Recently, later and thinner ice formation in the winter, combined with earlier break-up, has resulted in much of the area being seasonally free of ice. High interannual variability, however, occurs in the areal extent, age, and seasonal persistence of sea ice. Marine channels in the area have substantive shelves (< 200 m depths), and all areas are less than 500 meters deep with the exception of some smaller, isolated, deeper areas (all < 1,000 m). Productivity tends to be low overall.

South-Central Arctic Archipelago The south-central Arctic Archipelago includes Victoria Island, Boothia Peninsula, Melville Peninsula, and western Foxe Basin. The area is characterized by moderate (for the Canadian Arctic) annual irradiance (10–15 kLy), an annual mean temperature range of 36–45 Celsius degrees, January mean daily temperatures of −30°C to −35°C, June daily means of 8°C–5°C over water from south to north, winter beginning around late August to mid-September and ending in mid-June, and 100–200 millimeters of precipitation annually in the north and up to 300 millimeters in the southeast (Foxe Basin). Water bodies have limited nearshore shallow areas and moderately extensive shelves (< 200 m depths), and most areas are not much deeper. Sea ice is lost annually through much of the area but may persist in narrow channels “downstream” of source areas to the west. Productivity, particularly in areas where sea ice is absent, tends to be higher than that found further north.



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Baffin Island and Northeastern Arctic Archipelago The Baffin Island and northeastern Arctic Archipelago area includes all the eastern islands of the archipelago (i.e., from Baffin Island north to Ellesmere). This area is characterized by high south-tonorth gradients: irradiances of 30–5 kLy, mean annual temperature ranges of 23–37 Celsius degrees, January mean daily temperatures of −20°C to −33°C, June daily means of 8°C–5°C, winter beginning at around the end of September to the end of August and lasting until about 10 to 15 June, with 600 millimeters (southern mountain slopes) to 150 millimeters (northern over water) of mean annual precipitation. Extensive shelf areas are present nearshore and between most islands; however, deep channels are found in Lancaster and Jones Sounds. Sea ice consists mostly of first-year mobile ice in the south and multi-year mobile ice in the north. Except for the extreme northern portion, sea ice disappears completely over much of the area. Productivity appears to be moderate in southern areas, declining significantly in the north. Linkages in the south extend to Hudson Bay and Hudson Strait; eastern areas are linked to Baffin Bay and Davis Strait. The ice-bound northern areas are heavily influenced by the adjacent Arctic Ocean, which also exerts some influence via outflows through Nares Strait.

Hudson and James Bays and Hudson Strait The Hudson Bay marine system is a mostly enclosed area that is relatively colder than similar areas at the same latitude although a significant latitudinal gradient exists within the complex. From an atmospheric climate perspective, Foxe Basin is more closely allied with the central Arctic Archipelago and is treated in that description. The land-based adjacent Arctic Tundra climate zone influences the northwestern portion of Hudson Bay proper with colder polar winds and persistent low temperatures. The Northeastern Forest zone moderates climate in the southern and much of the eastern portion of the bay. High freshwater inputs characterize the area (i.e., the Churchill-Nelson system in the west, a number of large rivers in the south, and the Great Whale and other rivers in the east, with, for example, a 64 cm net gain in sea level over the entire bay annually) and affect oceanographic and ice conditions. These are further influenced by local winds and radiation inputs. The bay becomes completely ice covered annually except for some limited shore leads and local polynyas, and wind-driven ice motion is constant over most of the bay in winter. Seasonal ice melt is almost complete, and very little multi-year ice is present. Surface circulation is counter-clockwise (cyclonic) and fed by westwards currents along northern Hudson Strait and southward currents exiting Foxe Basin. Outflow from this mostly enclosed marine system occurs along the southern margin of Hudson Strait. Colder, more marine waters characterize Hudson Bay itself, whereas warmer waters with lower salinities characterize James Bay. Marine faunas accordingly are more Arctic in nature in the north, and boreal in the south. Waters below 100 meters depth are saline (33 psu) and cold (−1.4°C), with those of James Bay being more seasonally variable. Surface waters in

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Hudson Bay range from about 8°C as summer highs to −1.2°C at 36 meters depth with salinities spatially varying between 23 and 30 psu. Surface salinities in James Bay vary from 10 to 24 psu in summer and from 15 to 31 psu in winter. The average depth of Hudson Bay is 100 meters, with some areas approaching 300 meters; James Bay is shallow with less than 60 meter depths.

Baffin Bay and Davis Strait Prevailing currents, particularly the west Greenland Current bringing Atlantic water that is relatively warm (4°C in summer) and saline (about 34 psu), ameliorate the local climate of this system especially in the southern areas and towards the Greenland side. Waters from the Arctic Ocean entering from the north via Nares Strait, and freshened waters from the Arctic Archipelago entering from the west, tend to be colder and less saline (about 30 psu). These combine to form the deeper waters of the system with a temperature of −0.5°C and salinity of about 34 psu. Ice dominates the area in winter (about 80% coverage) particularly in the north, persisting as drifting ice in western areas (Canadian waters) well into the summer. Icebergs, sloughed from Ellesmere Island and Greenland, are also present throughout this area. Cyclonic weather patterns moving northwards through the area dominate weather most of the year with northwesterly winds. In comparison to other Canadian Arctic marine areas, this region is warmer with a narrower mean annual temperature range of 26–37 Celsius degrees from south to north. Daily mean temperatures in January (−20°C in the south, −33°C in the north) and July (8°C in the south, 3°C in the north) reflect this warmer climate, as does the comparatively short winter (from the end of September to the middle of June in the south). Annual precipitation (150– 300 mm) is also higher, particularly at higher elevations in the north. The northern area is also heavily influenced by the climate of the northern archipelago. Recent changes in annual average temperatures indicate that warming is occurring; these averaged a change of about 0.8°C over the last decade and have been accompanied by a 30% increase in average annual precipitation. Topographically the eastern Arctic islands generally exhibit higher elevations, many of which have significant ice masses and glaciers. These factors thus add a cooling influence generally to the environment that is transported eastwards and down slope, influencing the waters of coastal fiords and Baffin Bay and Davis Strait. Narrow shelves and deeper offshore waters of the marine system, however, tend to uncouple atmospheric climate drivers and marine responses especially for offshore areas. Oceanographic influences, particularly water inputs and currents as noted, are most relevant in this region.

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Overall Effects of Climate Change Significant and geologically recent increased concentrations of greenhouse gases (i.e., primarily carbon dioxide and methane) in the atmosphere have resulted in increases in mean annual temperatures globally and have altered precipitation patterns. These effects are amplified in the polar regions – such that the Arctic has experienced about a two-degree increase in annual mean temperature since the early 1900s – and are projected to increase in the future both in magnitude and in rates of change due to continued anthropogenic greenhouse-gas forcings and to the effects of interaction with the polar environment (e.g., sea-ice loss results in lower albedo, which then results in higher thermal absorption by Arctic marine waters). Anticipated changes for Arctic North America range from a further 2°C–4°C increase by the end of the century from the 1986–2005 mean, depending upon which anthropogenic emission scenario is used. Loss and draw-down of perennial ice stores (e.g., multi-year sea ice, permafrost, glaciers) and increased precipitation will continue to alter both the physical and the chemical components of Arctic marine ecosystems. Freshening, warming of the water column, and earlier ice loss will shift much of the primary production from sea ice to open water. Generally it appears that overall productivity may increase but will likely result more and more from pelagic than under-ice production. Nutrients, particularly iron, however, will become limiting, and productivity shifts may not be as large as expected. This is particularly so for offshore areas. Moreover, high atmospheric concentrations of carbon dioxide increase acidification, to which the Arctic Ocean is particularly sensitive. Acidification, in turn, will very likely affect both the nature and the efficacy of production pathways by reducing or eliminating key components in the pelagic and benthic foodwebs. This may also directly affect marine fishes (e.g., impacts on ichthyoplankton). Accordingly, significant shifts in marine ecosystems, their structural pathways, and the biodiversity of their constituent parts are all expected. Direct responses by marine fishes may also include colonization of more northerly areas as conditions change (i.e., overall diversity in an area accessible to sub-Arctic species may increase). Indirect responses will likely involve changes in population abundances and thus fisheries. From a human perspective, the climate-change effects on the marine system are expected to increase access both seasonally (e.g., longer open-water seasons) and geographically (e.g., the degradation of multi-year pack ice; see the record September sea-ice minimum in 2012 shown in fig. 7). Increased human activity (e.g., shipping, industrial development, possibly fishing) will have obvious consequences for marine fishes. Opportunities for research on marine fishes and their ecosystems, however, will also increase as access increases.

sources:Maxwell (1981); D.B. Stewart & Lockhart (2004); Arctic

Climate Impact Assessment (2005); Cobb et al. (2008); Arctic Monitoring and Assessment Programme (2011, 2012, 2014); Melling et al. (2012); Intergovernmental Panel on Climate Change (2014).

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ARCTIC FISH, NORTHERN CULTURES, AND TRADITIONAL ECOLOGICAL KNOWLEDGE Fikret Berkes

If Inuit art is any indication of the importance of fish in the lives of the Inuit, there is ample evidence in the form of carvings and prints from a variety of areas of the Arctic. However, fish are not among the spiritually powerful animals such as polar bears, seals, whales, and caribou and do not appear as frequently in Inuit art as do these other species. In addition, the number of fish-related stories in Inuit legends is relatively small. The Inuit print reflects the perspective that fish are a part of the community of animals that give both meaning and sustenance to the Inuit and other peoples of the North. Fish are especially important as a relatively dependable and predictable source of day-to-day food, a staple that people can count on, as the Cree would put it, even when other resources fail. Before the Inuit and other northern Indigenous peoples settled into permanent villages in the 1960s and 1970s, many groups moved with the seasons. Seasonal camps were located near good fishing sites especially in the summer months. The choice of sites for permanent villages also reflects this emphasis on fishing. For example, twelve of sixteen Cree villages on the Québec and Ontario sides of James and Hudson Bays are located on the estuary of a river and thus near a supply of fish. Nearly all present-day Inuit villages in the Hudson Bay area are on the coast or an estuary (Baker Lake is a notable exception). This pattern of seeking fish-rich locations is much the same for other Inuit, Inuvialuit, and Gwich’in villages on or near the coasts of Arctic and sub-Arctic Canada. In many cases, location names give away the importance of fishing sites. For example, the name of the capital of Nunavut, Iqaluit, means “the place for fishing.”

Local Knowledge and Fishing The importance of marine and anadromous fishes in the lives of Canadian northern Indigenous peoples is reflected not only in legends, beliefs, and art but also in the detailed environmental knowledge held by many groups. Traditional ecological knowledge (TEK) may be defined as “a cumulative body of knowledge,



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practice and belief, evolving by adaptive processes and handed down through generations by cultural transmission, about the relationship of living beings (including humans) with one another and with their environment” (Berkes 2012, p. 7). Many aspects of TEK are similar to those of scientific knowledge; both kinds of knowledge are based on observations. However, there are also certain fundamental differences. TEK is a knowledge, practice, and belief complex; it is experiential knowledge closely related to a way of life and conditioned by the beliefs held by its practitioners. It is multigenerational and passed on orally by cultural transmission, rather than by book learning. However, it is not static; it evolves. The spiritual aspect of the Indigenous relationship with fish is part of traditional ecological knowledge, and so is the knowledge of when and where to find the fish, their life cycles, natural history, and behaviour. Such knowledge of the fish is an integral part of the practice of fishing. Also integral to this practice is the way in which northern Indigenous peoples organize their fishing; the techniques and technologies they use; and the systems of rules that guide their practice. The spiritual aspects are of importance mainly to anthropologists; distributions and natural history are of interest to biologists; and the systems of fishing fall in the area of human ecology and resource management. The latter two are discussed further here. There are a number of sources that provide a glimpse of the Indigenous knowledge on fish distributions. Many groups of the Arctic and the sub-Arctic focus on different species at different times of the year, in different locations, sometimes using different techniques. For example, in the western Arctic Inuvialuit community of Sachs Harbour, people carry out ice-fishing on lakes in spring, catch Arctic Cod off sea-ice in early summer, and set gill-nets for Arctic Char, Arctic Cod, and Least Cisco in late summer. In the Cree community of Chisasibi, people carry out ice-fishing with nets and set lines in the lakes in winter; set gill-nets for anadromous Lake Whitefish along the retreating ice-edge in spring; set gill-nets for whitefish, anadromous Cisco (Coregonus artedi), and anadromous Brook Trout throughout the summer; and set gill-nets under the ice in the estuary for Cisco in early winter.

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Fig. 45. Fish lure, 1921, Copper Inuit, Coronation Gulf, Nunavut.

Riewe’s (1992) overview of Inuit land use, hunting, and fishing in Nunavut shows a great diversity of patterns of fishery use. Arctic Char is the main species in many areas. The catch may include fully marine species, such as Arctic Cod (often jigged) and sculpins, throughout Nunavut. Regionally there are other species as well; in Cumberland Sound, for example, Atlantic Cod and Greenland Halibut are also caught. Freshwater species are often part of the seasonal round of fishing activities. Anadromous species, as a group, seem to be the most important overall, not only in Nunavut but also in the Inuvialuit region, Hudson and Ungava Bays, and the Labrador coast. In the Mackenzie Delta, for example, the catch is dominated by anadromous Broad Whitefish (Coregonus nasus), Lake Whitefish, Arctic Cisco, and Pacific Herring (Clupea pallasii). In Ungava Bay the two main species, in addition to Arctic Char, are Atlantic Salmon and Brook Trout, both anadromous. Indigenous peoples of the North have a great deal of local knowledge about when and where the fish are found and how to catch them. However, some species are distributed widely, and one does not normally search for them. For example, the Report of the Inuit Land Use and Occupancy Project (Freeman, 1976) notes that, when Inuit fishers were asked to mark sculpin fishing areas, they disdainfully marked a few spots on the map as if to keep researchers happy – they did not look for sculpins; they just caught them. For

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Fig. 46. Sculpin, ivory, Middle Dorset Culture, Mansel Island, Nunavut.

important species, however, a great deal of natural history observation goes into making fishing a fine art: knowing when the fish are beginning to migrate, what they eat and where they find their feed, and how they move with the tide, so that netting can be timed to these movements. Indigenous fishers are good observers of fish biology and life cycles but do not of course have information about the times that the fish are not observable. In the old days many of the traditional fishing techniques required team work. In one of the classical books on Inuit life, Balikci (1970) described the fishing practice and social organization of the Netsilik, a central Arctic group that fished at stone weirs. The weirs were constructed or repaired every year and required the organized labour of an extended family group. In recent decades the use of stone weirs and the importance of social organization in fishing have declined. Much fishing takes place using gill-nets, usually a two-person operation, and increasingly by angling, an individual activity. However, Indigenous fishing is almost never a haphazard activity. Knowing when, where, and how to fish continues to be important. In addition, long-term studies indicate certain features of Indigenous systems of fishing that may be of interest in designing resource management strategies. In a series of papers (summarized in Berkes, 2012, chap. 7) Berkes showed that the

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Fig. 47. Woman with fish lithograph, Egevadluq Ragee.

subsistence fishery of the Chisasibi Cree of James Bay was more productive than comparable commercial fisheries in the sub-Arctic, and apparently had been sustainable since the 1930s. He found that three readily observable sets of locally devised management practices provided insights into the secrets of this sustainable fishery system. The first was the concentration of fishing effort, in space and time, on aggregations of fish. Indigenous fishers were maximizers of the catch per unit effort (CPUE); they monitored CPUE, switching fishing areas and target species. The second practice concerned rotational fishing or pulse fishing. A previously fished area would be rotated or rested so that the CPU (and the populations) would recover before fishers came back again. There is evidence that some Inuit groups practised pulse fishing on char rivers. The third practice involved the use of a mix of gill-net mesh sizes and a mix of catching techniques that removed a variety of size groups and age classes. The practice helps to maintain a mix of reproductive age



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classes that makes the population less vulnerable to reproductive failure in a highly variable environment.

Bridging Science and Traditional Ecological Knowledge Indigenous ecological knowledge poses a dilemma for the biologist. On the one hand, science and TEK have a great deal in common because they are both based on the observation of nature. On the other hand, the ground rules of the two are different. Science has a well-defined set of rules of evidence, and it has no room for the spiritual aspects of TEK. Can the observational components of TEK be separated from the rest of the knowledge, practice, and belief complex and incorporated into science? The consensus view among Indigenous experts and practitioners is that TEK should not be taken out of its cultural context. There is an alternative to trying to

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integrate TEK with science. We can look for ways in which they can enrich one another by sharing, bridging the two systems of knowledge while respecting the integrity of each. The Inuit Observations of Climate Change project illustrates how science and traditional ecological knowledge can be bridged. The five areas of potential convergence, as elucidated by Riedlinger and Berkes (2001) in relation to climate change research, are also relevant to the study of other environmental areas. They postulate the use of traditional knowledge as (1) local-scale expertise, (2) a source of baseline information, (3) potential for formulating research questions and hypotheses, (4) insight into impacts and adaptation, and (5) a basis of community-based environmental monitoring. In areas where they are active on the land, Indigenous peoples tend to generate knowledge of environmental processes at a finer and more detailed scale than do scientists. Such local-scale expertise was important, for example, in detecting the first appearance of two Pacific salmon species in Sachs Harbour in the western Arctic. Such observations can help establish a baseline. The study of environmental change, as in climate-change studies or impact assessments, requires establishing a historical record or a baseline against which change can be measured or assessed. In the Arctic in particular, such baseline data are rarely available from scientific sources. Traditional knowledge, through cumulative experience and oral history, can help fill some of the gaps, as history is embedded in Inuit narratives of animal species, harvesting records, travels, and extreme events. The time depth of Inuit knowledge provides a diachronic perspective, creating a baseline for expected deviations from “normal” conditions. This has proved very useful, in particular, for sea-ice research.

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Through observation of place-specific changes, and assessment of deviation from the expected, traditional ecological knowledge can contribute to research questions and hypotheses. Collaboration at the stage of formulating research objectives establishes a role for communities in research planning. For example, the observation by Dene fishers of liver abnormalities in fish led to the discovery by scientists that some Arctic species had extremely high concentrations of certain organochlorines. The Arctic ecosystem is an exceptionally variable environment, and Arctic peoples are well known by their ability to adapt to changes. In an era of rapid global change, traditional ecological knowledge provides insights into adaptation and change, addressing the issue in the context of food security, livelihoods, and community health. Such considerations suggest that TEK can be thought of as the basis of community-based monitoring in the Arctic and elsewhere. There may be certain advantages to environmental monitoring networks that rely on the observations of experienced and knowledgeable local experts to detect trends and changes, such as the extension of fish distributions, or to monitor long-term impacts of pollutants and development projects against the backdrop of a highly variable environment. Such monitoring can supplement science and may be more cost-effective than scientific research; in addition, it may contribute to fostering TEK by helping maintain the link between Arctic peoples and their environment.

sources:Balikci (1970); Freeman (1976); Johnson (1976a); Lockhart et al. (1987); Riewe (1992); Babaluk, Reist, Johnson, & Johnson (2000); Riedlinger & Berkes (2001); Nichols et al. (2004); Reid et al. (2006); Krupnik et al. (2010); Berkes (2012); Ford et al. (2015); Mistry & Berardi (2016); Rathwell & Armitage (2016).

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FISHERIES James D. Reist

Fish of all types play a significant role in the culture and tradition of Arctic Indigenous peoples. The significance of these fisheries results from their previous and ongoing importance in sustaining human presence in the Canadian Arctic. Marine fisheries also provided sustenance to non-natives, primarily Europeans, during their exploration of the Arctic. More recently, northern fisheries have been seen as both sustaining the traditional lifestyles of the Indigenous peoples and contributing to economic self-sufficiency in the North.

Pre-historical Indigenous Fisheries (18,000 to 300 Years Ago) People were probably present in Beringia near the glacial maximum perhaps as early as thirty thousand years ago. They undoubtedly exploited both freshwater and anadromous fishes of the area, but little direct evidence of this exists. Following deglaciation there were several waves of nomadic or semi-nomadic Arctic peoples who progressively colonized the Canadian Arctic from the west. These included the Paleo-Eskimo Culture (5,000– 3,000 years ago), Dorset Culture (3,000–1,000 years ago), and Thule Culture (1,000–300 years ago). These peoples were primarily associated with coastal areas throughout the Canadian Arctic, and evidence exists of the exploitation of marine fishes. It is quite likely that in most cases the fishing included or was focused upon anadromous fishes such as Arctic Char during their upstream migrations into fresh water. For example, remnants of stone weirs (saputits) and fish spears (leisters) dating to Thule times have been found at many Arctic archaeological sites including those in the High Arctic (e.g., Ruggles River draining Lake Hazen on the north end of Ellesmere Island). The Arctic peoples probably relied heavily upon the exploitation of the marine mammals that were their primary sources of livelihood; however, capture of coastal marine fishes was likely. Thus, exploitation of strictly marine fishes by these peoples, though significant, was not as high as in other Indigenous groups present in the Arctic.



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Although primarily coastal in association, at various times during this colonization and as they do today, Inuit peoples also occupied southern Arctic areas such as Foxe Basin, northern Hudson Bay, northern Québec, and Labrador. Many southern areas considered herein to be Arctic were also colonized by First Nations peoples from the south. Thus, during the later times of deglaciation the southern coastal areas of Hudson and James Bays were colonized by the antecedents of today’s Crees. In addition, many different First Nations groups colonized the inland regions of rivers that drain to the Arctic. They included antecedents of today’s Dene in the Mackenzie River basin, and northern Athapaskans in inland areas west of Hudson Bay. Given that many Arctic anadromous fishes migrate long distances up such rivers, such peoples would have exploited this segment of the Arctic marine fish fauna and perhaps coastal marine forms as well. Due to their ready availability, fish likely made up a correspondingly larger portion of the diet of these inland people as well as of coastal Inuit living in southern areas such as northern Québec. In comparison to today, the effects of these early peoples on local fish populations were probably minimal due to four factors. First, low human-population sizes resulted in correspondingly low needs. Second, a nomadic or semi-nomadic lifestyle resulted in the exploitation pressure on a particular population being limited to short periods of time (usually well within the generation time of the fish). Intervening periods of no exploitation would have allowed for full recovery of any of the effects of the exploitation. Third, limitations of the technology used to capture fish would have limited the rate of exploitation. Finally, during this time environmental effects from southern human societies on the Arctic would have been virtually non-existent. These factors, together with an ingrained respect for the organisms and their environment, ensured that fish populations were minimally perturbed by humans.

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Fisheries during the European Exploration (About 300 to 75 Years Ago) The journals of many of the Europeans who explored the Canadian Arctic contain occasional accounts of marine fishes found in the area. Although usually less explicit, these accounts also contain some information on the fisheries conducted by the explorers. European presence in the area can be divided into three general, but overlapping, periods that correspond to the differing principal aims of the activity: early exploration of the east by sea (e.g., Martin Frobisher, John Davis, and Henry Hudson, 1576–1700), commercial whaling enterprises in the eastern and western seas (ca. 1650–1850), and the search for the Northwest Passage and the lost Franklin expedition through the central Arctic (ca. 1810–75). Corresponding to the marine explorations of the latter two periods, exploration and exploitation of land and freshwater resources also occurred. This primarily involved the fur trade, but exploitation of marine, especially anadromous, Arctic fishes was regionally intensive to support the human populations present in the north. The period from about 1850 to 1925 also included several American expeditions to the north, primarily for whaling in the western Arctic and exploration and research in the eastern Arctic. The primary focus of the expeditions, especially the later ones during the 1800s, was the search for a northwest passage through to the Pacific Ocean. Thus, exploitation of marine fishes was primarily conducted for the sustenance of the expeditions. Similar to the practices of the Indigenous peoples in the area, such exploitation likely focused mostly upon anadromous species; for example, James Clark Ross’s diary of exploration of areas in western Foxe Basin in 1830 refers to fishing trips capturing a great many salmon (i.e., char). Such levels of exploitation were limited in both space and time and had little impact on local fish populations. It is also likely that low levels of exploitation of marine fishes occurred during much of this time in the northern portions of the Labrador Sea and southern Davis Strait. However, little specific data exist on sizes and frequencies of catches. Much of this period saw the exploration and exploitation of land-based resources, which was conducted primarily through the chartering of commercial companies and the awarding of fur-trading rights over large tracts of northern Canada (e.g., Hudson Bay and Northwest companies). These enterprises established a wide system of northern settlements along rivers and northern coastal areas of the mainland. Such posts acted as the focus for trade and local economic development in the North. Significant exploitation of fishery resources occurred, primarily upon anadromous fishes. Locally this exploitation was probably quite high and conducted primarily as food fisheries for both people and dogs. However, some economically based (i.e., commercial) exploitation also occurred and increased towards the end of this period as human populations increased. Also, records are better for more recent fisheries. For example, in the western Arctic the development of trading posts along the northern Mackenzie River valley (e.g., Fort Good Hope in 1836, Fort McPherson in 1840, and Aklavik in 1912), and missionary churches and schools (e.g., Arctic Red River or Tsiigehtchic in

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1901), increased and focused human populations in the area. This in turn increased exploitation of anadromous fishery resources. Initially such exploitation was likely for personal food, but, as populations grew, by the 1900s this exploitation was commercially based, with missionaries, trading companies, and the Northwest Mounted Police all either purchasing fish or hiring fishermen to provide fish. Target species included all anadromous species present in the area, but particular focus was upon Broad Whitefish in areas up to 500 km from the sea. Similar situations existed in the east throughout the low and central Arctic, e.g., Hudson and James Bays, Foxe Basin, northern Québec, and southern Baffin areas. Many of these fisheries appear to have been extensive and were conducted both in nearshore areas and along various river systems. The behaviour of Arctic anadromous fish, especially salmonids, set the stage for such fisheries. That is, the concentrated upstream migrations of the entire populations of these species in late summer and autumn provided for intensive fisheries in both space and seasonality. Furthermore, most populations of anadromous fishes were likely exploited sequentially as they migrated through different areas to spawning locations far upstream. These early circumstances set the basis for the development of fisheries that continue to the present: simultaneous exploitation by different types of fisheries, and locally high levels of exploitation.

Present-Day Fisheries (About 75 Years Ago to the Present) The following provides a general overview of the recent fisheries. Specifics pertinent to each taxon of Arctic marine fishes are enumerated in the “Family and Species Accounts” section. Fisheries of Arctic marine species are very localized and depend upon the suite of species present in the region; thus, although the generalities are similar across the Canadian Arctic, the specifics differ regionally. The human population of the Canadian Arctic is about 40,000 today, most of which (about 70%) is Indigenous and occupies about sixty, primarily coastal, communities throughout the area. Until about 1950 many groups of Inuit maintained a semi-nomadic lifestyle. However, at this time communities were established throughout the area to allow for the centralization of services to northern peoples. Thus, the semi-nomadic lifestyle was supplanted by residence in communities for much of the year along with the establishment of seasonal camps on the land. As a result, the exploitation of fishery resources tends now to be more focused on easily accessible fish populations, usually those near communities. In some cases this has led to over-exploitation of the local resources.

Types of Fisheries Present-day fisheries have been categorized according to various schemes that include the nature of the fish captured (e.g., freshwater, anadromous, marine), the nature of the end use for the fish (e.g., subsistence, commercial, sport), and the general location of

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the fishery (e.g., inshore vs. offshore). Such schemes are developed within the context of some form of fishery management that cannot be dissociated from the fishery. Previous to the settlement of comprehensive Indigenous land claims in the Canadian Arctic beginning in about 1975 in northern Québec, fishery management – and hence regulation of exploitation – was the purview of the relevant government department, primarily federal throughout much of the Arctic. Fisheries that occurred in offshore areas were subject to the relevant international convention and administered federally. With the enactment of land-claim legislation, formal co-management boards have been created and charged with the responsibility for fishery management. With the exception of parts of Hudson Bay and the offshore area of the Labrador Sea, all Arctic areas considered herein are now included under co-management umbrellas. Although included within the Nunavut land claim, offshore fisheries in the eastern Arctic are also regulated in part through the Northwest Atlantic Fisheries Organization (NAFO). Subsistence fisheries are defined as those conducted by an Aboriginal person (i.e., a member of the First Nations, an Inuk, or a person of mixed blood), by angling, nets, set lines, spears, snares, or dip nets for food for him or herself, family, or dogs. These fisheries are essentially unregulated and unlicensed although they are often subject to co-management regulation by the local community or formal board if population problems develop. Domestic fisheries are similar in that the goal is personal sustenance; however, these fisheries are conducted by non-Indigenous persons and are appropriately licensed. Commercial fisheries are those in which the product is sold or bartered. Commercial fisheries can also be designated as “local-sale” or “export,” with the product being sold in the community or sent south for sale. They are regulated by licence, species, gear type, area, seasons, and harvest levels. Moreover, commercial fishery products for export must be landed and processed at a federally approved fish plant. Sport fisheries are those conducted by individuals for recreation and may include both northern residents and non-residents; they are regulated through licensing, species, size, area closures, seasons, and harvest levels. Previous and present fishery policy recognizes that subsistence fisheries take precedence over other types of exploitation, and this is a primary factor in determining other development in the area.

Fisheries for Anadromous Species In the overall context of marine fish species, the most significant Arctic fisheries occur on anadromous fishes principally from the Salmonidae, owing to the widespread occurrence of such fish throughout the Arctic; the establishment of Arctic communities near rivers used by anadromous fishes, often near traditionally used fishing sites; the ease of capture due to the biology of these fish; local, relatively abundant populations; and the relatively limited technology necessary for success. Another major factor is the traditional aspect of anadromous fish exploitation by Indigenous peoples. Virtually all species encountered are exploited, especially in subsistence and domestic fisheries.



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Exploitation of anadromous fishes in subsistence fisheries has likely occurred since human occupation of the area. About twelve thousand Indigenous people from sixty communities participate directly in this type of fishery or benefit from it through the sharing of catches. Harvesting occurs throughout the year, peaking when fish are migrating upstream in fresh water during the late summer and autumn. Anadromous fishes are also exploited during their spring downstream migrations often under river ice, and in coastal and estuarine areas during their summer sojourns at sea. Fish are primarily captured by gill-nets and jigging with lures during both the open-water season and under ice, and in some cases by the more traditional methods of spearing or of using weirs. Harvests are quite variable, and in most cases only limited records are kept; thus, it is difficult to estimate the total harvest. One recent estimate for 1987–8 indicated that about 888,630 kg or 444,000 individual anadromous fishes were captured in Arctic subsistence fisheries during that year. The estimated economic value of this subsistence fishery as protein-equivalent items was about $6 million. Such figures are likely low in comparison to previous levels due to shifts in lifestyle, which include residence in towns and reduced reliance on dog teams for transportation. With the implementation of the requirements mandated under land-claim legislation, surveys of subsistence exploitation have been conducted within the landclaim settlement areas. The survey for the western Arctic Inuvialuit was conducted annually from 1986 to 1997, during which period over one million fish of all types were reported as caught in subsistence fisheries by Inuvialuit fishers from six communities. Seven species of anadromous salmonids in this area represented 77% of this total. Annual reported catches of anadromous fishes varied between 36,000 and 110,000 fish with an average weight of about 2 kg, and the mean number of fishers varied between fifteen and fortythree. Such numbers likely under-estimate the actual subsistence harvest by an unknown but substantial amount, perhaps 100%, due to lack of reporting, under-estimation of catches, and similar factors. These figures also do not include simultaneous catch of some species of anadromous fishes (primarily coregonines) of the western Arctic by other Indigenous groups in areas further upstream in the Mackenzie River basin. Levels of such catches likely approach those captured by the Inuvialuit. Commercial exploitation of anadromous fishes also occurs throughout the Canadian Arctic. Commercial catches may be sold locally or exported from the area and sold directly to southern markets or through the Freshwater Fish Marketing Corporation in Winnipeg. About five hundred people annually participate in commercial fisheries for anadromous fishes that are usually conducted by individuals gill-netting from small boats. The oldest commercial fishery in the Canadian Arctic was for Atlantic Salmon in Ungava Bay, established by the Hudson’s Bay Company; it was conducted in various forms from 1867 to the Second World War and then re-established in 1961. Early average catches were about 21,000 kg annually. Similar-sized commercial fisheries have been sporadically conducted throughout the Canadian Arctic. In most cases the local markets are small, which necessitates export of the product to the south. This entails great costs for transport that, together with

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competitive pricing in the south, often make the fishery economically non-viable. Thus, the typical history of commercial fisheries for Arctic anadromous fishes has been one of subsidized development for a few years, followed by operation for a few years, and then closure due to a lack of profits. Nevertheless, some commercial fisheries have been successful. For example, commercial fisheries for Arctic Char have been conducted more or less continuously in some areas of the Canadian Arctic since 1932. Total annual harvests have varied from 1,000 to 183,000 kg. In 1987–8 the commercial harvest of Arctic Char was at least 142,000 kg. Approximately 42% of this catch was exported south and had a gross value of $659,000. Sport fisheries for anadromous fishes primarily exploit Arctic Char, Brook Trout, and Atlantic Salmon. In 1985 about 16,000 Arctic Char were harvested in sport fisheries throughout the then Northwest Territories (including present-day Nunavut, but excluding provincial records for Arctic areas). These fisheries are usually conducted in fresh waters, although some may be coastal. This seemingly small number belies the actual economic benefit of such sport fishing, which was estimated to be $1.5 million. Much of this comes from value added to the fishery in the form of travel, services, and supplies purchased by anglers, many of whom are from the south. The diversity of fish species in the north tends to be lower than in the south. Combined with regional differences in the species’ complement, this results in all fisheries in a particular area tending to exploit the same mix of taxa. Thus, the potential for over-exploitation is constantly present. It is further complicated by incomplete understanding of the biology, population structure, dynamics, and abundances of most Arctic taxa. Potential over-exploitation is especially acute for many anadromous populations that are easily accessed during the freshwater phases of their life history. The potential for over-exploitation is exacerbated by additional impacts that appear to be significantly affecting Arctic anadromous fishes, including climate change, contaminant deposition, and local habitat changes that result from increased human populations and industrial development of the north.

Fisheries for Wholly Marine Species Marine fish species are also exploited in the Canadian Arctic, typically only in commercial and domestic fisheries. As for other fisheries, exploitation is based upon the availability of various species and therefore varies regionally across the area. Such fisheries generally are local in nature and with one exception are relatively small in scale, being conducted by a few residents. With a few exceptions, most wholly marine fishes found in the Canadian Arctic are relatively small bodied. As a result, substantial commercial fisheries have generally not been developed except for those of the eastern Arctic based on Greenland Halibut. Subsistence harvests of marine fishes in the Arctic are limited by an even greater lack of information than that for anadromous fishes. These fisheries capture about twelve wholly marine species from the following families: Clupeidae (two species), Gadidae (five species),

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Cottidae (two to four species), Cyclopteridae (one species), and Pleuronectidae (two species). Catch and thus the economic value of marine subsistence fisheries cannot be assessed, due to a general lack of information. However, some data are available: in 1987–8 about 30,000 kg of Atlantic Cod and 10,000 kg of Sculpins were harvested in northern Québec. Also, as for subsistence anadromous fisheries, harvests by the Inuvialuit in the western Arctic have been documented through the harvest study. From 1986 to 1997 about 66,000 marine fishes were reported as captured (about 6% of the total reported catch of all fish). These represented five species, with the vast majority (90%) being Pacific Herring; in 1987–8 about 1,880 kg of Pacific Herring were captured by about ten fishers. Arctic Inuit have little history of commercial fishing. Attempted commercial fisheries for wholly marine species, usually fostered by attempts at local economic development, have generally experienced the same boom-and-bust cycle described above for anadromous fisheries. Similar factors have driven this cycle, principally high costs – especially transportation to the south – low economic return, uncertain markets, changing human demography and economy, and limited supply of the resource. An additional factor for most marine fisheries is the need for high initial capital costs, larger boats, larger and more complicated fishing gear, and so forth. A commercial fishery for Atlantic Cod in Killiniq, northern Québec, was begun in 1950 and was active throughout the 1960s, catching up to 27,000 kg of cod, but was closed in 1975 as the settlement was slowly abandoned. Many recent commercial fisheries in the Arctic have been developed through an exploratory fishery process, which recognizes the uncertain information base for most Arctic fishery stocks and involves setting provisional harvest levels upon specific requests from proponents. The early years of such fisheries are intended for assessment of the resource base and usually have an integral biological component; they also involve the development of fishery skills and infrastructure (e.g., processing plants, markets). As the fishery proceeds, the harvest levels are adjusted according to biological findings. Normally used to develop anadromous fisheries, this exploratory process has also been applied to the development of marine fisheries. For example, in the western Arctic an experimental fishery for Pacific Herring, conducted in 1963, harvested about 8,000 kg. In the 1980s it was revived as a roe fishery, and about 4,600 kg of fish (about 400 kg roe) were harvested. However, estimates of the spawning biomass necessary to support a viable fishery are on the order of 1,000 t, and no stock of such size has been found in this region. The exploratory process has been applied to marine fishes in the eastern Arctic with more success. In particular, fisheries based on Greenland Halibut have been developed. These consist of two separate types of commercial fishery: an inshore fishery conducted primarily in fiords of southern Baffin Island, and an offshore fishery conducted in Davis Strait and Baffin Bay (NAFO areas 0A and 0B, since the early 2000s and 1990s respectively). The inshore fishery, beginning in 1986, has typically been conducted in late winter by Indigenous fishers deploying baited long lines through the ice, and although Greenland Halibut is the target species, substantial numbers of Greenland Shark are captured as by-catch. In 1989 about sixty

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individuals participated in this fishery, capturing about 180,000 kg of Greenland Halibut. Recent catches in this fishery are highly variable due to uncertain and unsafe ice conditions. The present (2016) inshore quota for this winter and any summer fisheries in the inshore area is 500 metric tonnes (t). Offshore fisheries were conducted in late summer, originally by trawling but presently by gill-netting and trawling from suitably sized vessels. Similar to offshore fisheries in more southern regions, Greenland Halibut are targeted, but the by-catch includes redfishes (Sebastes spp.), Atlantic Cod, Grenadiers, and Skates. Commercial quotas for the Canadian allocations in areas 0A and 0B for 2016 are 8,000 and 7,000 t, respectively. There are no known sport fisheries conducted for wholly marine species in the Canadian Arctic, although some are likely taken.

Arctic Marine Fisheries in the Canadian Context In the context of Canadian fisheries generally, Arctic marine fisheries are relatively small when measured in simple tonnage and economic terms. For example, in 1987 all Canadian commercial fisheries for all species, including finfish, shellfish, and marine mammals, captured about 1.6 million t, worth about $1.6 billion. The vast majority of these came from fisheries in the Atlantic Ocean. This compares with a commercial harvest of about 780 t for Arctic fisheries, worth about $8 million to the Canadian economy. The figures do not include estimates associated with the subsistence, domestic, or recreational fisheries. As noted, the contributions of these fisheries increase incrementally the simple economic value of Arctic fisheries, most of which results from the protein-replacement value that is primarily derived from anadromous species in subsistence fisheries. To the Arctic the proportional importance of subsistence fisheries is far greater than it is to either the east or the west coast, but is similar to the importance of freshwater fisheries that have a significant domestic component. The relative importance of Arctic marine sport fisheries is very high and parallels that of both marine (i.e., anadromous species) and freshwater sport fisheries in southern areas of Canada. A major difference between Arctic marine fisheries generally, relative to those of southern Canada, is the reliance upon comparatively few species in the Arctic. This is especially true when one considers that almost all Arctic fisheries exploit no more than nine anadromous taxa and only one wholly marine species. In comparison, Atlantic commercial fisheries are dominated by wholly marine finfish species (i.e., 8–10 species of groundfish accounted for 55%, and 4–5 species of pelagic fish for 30%, of the total catch in 1988, measured as tonnage), with virtually no contribution of anadromous taxa. Pacific fisheries tend to be more equitably distributed for groundfish (44% from about 8 species), pelagic fishes (12% from about 3 species), and anadromous fishes (33% from five species).



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Canadian Arctic Marine Fisheries in the Global Arctic Context Estimating the magnitude of Arctic fisheries generally and those of Canada in particular is difficult due to a general paucity of documentation of catches (except for regionally managed commercial species), absence of data for key fisheries (e.g., subsistence fisheries) that may be extensive, confusion regarding taxonomy in the synthesis of statistics, and varying definitions of what constitutes an “Arctic” fishery. A recent attempt based upon Area 18 of the Food and Agriculture Organization of the United Nations (FAO) (i.e., the Arctic Ocean basin only), thus excluding Hudson Bay, Baffin Bay, Davis Strait, and northeast Atlantic waters, extrapolated catches from all types of Arctic fisheries. With these caveats in mind, and the likelihood that significant error exists in such reconstructed estimates, the reconstructed total catch from 1950 to 2006 was estimated as 950,000 t for USSR, U.S., and Canadian fisheries. Of this, 94,000 t (9.9%) was estimated to be Canadian, which is comparable to the estimate for Arctic Alaska (USA) at 89,000 t, and around 12% of that for the USSR (Russia) at 770,000 t. Annual reconstructed catches in the Beaufort Sea peaked at 250 t per year in 1961, declining to 50 t recently, and were dominated by whitefishes (Broad Whitefish and Inconnu). Those for the Arctic Archipelago peaked at 700 t in the 1960s, declining to around 200 t recently, with Arctic Char being the dominant species. For Hudson Bay the catches were around 2,300 t annually in the 1960s, declining to around 600 t recently, with Arctic Char and Atlantic Salmon being the dominant species. Shifts over time are likely due to the replacement of dog teams with mechanized travel over land and the declining need for dog food; that is, they result from shifts within the subsistence component of the overall fishery. These figures do not include commercial catches for the Greenland Halibut fisheries in Baffin Bay and Davis Strait discussed earlier, which are presently around 15,000 t, the total allowable catch allocated to Canada. Moreover, in comparison to fisheries in sub-Arctic seas such as the Bering Sea, Barents Sea, or Greenland Sea, these catches are small.

Impacts other than Fisheries on Arctic Marine Fish In addition to exploitation, Canadian Arctic marine fish populations experience a number of other impacts similar to those experienced by northern fish populations throughout the globe. These include physical and chemical changes in their freshwater and marine environments as a result of industrial development in southern latitudes. Hydro-electric development of rivers flowing to the Arctic affects physical water flows, with consequences for estuarine and nearshore habitats. Chemical changes have resulted from the transmission of contaminants northwards through both atmospheric and hydrologic processes. Thus, despite the seeming remoteness of the Arctic, some species of Arctic marine fishes have very high levels of certain types of persistent contaminants.

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Local industrial development, particularly oil and gas exploration and production, has been the greatest single industrial activity present in the marine and adjacent coastal areas of the Canadian Arctic. This has likely resulted in local impacts on Arctic marine fish populations, but to date they have been relatively limited. Additionally, local mining operations, though few in number, are of increasing interest, and the development of related infrastructure (e.g., harbours and shipping) may affect local populations of marine fish. Human populations in the north are increasing through both high birth rates among native groups and immigration; thus, increased pressure on Arctic marine and anadromous fish populations can be expected from exploitation as well as infrastructure development, for example, increased shipping and increased local sewage disposal. A pervasive impact that is expected to be proportionately greater in the Arctic is that of global change including climate change, related acidification of marine waters, and increased levels of surface ultraviolet radiation due to ozone depletion. Increases in temperature and changes in precipitation are predicted to be very high in the Arctic, which will in turn affect habitats and annual production cycles in the area. Many of these will occur in terrestrial and freshwater environments, the effects of which will also be manifested downstream in marine environments, whereas other effects will directly occur on the marine environment. All of these in turn will directly affect marine fish populations. Among such effects will likely be productivity shifts; for example, if climate warming results in thinning of sea-ice and/or shorter durations of seasonal ice, the productivity of the marine environment will likely decrease due to ice loss and seasonal shifts in ice-associated primary production. Associated shifts in nearshore currents and nutrient-replenishment mechanisms such as upwelling may also occur. Furthermore, as climates ameliorate, southern fish taxa may become established in the Arctic and could have negative effects on existing taxa; this situation may already be occurring in the western Arctic where there are increasingly frequent reports of Pacific Salmons. An ancillary outcome of increased atmospheric concentrations of carbon dioxide (now globally around 400 ppm), a greenhouse gas, is the prospect of the growing acidification of the Arctic Ocean, to which it is especially susceptible given the cold temperatures of the water, the ice cover limiting gas exchange with the atmosphere, and the presently low dissolved levels. Significant effects upon the underlying ecosystem and thus the fishes present may ensue from both drivers.

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Finally, although we tend to view such impacts as individual effects limited in space and time, all the above possibilities including exploitation may cumulate in various ways to affect the same populations of Arctic marine fish. Such cumulative impacts are of particular concern because they may be substantive, yet difficult to demonstrate as having cause-effect relationships, and thus are almost impossible to manage adequately. Such scenarios are further complicated by the fact that generally Arctic fish populations exhibit limited production and relatively slow growth due to their environment. Thus, the capacity for Arctic fish taxa to meet challenges such as these may be limited.

Summary A number of generalities can be made regarding Arctic marine fisheries. First, the complement of species suitable for fisheries is generally limited, consisting primarily of anadromous species and regionally occurring marine species. Second, within a particular region all types of fisheries tend to co-occur, with most targeting the same species and the same stocks within species. This results in potential conflicts in resource allocation among fisheries, with subsistence use being the highest priority. However, subsistence utilization appears to have decreased from the 1970s to the present. Third, the biomass to support sustainable fisheries is often not present, is present erratically or unpredictably, or may be present but is unknown with respect to location. Fourth, limited biomass restricts catch, thereby restricting the development of the infrastructure necessary to support commercial fisheries. Nevertheless, Arctic marine fisheries provide a wide range of benefits to northern Indigenous peoples and Canada at large, including the maintenance of the culture and traditions of northern peoples, food, local self-sufficiency, cash income, and economic development in the north. Arctic fish also provide a number of additional non-consumptive benefits such as educational and scientific benefits, ecological benefits (e.g., species such as Arctic Cod are pivotal in Arctic marine food chains leading to marine mammals), and societal benefits derived from the knowledge that such populations exist.

sources: Corkum & McCart (1981); Crawford (1989); Clarke

(1993); Parsons (1993); Tallman & Reist (1997); Reist, Low, Johnson, & McDowell (2002).

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SCIENTIFIC NAMES Brian W. Coad

Scientific names are an essential retrieval mechanism for information on fishes, in particular because common names vary between localities and between languages and are prone to misunderstandings. Scientific names also reflect relationships between species and are part of a higher classification system. This book follows the higher classification used by Nelson (2006) and the names used by McAllister (1990), Robins et al. (1991), Eschmeyer (1998), Page et al. (2013), and Coad (1995, 2014). These works should be consulted by any serious student of the scientific names of fishes. The scientific name is unique to a single species and is used worldwide. It comprises two words, the first of which is the genus (plural, genera) and the second the species name, species epithet, or trivial name. The genus is always spelled with a capital first letter, and the species name with a lower-case first letter; the name should be printed in italics or underlined. The scientific name always appears in Latin script even where orthography differs as in Inuktitut, Russian, or Japanese. For example, Salvelinus alpinus is the scientific name of the Arctic Char (also known as Alpine Char, Silver Char, Salmon Trout, Hearne’s Salmon, Sea Trout, Coppermine River Salmon, European Char, Hudson Bay Salmon, Mountain Char, Arctic Salmon, Blueback Trout, Greenland Char, Québec Red Trout, omble chevalier,



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Ekalluk, Ikalukpik, Kaloarpok, Ivatarak, Iluuraq, Aniaq, Nutilliq, and many others). The Brook Trout or Char, Salvelinus fontinalis, is a close relative of the Arctic Char and is placed in the same genus Salvelinus, but this is uniquely combined with the species epithet fontinalis to show that we are dealing with a fish that is distinct anatomically and does not normally interbreed with other, related species. Other members of the Salmonidae or Salmon Family are more distantly related to the Arctic Char, and some are placed, for example, in a distinct genus Coregonus that includes whitefishes and ciscoes. The scientific name is followed by the name of the person who first described it scientifically and by the year of the published description: hence, Salvelinus alpinus (Linnaeus, 1758). When the describer and the date are in parentheses as here, this species was originally described in another genus (Salmo, in this case), but it has since been reclassified. A particular species may have two or more scientific names. This can arise when authors are ignorant of each other’s work or genuinely believe that more than one species exists. Subsequent studies may demonstrate that there is, in fact, only one species. The first name published has priority, and the second and subsequent names are called synonyms and are no longer used.

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FISH STRUCTURE Brian W. Coad

This section gives a brief overview of the external structures that are used generally for fish identifications and descriptions. Some internal structures are also described. The glossary gives an alphabetical listing of structures with more explanation, and many are illustrated in the “Keys” section of the book. General body shapes and external structures of a soft-rayed bony fish, a spiny-rayed bony fish, a shark, and a skate are shown in the “Keys” section. The head of a fish comprises several structures. The eyes are generally without eyelids, although sharks have a protective membrane, the nictitating membrane, which acts as an eyelid, and Herrings have a fatty or adipose eyelid partially covering the eye. The nostrils lie on the snout, that part of the head before the eyes. Nostrils do not connect with the mouth cavity. They may consist of a blind sac with one opening, or, most commonly, there is an entryand-exit nostril through which water may flow. Barbels are slender, fleshy structures like feelers on the snout or chin and are used for touch and taste. Some fish can also taste with the aid of sensitive cells in their skin. Sharks and Skates have a small opening near the eye called the spiracle. In Skates this is most useful as a means of taking in water for respiration. Most fish respire by taking water in through the mouth, but the mouth of a skate is on its underside, and inhaling water through it would take in gill-clogging mud and sand. Teeth may be found variously on the tongue, roof, and floor of the mouth, on the bones lining the gape, or even in the throat (see the “Keys” section). On the side of the head behind the eye and over the gills in bony fishes is the gill cover, which is composed of four bones – the opercle or operculum (usually the largest), the preoperculum, the suboperculum, and the interoperculum – all serving to protect the gills. The gill cover, hinged at the front, opens posteriorly and allows inhaled water to pass out after oxygen and carbon dioxide have been exchanged at the gills. Sharks have a row of gill slits behind the eye, and Skates have them on the underside of the head. The cheek is the area just below the eye, and between the eye and the gill cover. The head may be variously armed and ornamented by spines and fleshy tabs. The branchiostegal membrane is found below each gill cover, supported by thin slivers of bone or branchiostegals, and connected

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with the gill cover on the other side of the head. Under the gill cover or behind the gill slits lie the gills, which are made up of an arch of bone or cartilage supporting the gill filaments behind the gill rakers (see the “Keys” section). Rakers may be short and widely spaced in fish whose food items are large and easily deflected, or long and close together in fish whose food items are minute, like plankton, and are sieved from the water. The head is joined directly to the body; there is no neck. The body consists mostly of trunk. The caudal peduncle starts behind the anal fin and ends at the tail or caudal fin. The number and presence of different types of fins on the body varies with the species of fish and is often a useful character for identification. The back may carry one to three dorsal fins and an adipose fin between the dorsal fin and the tail fin. The tail (or caudal) fin is at the end of the body and may be forked, square cut, rounded, pointed, or lunate. Its skeletal structure may be almost symmetrical or upturned at the end. This upturn is obvious in sharks that also have a large upper lobe to the fin and a small lower lobe. The anal fin, or fins, lies on the underside of the body behind the vent that is the exit for the intestine, kidney ducts, and gonads. The pectoral fins are found behind the gill cover on each side of the body, and a pair of pelvic fins is behind, below, or in front of the pectorals on the lower body. An axillary pelvic scale above the pelvic fin in some fishes streamlines the fin when the fin is pressed against the body. Certain fishes have the pelvic fins modified as an adhesive sucker or disc, while others lack pelvic fins entirely, for example, the Snailfish Family (Liparidae). All the fins except the fleshy adipose fin are supported by rays. Soft rays are flexible and jointed, and spines are rigid, pointed, and unjointed. The number of spines and soft rays in the various fins are useful for identification purposes. Most fishes have a body covering of scales that may extend onto the head and certain fins. Rounded, smooth scales are called cycloid and are found in the less advanced bony fishes. Large cycloid scales may easily detach, as in Herrings, but small cycloid scales can be embedded and hard to see. Ctenoid scales bear small teeth on the posterior margin and feel rough to the touch. Such scales are found in the more advanced bony fishes. Sharks and Skates have placoid

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scales that can be so rough as to scrape the skin off a human. Skates have spines and thorns (large spines) on the head, body, and tail. Most fish have a distinctive colour pattern, but this can change with age, maturity, behaviour, background, between sexes, and after death. Fishes have a lateral-line system that runs along the flank and a similar system on the head. The extent and development of these systems varies with the species of fish. The lateral line is a tube in the skin with openings to the outside through pores in the scales. A lateral-line pore count is often used in separating fish species. The tube contains sensory cells that serve the fish in detecting low-frequency vibrations, currents, and waves and, in sharks and Skates only, temperature changes. Light organs are found in some species that live in deeper and darker waters. The number and disposition of these organs on the head and body are often critical in identification. The organs are individually and, when in rows or groups, collectively named with arcane abbreviations. These are explained in the “Keys” section and in the “Family and Species Accounts” section for deep-sea fishes, for example, Myctophidae. Internally, the mouth cavity narrows to an oesophagus that passes to a straight, U-shaped, or J-shaped stomach. Pyloric caeca may be attached at the junction of the stomach and the intestine in some fishes, and these blind sacs aid digestion by adding enzymes to the food. Their number may aid in species identification. The intestine may be a short S-shaped loop or an extremely long and complex coil. It ends at the vent that is seen on the abdomen, usually in front of the anal fin but, in some fishes, far anterior. Fish have a liver, an orange or reddish organ, at the front of the body cavity. In sharks this may be very large and form a significant part of the body weight. The gas bladder (swimbladder) is a gas-filled sac with thin walls lying near the top of the body cavity. It functions as a buoyancy organ and in some fishes is used to transmit sounds to the inner ear or to produce sounds by means of special muscles. Just below the backbone, above the gas bladder, are two long, dark-coloured



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kidneys that are connected to a small urinary bladder at the end of the body cavity. The two kidneys may appear as an indistinguishable black mass below the backbone. Below the kidneys are the ovaries, which may be filled with eggs, or the testes, which produce the sperm. The body cavity is lined with a membrane that may vary in colour from silvery white to jet black. The main body muscles are in the form of Σ-shaped, interlocking blocks and their contractions help to produce the sinuous body movements by which fish swim. Sharks and Skates have somewhat different structures from those of bony fishes. Some species produce living young rather than eggs, and in others the embryo is laid in a horny egg case, known as a mermaid’s purse when it washes up on a beach. Male sharks and Skates have claspers derived from the pelvic fins, which serve to deliver the sperm to the female during copulation. The length of time that food stays in the gut of sharks is increased by a spiral valve. The food follows the spiral around rather than straight through the gut, and so there is more time for digestion and more surface for absorption. The pectoral fins of Skates are greatly enlarged and enable them to “fly” through the water. There is no gas bladder in sharks and Skates, which have to swim constantly to stay above the bottom. Sharks and Skates produce teeth in multiple rows, and as older teeth at the front of the jaw fall out or are broken off, new ones move forward to replace them. The skeleton of a bony fish is illustrated in the “Keys” section. It includes the skull, comprising the cranium (which contains the brain), jaws, gill arches, operculum, and other associated bones. The cranium also contains small objects known as otoliths in the inner ear. These and their associated structures aid in sensing change of direction and in maintaining balance. There is a vertebral column with ribs anteriorly enclosing and protecting the body cavity and its contents. A tail skeleton supports the tail fin, and the pectoral and pelvic girdles support their respective fins. There are fin supports too for the dorsal and anal fins. Sharks and Skates have a skeleton composed of cartilage, a substance that is not as strong as bone but quite hard when impregnated with salts.

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COLLECTING AND PRESERVING FISHES Brian W. Coad

The collecting methods for fishes are many and varied. An entry to this field is given by Coad (1995), which describes some of the more elementary techniques and has a list of references for others. The deep-sea fishes are only accessible to research vessels using specialized, heavy equipment. The most fragile species from these deeper waters are often damaged when they are brought on deck and may be difficult to identify in this state. Captured fishes that cannot be identified or seem unusual enough to warrant further attention should be preserved. Labelled, preserved specimens that are deposited in a museum are a permanent record of species identity and distribution. Some taxa present problems of identification even for experts, and misidentifications are often a nuisance if there is no material to examine. Samples from ecological or experimental studies as well as systematic and distributional works may be preserved and sent to a museum where their identity can be confirmed and where they are available to workers in the future. Some species are rarely caught, and it can take many years to build up collections of sufficient size to enable the resolution of systematic and taxonomic problems. This applies particularly to the more remote and difficult-to-sample areas such as the Arctic. Specimens should be preserved whole, without removal of the guts or gills, so that no key characters are lost. Specimens may be frozen, but the best method is to drop fish into one part of fullstrength formalin to kill the fish quickly and then immediately add nine parts of water to form a 10% preserving solution. Anaesthetics (MS222) can be used before immersion in preservative. Large specimens (larger than about 15 cm) should have a small slit made in the right side of the belly after death to allow formalin to penetrate the tissues. Ichthyologists cut the right side of the fish and leave the left side undamaged for illustration and scale counting. Hypodermic syringes are used to inject the abdominal cavity and muscle blocks of very large fish with formalin; otherwise the preservative will not penetrate all the tissues before decay sets in. Syringes should have a capacity of up to 100 milliliters and be capable of taking needles of various sizes. Particular care should be taken when injecting formalin into tissues; the needle should be withdrawn gradually while

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injecting the formalin solution to avoid a sudden spurt of liquid under pressure from the injection site. Formalin should be handled with care as it is a noxious chemical that irritates the eyes and nose and is painful in skin cuts. It may be carcinogenic, and repeated exposure can trigger allergic reactions in the skin. Gloves and safety glasses are useful when diluting full-strength formalin. It should only be handled in well-ventilated rooms or in the open air. In the field, care should be exercised in packing specimens for transport so that leakages do not occur. Long-term preservation in formalin is not advisable as the solution becomes acidic, decalcifies bones, and rots the fish. It also wrinkles and hardens the specimens. Specimens preserved in formalin cannot generally be used for recovery of DNA. Freezing or 95% ethanol is better for that purpose although less desirable for morphology and long-term storage. Often DNA samples are taken from captured fish before formalin preservation. Most museums store their specimens in alcohol for the long term. The formalin-fixed specimens are washed briefly in water and then transferred to 45% isopropyl alcohol or preferably 70% ethanol. These chemicals are more pleasant to work with. Some care should be taken so that specimens are not twisted and bent inside the preserving container. It is difficult to make the counts and the measurements necessary for identification on badly deformed specimens. Each specimen or group of specimens should have at least an equal volume of preservative because water in the fish tissues tends to dilute the preserving fluid. Specimens may be stepped through solutions of 30%, 50%, and 70% alcohol to reduce wrinkling and ensure a fuller penetration of alcohol into tissues and a final storage solution of at least 70% ethanol. The best containers for long-term storage are made of glass with tightly sealing polypropylene lids. Plastic containers deteriorate with time and tend to crack. Metal containers and metal lids eventually rust. In the field, large plastic buckets with tightly sealed lids are less likely to break than glass containers and are not as heavy. Very large fish may require some sort of drum, such as a clean oil drum, but it should be noted that formalin corrodes metal, and the

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drums should be lined with plastic or lacquered. Fluid levels in the collection should be checked regularly, and alcohol concentrations maintained at the recommended values, or the specimens will deteriorate. Collections should be kept in the dark to reduce fading of pigments and at a constant, cool temperature. Fish that have been preserved for a week (more for larger fishes) in formalin or transferred to alcohol can be sent to a museum for identification. Glass containers full of formalin or alcohol should not be mailed because of the danger of breakage. The fish should be wrapped in cheesecloth or some other absorbent packaging, and the packaging dampened with preservative. The label must be included with the fish. The package is then tightly sealed in several, leak-proof plastic bags before being placed in a padded box for mailing. Spiny fish should be especially well wrapped to avoid puncturing the plastic bags. A tightly sealed package retains the preservative that keeps the fish in good condition. The box should be labelled according to the appropriate regulations, for example, “Research Specimens, Not Restricted, Special Provision A180 applies,” and the shipping documentation state that “this package contains dead preserved fishes for scientific research, which were originally pre-fixed in a formalin solution and then transferred into 70% ethanol for further preservation; these specimens are not infectious due



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to the original preservation technique … They have no commercial value, and are not for resale.” The label is as important as the fish itself. An interesting specimen is of little or no scientific value if there is no locality data. Labels should be written at the time of capture. Faulty memory and good intentions to label specimens later make a poor combination and often result in collections with no data or, worse, with incorrect data. The label should bear the place of capture, including a reference to the nearest inhabited place or major geographical feature, latitude and longitude (a Global Positioning System, GPS, can be used), province or territory, date, name of collector, notes on the habitat and live colour of the specimens, and any other items likely to be useful. Colour photographs of fresh fish are most useful, especially if the fins are pinned erect. Pencil or India ink should be used on stout, waterproof paper that will not disintegrate in liquid. The label must be dropped in the jar with the fish. Labels on the outside of jars always fall off, and lids with labels always get put on the wrong jar! Collectors should check with the museum to which they wish to send fish. Identification, cataloguing, and permanent storage of specimens is expensive, and it is not always possible for a museum to accept extensive and unexpected collections.

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Checklists of Species

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MARINE SPECIES Brian W. Coad

Marine species are those that spend all or a significant part of their life in the sea. Some species, notably members of the Trouts and Salmons Family (Salmonidae), can live wholly in fresh water or spend part of their life in the sea and return to fresh water to breed. All such species with a marine stage in their life are included here. Some species of fish in Canada are primarily freshwater species but do have a tolerance (of varying degrees) for saline conditions and are listed separately below and are not covered by this book. Spelling of scientific names, authors, and dates have been checked against the American Fisheries Society (AFS) list (Page et al., 2013) as the primary source for these names in North America and also for English and French common names. Page et al. (2013) gives explanations for recent taxonomic changes and common-name usage. Earlier changes can be tracked in previous editions of the work. However, the AFS list “exclude[s] species known only from beyond continental shelf waters over bottoms exceeding 200 m, even if found in the midwater of less than 200 m” and does not always have French names. Scientific names for these excluded species were taken from Eschmeyer (1998; 11 February 2013 online version), which is an entry point to the literature on scientific names that is not reviewed extensively here.

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The source for English and French common names that are not in the AFS list is Coad, with Waszczuk and Labignan (1995), and then FishBase (Froese & Pauly, 2013) for fish not in that book. Deviations from names used in Coad et al. (1995) are explained in the introduction to the “Family and Species Accounts” section, under “Taxonomy.” In the absence of “official” common names, the choice of name to use is somewhat arbitrary and varies between sources. There are several countries that use English or French common names, or slightly different spellings of similar names. The “common” names of deep-sea species are only common in contrast to the scientific name and are not names in common or popular use. The fishes are only seen by scientists on research vessels or working in museum collections. Some French common names of deep-sea species were coined for this work, and these are indicated in the “Family and Species Accounts” section. In a few instances, recognition of species varies from the sources listed above, and this is discussed in the “Family and Species Accounts” section. The principal differences are Gadus ogac, which is recognized as a distinct species, and the retention of (but with questions about) some zoarcid species.

Checklists Of Sp ecies

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The marine species of Arctic Canada are as follows: Family 1. Myxinidae – Hagfishes, Myxines 1. Myxine glutinosa Linnaeus, 1758 – Atlantic Hagfish, myxine du nord

Family 9. Acipenseridae – Sturgeons, Esturgeons 1. Acipenser fulvescens Rafinesque, 1817 – Lake Sturgeon, esturgeon jaune

Family 2. Petromyzontidae – Lampreys, Lamproies 1. Lethenteron camtschaticum (Tilesius, 1811) – Arctic Lamprey, lamproie arctique

Family 10. Notacanthidae – Deep-sea Spiny Eels, Poissons-tapirs à épines 1. Notacanthus chemnitzii Bloch, 1788 – Snubnosed Spiny Eel, tapir à grandes écailles 2. Polyacanthonotus rissoanus (de Filippi and Vérany, 1857) – Shortspine Tapirfish, tapir à petites épines

Family 3. Rhinochimaeridae – Longnose Chimaeras, Chimères à long nez 1. Harriotta raleighana Goode and Bean, 1895 – Longnose Chimaera, chimère-spatule 2. Rhinochimaera atlantica Holt and Byrne, 1909 – Knifenose Chimaera, chimère-couteau

Family 11. Synaphobranchidae – Cutthroat Eels, Anguilles égorgées 1. Simenchelys parasitica Gill, 1879 – Snubnose Eel, anguille à nez court 2. Synaphobranchus kaupii Johnson, 1862 – Northern Cutthroat Eel, anguille égorgée bécue

Family 4. Chimaeridae – Shortnose Chimaeras, Chimères 1. Hydrolagus affinis (de Brito Capello, 1868) – Deepwater Chimaera, chimère de profondeur

Family 12. Nemichthyidae – Snipe Eels, Poissons-avocettes 1. Nemichthys scolopaceus Richardson, 1848 – Slender Snipe Eel, avocette ruban

Family 5. Scyliorhinidae – Cat Sharks, Roussettes 1. Apristurus profundorum (Goode and Bean, 1896) – Deepsea Cat Shark, roussette de profondeur

Family 13. Serrivomeridae – Sawpalates, Serrivomers 1. Serrivomer beanii Gill and Ryder, 1883 – Stout Sawpalate, serrivomer trapu

Family 6. Etmopteridae – Lantern Sharks, Requins-lanternes 1. Centroscyllium fabricii (Reinhardt, 1825) – Black Dogfish, aiguillat noir

Family 14. Saccopharyngidae – Swallowers, Avaleurs 1. Saccopharynx ampullaceus (Harwood, 1827) – Taillight Gulper, avaleur feu-arrière

Family 7. Somniosidae – Sleeper Sharks, Somniosidés 1. Centroscymnus coelolepis Barbosa du Bocage and de Brito Capello, 1864 – Portuguese Shark, pailona 2. Somniosus microcephalus (Bloch and Schneider, 1801) – Greenland Shark, laimargue atlantique 3. Somniosus pacificus Bigelow and Schroeder, 1944 – Pacific Sleeper Shark, laimargue du Pacifique

Family 15. Eurypharyngidae – Gulpers, Grandgousiers 1. Eurypharynx pelecanoides Vaillant, 1882 – Pelican Gulper, grandgousier pélican

Family 8. Rajidae – Skates, Raies 1. Amblyraja hyperborea (Collett, 1879) – Darkbelly Skate, raie boréale 2. Amblyraja jenseni (Bigelow and Schroeder, 1950) – Shorttail Skate, raie à queue courte 3. Amblyraja radiata (Donovan, 1808) – Thorny Skate, raie épineuse 4. Bathyraja sp. – unknown 5. Bathyraja spinicauda (Jensen, 1914) – Spinytail Skate, raie à queue épineuse 6. Malacoraja spinacidermis (Barnard, 1923) – Soft Skate, raie molle 7. Rajella bathyphila (Holt and Byrne, 1908) – Abyssal Skate, raie bathyale 8. Rajella fyllae (Lütken, 1887) – Round Skate, raie ronde 9. Rajella lintea (Fries, 1838) – Linen Skate, raie linon



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Family 16. Clupeidae – Herrings, Harengs 1. Clupea harengus Linnaeus, 1758 – Atlantic Herring, hareng atlantique 2. Clupea pallasii Valenciennes, 1847 – Pacific Herring, hareng du Pacifique Family 17. Argentinidae – Argentines, Argentines 1. Argentina silus (Ascanius, 1775) – Atlantic Argentine, grande argentine Family 18. Microstomatidae – Pencilsmelts, Microbecs  1. Bathylagus euryops Goode and Bean, 1896 – Goitre Blacksmelt, garcette-goître Family 19. Platytroctidae – Tubeshoulders, Circés 1. Holtbyrnia anomala Krefft, 1980 – Bighead Searsid, tube-épaule grosse tête 2. Maulisia mauli Parr, 1960 – Maul’s Searsid, tube-épaule de Maul 3. Maulisia microlepis Sazonov and Golovan, 1976 – Smallscale Searsid, tube-épaule petites écailles

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4. Normichthys operosus Parr, 1951 – Longsnout Manypitshoulder, épaule-criblée long nez 5. Platytroctes apus Günther, 1878 – Legless Searsid, tube-épaule apode

Family 20. Alepocephalidae – Slickheads, Alépocéphales 1. Alepocephalus agassizii Goode and Bean 1883 – Dusky Slickhead, alépocéphale obscur 2. Alepocephalus bairdii Goode and Bean, 1879 – Manyray Smoothhead, alépocéphale multirai 3. Bajacalifornia megalops (Lütken, 1898) – Bigeye Smoothhead, alépocéphale à grands yeux 4. Bathytroctes sp. – unknown 5. Rouleina maderensis Maul, 1948 – Madeiran Smoothhead, alépocéphale de Madère 6. Xenodermichthys copei (Gill, 1884) – Atlantic Gymnast, gymnaste atlantique Family 21. Osmeridae – Smelts, Éperlans 1. Mallotus villosus (Müller, 1776) – Capelin, capelan 2. Osmerus dentex Steindachner and Kner, 1870 – Pacific Rainbow Smelt, éperlan du Pacifique 3. Osmerus mordax (Mitchill, 1814) – Rainbow Smelt, éperlan arc-en-ciel Family 22. Salmonidae – Trouts and Salmons, Truites et Saumons 1. Coregonus artedi Lesueur, 1818 – Cisco, cisco de lac 2. Coregonus autumnalis (Pallas, 1776) – Arctic Cisco, cisco arctique 3. Coregonus clupeaformis (Mitchill, 1818) – Lake Whitefish, grand corégone 4. Coregonus nasus (Pallas, 1776) – Broad Whitefish, corégone tschir 5. Coregonus sardinella Valenciennes, 1848 – Least Cisco, cisco sardinelle 6. Oncorhynchus gorbuscha (Walbaum, 1792) – Pink Salmon, saumon rose 7. Oncorhynchus keta (Walbaum, 1792) – Chum Salmon, saumon kéta
 8. Oncorhynchus kisutch (Walbaum 1792) – Coho Salmon, saumon coho 9. Oncorhynchus nerka (Walbaum, 1792) – Sockeye Salmon, saumon rouge 10. Oncorhynchus tshawytscha (Walbaum, 1792) – Chinook Salmon, saumon chinook 11. Prosopium cylindraceum (Pennant, 1784) – Round Whitefish, ménomini rond 12. Salmo salar Linnaeus, 1758 – Atlantic Salmon, saumon atlantique 13. Salvelinus alpinus (Linnaeus, 1758) – Arctic Char, omble chevalier 14. Salvelinus fontinalis (Mitchill, 1814) – Brook Trout, omble de fontaine 15. Salvelinus malma (Walbaum, 1792) – Dolly Varden, omble malma 16. Salvelinus namaycush (Walbaum, 1792) – Lake Trout, touladi 17. Stenodus leucichthys (Güldenstädt, 1772) – Inconnu, inconnu

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Family 23. Gonostomatidae – Bristlemouths, Cyclothones 1. Cyclothone microdon (Günther, 1878) – Veiled Anglemouth, cyclothone à petites dents 2. Sigmops bathyphilus (Vaillant, 1884) – Spark Anglemouth, gonostome étincelé Family 24. Sternoptychidae – Marine Hatchetfishes, Haches d’argent 1. Argyropelecus gigas Norman, 1930 – Greater Silver Hatchetfish, grande hache d’argent Family 25. Stomiidae – Dragonfishes, Dragons à écailles 1. Astronesthes cf. richardsoni (Poey, 1852) – Richardson’s Snaggletooth, dragon-saumon de Richardson 2. Borostomias antarcticus (Lönnberg, 1905) – Large-eye Snaggletooth, dragon-saumon à grands yeux 3. Chauliodus sloani Bloch and Schneider, 1801 – Manylight Viperfish, chauliode très-lumineux 4. Malacosteus niger Ayres, 1848 – Stoplight Loosejaw, drague rouge-verte 5. Rhadinesthes decimus (Zugmayer, 1911) – Slender Snaggletooth, dragon-saumon élancé 6. Stomias boa (Risso, 1810) – Boa Dragonfish, dragon-boa. Subspecies Stomias boa ferox Reinhardt, 1842 Family 26. Notosudidae – Waryfishes, Guetteurs 1. Scopelosaurus lepidus (Krefft and Maul, 1955) – Blackfin Waryfish, guetteur à nageoire noire Family 27. Paralepididae – Barracudinas, Lussions 1. Anotopterus pharao Zugmayer, 1911 – Daggertooth, pharaon 2. Arctozenus risso (Bonaparte, 1840) – White Barracudina, lussion blanc 3. Magnisudis atlantica (Krøyer, 1868) – Duckbill Barracudina, lussion à bec de canard 4. Paralepis coregonoides Risso, 1820 – Sharpchin Barracudina, lussion à menton. Subspecies Paralepis coregonoides borealis Reinhardt, 1837 Family 28. Myctophidae – Lanternfishes, Poissons-lanternes 1. Benthosema glaciale (Reinhardt, 1837) – Glacier Lanternfish, lanterne glaciaire 2. Lampanyctus crocodilus (Risso, 1810) – Jewel Lanternfish, lanterne-joyau 3. Lampanyctus intricarius Tåning, 1928 – Diamondcheek Lanternfish, lanterne à joue pailletée 4. Lampanyctus macdonaldi (Goode and Bean, 1896) – Rakery Lanternfish, lanterne-bouée râtelière 5. Myctophum punctatum Rafinesque, 1810 – Spotted Lanternfish, lanterne ponctuée 6. Notoscopelus kroyeri (Malm, 1861) – Northern Saillamp, lampe-voilière du nord 7. Protomyctophum arcticum (Lütken, 1892) – Arctic Telescope, télescope arctique

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8. Symbolophorus veranyi (Moreau, 1888) – North Atlantic Cornerlantern, lanterne-de-coin nord-atlantique

Family 29. Macrouridae – Grenadiers, Grenadiers 1. Coryphaenoides armatus (Hector, 1875) – Russet Grenadier, grenadier roux 2. Coryphaenoides brevibarbis (Goode and Bean, 1896) – Shortbeard Grenadier, grenadier à barbe courte 3. Coryphaenoides carapinus Goode and Bean, 1883 – Carapine Grenadier, grenadier à barbillon court 4. Coryphaenoides guentheri (Vaillant, 1888) – Günther’s Grenadier, grenadier de Günther 5. Coryphaenoides rupestris Gunnerus, 1765 – Rock Grenadier, grenadier de roche 6. Gadomus longifilis (Goode and Bean, 1885) – Threadfin Grenadier, grenadier à filaments 7. Macrourus berglax Lacepède, 1801 – Roughhead Grenadier, grenadier berglax 8. Nezumia bairdii (Goode and Bean, 1877) – Marlin-spike, grenadier du Grand Banc 9. Trachyrincus murrayi Günther, 1887 – Roughnose Grenadier, grenadier-scie

Family 34. Lophiidae – Goosefishes, Baudroies 1. Lophius americanus Valenciennes, 1837 – Goosefish, baudroie d’Amérique Family 35. Himantolophidae – Footballfishes, Poissons-football 1. Himantolophus groenlandicus Reinhardt, 1837 – Atlantic Footballfish, football fine-lampe Family 36. Oneirodidae – Dreamers, Rêveurs 1. Chaenophryne longiceps Regan, 1925 – Can-opener Smoothdream, doux-rêve ouvre-boîte 2. Oneirodes sp. – unknown 3. Spiniphryne gladisfenae (Beebe, 1932) – Prickly Dreamer, rêveur piquant Family 37. Ceratiidae – Seadevils, Poissons-pêcheurs 1. Ceratias holboelli Krøyer, 1845 – Northern Giant Seadevil, pêcheur à deux massettes Family 38. Gigantactinidae – Whipnoses, Tacts géants 1. Gigantactis vanhoeffeni Brauer, 1902 – Vanhoeffen’s Whipnose, tact géant de Vanhoeffen

Family 30. Moridae – Codlings, Moros 1. Antimora rostrata (Günther, 1878) – Blue Antimora, antimore bleu 2. Halargyreus johnsonii Günther, 1862 – Dainty Mora, more délicat 3. Lepidion eques (Günther, 1887) – Largeye Lepidion, lépidion à grands yeux

Family 39. Melamphaidae – Ridgeheads, Poissons-heaumes 1. Scopeloberyx robustus (Günther, 1887) – Longjaw Bigscale, tête-à-crète robuste

Family 31. Phycidae – Phycid Hakes, Phycidés 1. Enchelyopus cimbrius (Linnaeus, 1766) – Fourbeard Rockling, motelle à quatre barbillons 2. Gaidropsarus argentatus (Reinhardt, 1837) – Silver Rockling, mustèle argentée 3. Gaidropsarus ensis (Reinhardt, 1837) – Threebeard Rockling, mustèle arctique à trois barbillons 4. Phycis chesteri Goode and Bean, 1878 – Longfin Hake, merluche à longues nageoires

Family 41. Trachichthyidae – Slimeheads, Hoplites 1. Hoplostethus atlanticus Collett, 1889 – Orange Roughy, hoplostète orange

Family 32. Gadidae – Cods, Morues 1. Arctogadus glacialis (Peters, 1872) – Polar Cod, saïda imberbe 2. Boreogadus saida (Lepechin, 1774) – Arctic Cod, saïda franc 3. Brosme brosme (Ascanius, 1772) – Cusk, brosme 4. Eleginus gracilis (Tilesius, 1810) – Saffron Cod, navaga jaune 5. Gadus morhua Linnaeus, 1758 – Atlantic Cod, morue franche 6. Gadus ogac Richardson, 1836 – Greenland Cod, ogac 7. Lota lota (Linnaeus, 1758) – Burbot, lotte 8. Micromesistius poutassou (Risso, 1827) – Blue Whiting, merlan bleu

Family 43. Scorpaenidae – Scorpionfishes, Scorpènes 1. Sebastes fasciatus Storer, 1854 – Acadian Redfish, sébaste acadien 2. Sebastes mentella Travin, 1951 – Deepwater Redfish, sébaste atlantique 3. Sebastes norvegicus (Ascanius, 1772) – Golden Redfish, sébaste orangé

Family 33. Bythitidae – Viviparous Brotulas, Donzelles vivipares 1. Bythites fuscus Reinhardt, 1837 – Arctic Brotula, donzelle arctique



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Family 40. Anoplogastridae – Ogrefishes, Ogres 1. Anoplogaster cornuta (Valenciennes, 1833) – Fangtooth, ogre

Family 42. Gasterosteidae – Sticklebacks, Épinoches 1. Gasterosteus aculeatus Linnaeus, 1758 – Threespine Stickleback, épinoche à trois épines 2. Pungitius pungitius (Linnaeus, 1758) – Ninespine Stickleback, épinoche à neuf épines

Family 44. Cottidae – Sculpins, Chabots 1. Artediellus atlanticus Jordan and Evermann, 1898 – Atlantic Hookear Sculpin, hameçon atlantique 2. Artediellus scaber Knipowitsch, 1907 – Hamecon, hameçon rude 3. Artediellus uncinatus (Reinhardt, 1835) – Arctic Hookear Sculpin, hameçon neigeux

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4. Gymnocanthus tricuspis (Reinhardt, 1830) – Arctic Staghorn Sculpin, tricorne arctique 5. Icelus bicornis (Reinhardt, 1840) – Twohorn Sculpin, icèle à deux cornes 6. Icelus spatula Gilbert and Burke, 1912 – Spatulate Sculpin, icèle spatulée 7. Myoxocephalus aenaeus (Mitchill, 1814) – Grubby, chaboisseau bronzé 8. Myoxocephalus octodecemspinosus (Mitchill, 1814) – Longhorn Sculpin, chaboisseau à dix-huit épines 9. Myoxocephalus quadricornis (Linnaeus, 1758) – Fourhorn Sculpin, chaboisseau à quatre cornes 10. Myoxocephalus scorpioides (Fabricius, 1780) – Arctic Sculpin, chaboisseau arctique 11. Myoxocephalus scorpius (Linnaeus, 1758) – Shorthorn Sculpin, chaboisseau à épines courtes 12. Triglops murrayi Günther, 1888 – Moustache Sculpin, faux-trigle armé 13. Triglops nybelini Jensen, 1944 – Bigeye Sculpin, faux-trigle aux grands yeux 14. Triglops pingelii Reinhardt, 1837 – Ribbed Sculpin, faux-trigle bardé

Family 45. Agonidae – Poachers, Poissons-alligators 1. Aspidophoroides monopterygius (Bloch, 1786) – Alligatorfish, poisson-alligator atlantique 2. Aspidophoroides olrikii Lütken, 1877 – Arctic Alligatorfish, poisson-alligator arctique 3. Leptagonus decagonus (Bloch and Schneider, 1801) – Atlantic Poacher, agone atlantique Family 46. Psychrolutidae – Fathead Sculpins, Chabots veloutés 1. Cottunculus microps Collett, 1875 – Polar Sculpin, cotte polaire 2. Cottunculus thomsonii (Günther, 1882) – Pallid Sculpin, cotte blême 3. Psychrolutes phrictus Stein and Bond, 1978 – Giant Blob Sculpin, chabot maculé Family 47. Cyclopteridae – Lumpfishes, Poules de mer 1. Cyclopteropsis jordani Soldatov, 1929 – Smooth Lumpfish, petite poule de mer douce 2. Cyclopteropsis mcalpini (Fowler, 1914) – Arctic Lumpsucker, petite poule de mer McAlpine 3. Cyclopterus lumpus Linnaeus, 1758 – Lumpfish, grosse poule de mer 4. Eumicrotremus derjugini Popov, 1926 – Leatherfin Lumpsucker, petite poule de mer arctique 5. Eumicrotremus spinosus (Fabricius, 1776) – Atlantic Spiny Lumpsucker, petite poule de mer atlantique Family 48. Liparidae – Snailfishes, Limaces de mer 1. Careproctus kidoi Knudsen and Møller, 2008 – Kido’s Snailfish, limace de Kido

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2. Careproctus longipinnis Burke, 1912 – Longfin Snailfish, limace à longues nageoires 3. Careproctus reinhardti (Krøyer, 1862) – Sea Tadpole, petite limace de mer 4. Liparis atlanticus (Jordan and Evermann, 1898) – Atlantic Seasnail, limace atlantique 5. Liparis fabricii Krøyer, 1847 – Gelatinous Seasnail, limace gélatineuse 6. Liparis gibbus Bean, 1881 – Variegated Snailfish, limace marbrée 7. Liparis tunicatus Reinhardt, 1836 – Kelp Snailfish, limace des laminaires 8. Paraliparis bathybius (Collett, 1879) – Black Seasnail, limace noire 9. Paraliparis copei Goode and Bean, 1896 – Blacksnout Snailfish, limace à museau noir 10. Paraliparis garmani Burke, 1912 – Pouty Snailfish, limace pote 11. Rhodichthys regina Collett, 1879 – Threadfin Snailfish, limace à filaments

Family 49. Caristiidae – Manefishes, Caristes 1. Caristius fasciatus (Borodin, 1930) – Banded Manefish, cariste barré Family 50. Zoarcidae – Eelpouts, Lycodes 1. Gymnelus barsukovi Chernova, 1999 – Barsukov’s Pout, unernak de Barsukov 2. Gymnelus bilabrus Andriashev, 1937 – Twolip Pout, unernak à deux lèvres 3. Gymnelus knipowitschi Chernova, 1999 – Knipowitsch’s Pout, unernak de Knipowitsch 4. Gymnelus retrodorsalis Le Danois, 1913 – Aurora Pout, unernak aurore 5. Gymnelus viridis (Fabricius, 1780) – Fish Doctor, unernak caméléon 6. Lycenchelys kolthoffi Jensen, 1904 – Checkered Wolf Eel, lycode quadrillée 7. Lycenchelys muraena (Collett, 1878) – Moray Wolf Eel, lycode murène 8. Lycenchelys paxillus (Goode and Bean, 1879) – Common Wolf Eel, lycode commune 9. Lycenchelys sarsii (Collett, 1871) – Theologian Eelpout, lycode de Sars 10. Lycodes adolfi Nielsen and Fosså, 1993 – Adolf ’s Eelpout, lycode d’Adolf 11. Lycodes esmarkii Collett, 1875 – Greater Eelpout, grande lycode 12. Lycodes eudipleurostictus Jensen, 1902 – Doubleline Eelpout, lycode à deux lignes 13. Lycodes frigidus Collett, 1879 – Glacial Eelpout, lycode glaciale 14. Lycodes jugoricus Knipowitsch, 1906 – Shulupaoluk, lycode plume 15. Lycodes lavalaei Vladykov and Tremblay, 1936 – Laval Eelpout, lycode de Laval 16. Lycodes luetkenii Collett, 1880 – Pink Eelpout, lycode rose

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17. Lycodes marisalbi Knipowitsch, 1906 – White Sea Eelpout, lycode de la mer Blanche 18. Lycodes mcallisteri Møller, 2001 – McAllister’s Eelpout, lycode de McAllister 19. Lycodes mucosus Richardson, 1855 – Saddled Eelpout, lycode à selles 20. Lycodes paamiuti Møller, 2001 – Paamiut Eelpout, lycode de Paamiut 21. Lycodes pallidus Collett, 1879 – Pale Eelpout, lycode pâle 22. Lycodes polaris (Sabine, 1824) – Canadian Eelpout, lycode polaire 23. Lycodes reticulatus Reinhardt, 1835 – Arctic Eelpout, lycode arctique 24. Lycodes sagittarius McAllister, 1975 – Archer Eelpout, lycode à arc 25. Lycodes seminudus Reinhardt, 1837 – Longear Eelpout, lycode à oreilles 26. Lycodes squamiventer Jensen, 1904 – Scalebelly Eelpout, lycode ventre-écaillé 27. Lycodes terraenovae Collett, 1896 – Newfoundland Eelpout, lycode de Terre-Neuve 28. Lycodes vahlii Reinhardt, 1831 – Checker Eelpout, lycode à carreaux 29. Lycodonus mirabilis Goode and Bean, 1883 – Chevron Scutepout, lycaspine à chevrons 30. Melanostigma atlanticum Koefoed, 1952 – Atlantic Soft Pout, mollasse atlantique 31. Zoarces americanus (Bloch and Schneider, 1801) – Ocean Pout, loquette d’Amérique

Family 53. Anarhichadidae – Wolffishes, Poissons-loups 1. Anarhichas denticulatus Krøyer, 1845 – Northern Wolffish, loup à tête large 2. Anarhichas lupus Linnaeus, 1758 – Atlantic Wolffish, loup atlantique 3. Anarhichas minor Olafsen, 1772 – Spotted Wolffish, loup tacheté 4. Anarhichas orientalis Pallas, 1814 – Bering Wolffish, loup de Béring

Family 51. Stichaeidae – Pricklebacks, Stichées 1. Acantholumpenus mackayi (Gilbert, 1896) – Blackline Prickleback, terrassier à six lignes 2. Anisarchus medius (Reinhardt, 1837) – Stout Eelblenny, lompénie naine 3. Chirolophis ascanii (Walbaum, 1792) – Atlantic Warbonnet, toupet marbré 4. Eumesogrammus praecisus (Krøyer, 1837) – Fourline Snakeblenny, quatre-lignes atlantique 5. Leptoclinus maculatus (Fries, 1838) – Daubed Shanny, lompénie tachetée 6. Lumpenus fabricii Reinhardt, 1836 – Slender Eelblenny, lompénie de Fabricius 7. Lumpenus lampretaeformis (Walbaum, 1792) – Snakeblenny, lompénie-serpent 8. Stichaeus punctatus (Fabricius, 1780) – Arctic Shanny, stichée arctique 9. Ulvaria subbifurcata (Storer, 1839) – Radiated Shanny, ulvaire deux-lignes

Family 58. Pleuronectidae – Righteye Flounders, Plies 1. Glyptocephalus cynoglossus (Linnaeus, 1758) – Witch Flounder, plie grise 2. Hippoglossoides platessoides (Fabricius, 1780) – American Plaice, plie canadienne 3. Hippoglossoides robustus Gill and Townsend, 1897 – Bering Flounder, plie de Béring 4. Hippoglossus hippoglossus (Linnaeus, 1758) – Atlantic Halibut, flétan atlantique 5. Limanda proboscidea Gilbert, 1896 – Longhead Dab, limande carline 6. Platichthys stellatus (Pallas, 1788) – Starry Flounder, flet étoilé 7. Pleuronectes glacialis Pallas, 1776 – Arctic Flounder, plie arctique 8. Pleuronectes putnami (Gill, 1864) – Smooth Flounder, plie lisse 9. Reinhardtius hippoglossoides (Walbaum, 1792) – Greenland Halibut, flétan du Groenland

Family 54. Chiasmodontidae – Black Swallowers, Grands avaleurs 1. Chiasmodon harteli Melo, 2009 – Hartel’s Swallower, avaleur d’Hartel Family 55. Ammodytidae – Sand Lances, Lançons 1. Ammodytes dubius Reinhardt, 1837 – Northern Sand Lance, lançon du nord 2. Ammodytes hexapterus Pallas, 1814 – Pacific Sand Lance, lançon gourdeau Family 56. Trichiuridae – Cutlassfishes, Sabres de mer 1. Aphanopus carbo Lowe, 1839 – Black Scabbardfish, aphanope charbon Family 57. Stromateidae – Butterfishes, Stromatées 1. Peprilus triacanthus (Peck, 1804) – Butterfish, stromatée à fossettes

Family 52. Pholidae – Gunnels, Sigouines 1. Pholis fasciata (Bloch and Schneider, 1801) – Banded Gunnel, sigouine rubanée



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EXTRALIMITAL SPECIES Brian W. Coad

The record in R. Bell (1881) of a dead Haddock – Melanogrammus aeglefinus (Linnaeus, 1758) – floating in Hudson Strait has not been repeated with voucher specimens. A single record interpreted as Acipenser oxyrinchus Mitchill, 1815, from the George River in Ungava Bay by Scott and Scott (1988), has never been duplicated; the original record (Low, 1896) refers only to Accipenser sp. (sic), and other records in the same paragraph are from James Bay. Note that La Grande River of James Bay was formerly the Fort George River, and there is evident confusion over localities. The record of Pollachius virens (Linnaeus, 1758) “as far north as Hudson and Davis Straits” in Bigelow and Welsh (1925), and subsequently in Collette and Klein-MacPhee (2002), probably dates back to Sabine (1824), but we have no recent records despite extensive surveys. Envirocon Ltd. (1977) lists Delolepis gigantea Kittlitz, 1858 (the same as Cryptacanthodes giganteus, Family Cryptacanthodidae), cryptacanthodid larvae, and Liparis rutteri (Gilbert & Snyder, in Jordan & Evermann, 1898) from the Beaufort Sea (16.5 km north of Pullen Island), but these are probably misidentifications. The records of Microgadus tomcod (Walbaum, 1792) from Port Burwell (Killiniq) (Imperial Oil Limited, Aquitane Co. of Canada Ltd., & Canada-Cities Service Ltd., 1978a; MacLaren Atlantic Limited, 1978b) have not been confirmed by voucher specimens and could be misidentified. MacLaren Marex (1979a, 1979b) report Ceratoscopelus maderensis (Lowe, 1839) (Family Myctophidae) in Davis Strait as gut contents of Ivory Gulls, and Cairns (1982) reports Cryptacanthodes maculatus Storer, 1839 (Family Cryptacanthodidae) from Black Guillemot chick food near the Nuvuk Islands, Nunavut, but these need confirmation as there are no neighbouring records for these species, and the gut contents may be in poor condition. Records of Lycenchelys verrillii (Bean, 1877) from eastern Hudson Strait and Ungava Bay need verification

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(Parsons, 1982; Stewart, Dunbar, and Bernier, 1993). A paper by Siferd (2010) summarizing shrimp by-catches from 1979 to 2009 off southern Baffin Island records numerous species not found in other works on this area. The fish species were evidently identified by various workers aboard ship, incidental to the main work. While some of these fish species records may be correct, others are patently unlikely, and in the absence of voucher specimens none are included here. The following taxa have been recorded from waters near Arctic Canada but lack definitive Canadian records or voucher specimens. Key characters and descriptions for most can be found in Coad, with Waszczuk and Labignan (1995) and other references in the bibliography herein. The record of a Patagonian Toothfish (Dissostichus eleginoides Smitt, 1898), an Antarctic stray, is from Møller, Nielsen, and Fossen (2003). The waters of western Greenland harbour a number of species not yet reported from the Canadian side of the Davis Strait; this fauna has most recently been summarized in Nielsen, Bertelsen, and Nystrøm (1992), Jørgensen, Hvingel, Møller, and Treble (2005), and Møller et al. (2010; only species reported from their NW and SW area). The western Beaufort Sea of Alaska also contains some species not yet reported from Canada – this fauna has been documented by Mecklenburg et al. (2002) and Rand and Logerwell (2011) – and even more species occur in the Chukchi Sea (but are not listed here). Andriyashev and Chernova (1995) give an annotated list of species in circumpolar Arctic seas and adjacent waters. Note that the Zoarcidae are currently under study by several students of this family, and views as to species validity and distribution vary. Eastern Arctic (eastern Davis Strait and eastern Baffin Bay) species are marked by (D), western Arctic (western Beaufort Sea of Alaska) species by (B).

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Family 1. Myxinidae 1. Myxine jespersenae Møller, Feld, Poulsen, Thomsen, and Thormar, 2005 (D) Family 2. Petromyzontidae 1. Petromyzon marinus Linnaeus, 1758 (D)

Family 17. Alepocephalidae 1. Bathylaco nigricans Goode and Bean, 1896 (D) 2. Einara edentula (Alcock, 1892) (D) 3. Photostylus pycnopterus Beebe, 1933 (D) 4. Rouleina attrita (Vaillant, 1888) (D)

Family 3. Rhinochimaeridae 1. Harriotta haeckeli Karrer, 1972 (D)

Family 18. Salmonidae 1. Coregonus laurettae Bean, 1881 (B) 2. Coregonus pidschian (Gmelin, 1789) (B)

Family 4. Chimaeridae 1. Hydrolagus pallidus Hardy and Stehmann, 1990 (D)

Family 19. Gonostomatidae 1. Cyclothone braueri Jesperson and Tåning, 1926 (D)

Family 5. Cetorhinidae 1. Cetorhinus maximus (Gunnerus, 1765) (D)

Family 7. Scyliorhinidae 1. Apristurus laurussonii (Saemundsson, 1922) (D)

Family 20. Sternoptychidae 1. Argyropelecus aculeatus Valenciennes, 1850 (D) 2. Argyropelecus hemigymnus Cocco, 1829 (D) 3. Argyropelecus olfersi (Cuvier, 1829) (D) 4. Maurolicus muelleri (Gmelin, 1789) (D) 5. Polyipnus asteroides Schultz, 1938 (D) 6. Polyipnus polli Schultz, 1961 (D) 7. Sternoptyx pseudobscura Baird, 1971 (D)

Family 8. Squalidae 1. Squalus acanthias Linnaeus, 1758 (D)

Family 21. Phosichthyidae 1. Polymetme corythaeola (Alcock, 1898) (D)

Family 9. Etmopteridae 1. Etmopterus princeps Collett, 1904 (D)

Family 22. Stomiidae 1. Melanostomias bartonbeani Parr, 1927 (D) 2. Trigonolampa miriceps Regan and Trewavas, 1930 (D)

Family 6. Lamnidae 1. Lamna nasus (Bonnaterre, 1788) (D)

Family 10. Rajidae 1. Rajella bigelowi (Stehmann, 1978) (D)

Family 23. Synodontidae 1. Bathysaurus ferox Günther, 1878 (D)

Family 11. Halosauridae 1. Aldrovandia phalacra (Vaillant, 1888) (D)

Family 24. Alepisauridae 1. Alepisaurus brevirostris Gibbs, 1960 (D) 2. Alepisaurus ferox Lowe, 1833 (D)

Family 12. Anguillidae 1. Anguilla rostrata (Lesueur, 1817) (D) Family 13. Synaphobranchidae 1. Histiobranchus bathybius (Günther, 1877) (D)

Family 25. Myctophidae 1. Ceratoscopelus maderensis (Lowe, 1839) (D) 2. Lampadena speculigera Goode and Bean, 1896 (D)

Family 14. Nemichthyidae 1. Avocettina infans (Günther, 1878) (D)

Family 26. Lampridae 1. Lampris guttatus (Brünnich, 1788) (D)

Family 15. Microstomatidae 1. Melanolagus bericoides (Borodin, 1929) (D) 2. Nansenia sp. (D) 3. Nansenia groenlandica (Reinhardt, 1840) (D) 4. Nansenia oblita (Facciolà, 1887) (D)

Family 27. Trachipteridae 1. Trachipterus arcticus (Brünnich, 1788) (D) Family 28. Macrouridae 1. Coryphaenoides longifilis Günther, 1878 (D) 2. Nezumia aequalis (Günther, 1878) (D)

Family 16. Platytroctidae 1. Holtbyrnia macrops Maul, 1957 (D) 2. Sagamichthys schnakenbecki (Krefft, 1953) (D) 3. Searsia koefoedi Parr, 1937 (D)



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Family 29. Phycidae 1. Ciliata septentrionalis (Collett, 1875) (D) 2. Urophycis regia (Walbaum, 1792) (D) 3. Urophycis tenuis (Mitchill, 1814) (D)

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Family 30. Gadidae 1. Gadus chalcogrammus Pallas, 1814 (B) 2. Gadus macrocephalus Tilesius, 1810 (B) 3. Melanogrammus aegelfinus (Linnaeus, 1758) (D) 4. Merlangius merlangus (Linnaeus, 1758) (D) 5. Molva dypterygia (Pennant, 1784) (D) 6. Molva molva (Linnaeus, 1758) (D) 7. Pollachius virens (Linnaeus, 1758) (D) 8. Trisopterus esmarkii (Nilsson, 1855) (D)

Family 40. Cetomimidae 1. Gyrinomimus myersi Parr, 1934 (D) Family 41. Diretmidae 1. Diretmoides pauciradiatus (Woods, 1973) (D) Family 42. Oreosomatidae 1. Allocyttus verrucosus (Gilchrist, 1906) (D)

Family 31. Lophiidae 1. Lophius piscatorius Linnaeus, 1758 (D)

Family 43. Scorpaenidae 1. Helicolenus dactylopterus (Delaroche, 1809) (D) 2. Sebastes viviparus Krøyer, 1845 (D)

Family 32. Caulophrynidae 1. Caulophryne jordani Goode and Bean, 1896 (D)

Family 44. Syngnathidae 1. Entelurus aequoreus (Linnaeus, 1758) (D)

Family 33. Melanocetidae 1. Melanocetus johnsonii Günther, 1864 (D) 2. Melanocetus murrayi Günther, 1887 (D)

Family 45. Hexagrammidae 1. Hexagrammos stelleri Tilesius, 1810 (B)

Family 34. Oneirodidae 1. Chaenophryne draco Beebe, 1932 (D) 2. Danaphryne nigrifilis (Regan and Trewavas, 1932) (D) 3. Dolopichthys longicornis Parr, 1927 (D) 4. Lophodolos acanthognathus Regan, 1925 (D) 5. Oneirodes eschrichtii Lütken, 1871 (D) 6. Oneirodes macrosteus Pietsch, 1974 (D) 7. Phyllorhinichthys balushkini Pietsch, 2004 (D) 8. Phyllorhinichthys micractis Pietsch, 1969 (D)

Family 46. Cottidae 1. Artediellus gomojunovi Taranetz, 1933 (B) 2. Enophrys diceraus (Pallas, 1788) (B) 3. Megalocottus platycephalus (Pallas, 1814) (B) 4. Myoxocephalus jaok (Cuvier, 1829) (B) 5. Myoxocephalus verrucosus (Bean, 1881) (B). Or a synonym of Myoxocephalus scorpius (Linnaeus, 1758) Family 47. Hemitripteridae 1. Nautichthys pribilovius (Jordan and Gilbert, 1898) (B)

Family 35. Ceratiidae 1. Cryptopsaras couesii Gill, 1883 (D)

Family 48. Agonidae 1. Podothecus veternus Jordan and Starks, 1895 (B)

Family 36. Linophrynidae 1. Haplophryne mollis (Brauer, 1902) (D) 2. Linophryne algibarbata Waterman, 1939 (D) 3. Linophryne bicornis Parr, 1927 (D) 4. Linophryne coronata Parr, 1927 (D) 5. Linophryne lucifer Collett, 1886 (D)

Family 49. Cyclopteridae 1. Eumicrotremus andriashevi Perminov, 1936 (B)

Family 37. Melamphaidae 1. Melamphaes microps (Günther, 1878) (D) 2. Poromitra capito Goode and Bean, 1883 (D) 3. Poromitra crassiceps (Günther, 1878) (D) 4. Scopelogadus beani (Günther, 1887) (D) Family 38. Rondeletiidae 1. Rondeletia loricata Abe and Hotta, 1963 (D) Family 39. Barbourisiidae 1. Barbourisia rufa Parr, 1945 (D)

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Family 50. Liparidae 1. Careproctus cf. rastrinus Gilbert and Burke, 1912 (B) (from Rand & Logerwell, 2011) 2. Liparis bathyarcticus Parr, 1931 (B, D) 3. Liparis coheni Able, 1976 (D) 4. Liparis marmoratus Schmidt, 1950 (B) 5. Paraliparis hystrix Merrett, 1983 (D) 6. Psednos groenlandicus Chernova, 2001 (D) 7. Psednos harteli Chernova, 2001 (D) 8. Psednos melanocephalus Chernova and Stein, 2002 (D) 9. Psednos micruroides Chernova, 2001 (D) Family 51. Caristiidae 1. Paracaristius maderensis (Maul, 1949) (D) 2. Platyberyx opalescens Zugmayer, 1911 (D)

Checklists Of Sp ecies

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Family 52. Zoarcidae 1. Lycenchelys alba (Vaillant, 1888) (as L. labradorensis Geistdoerfer, Hureau, and Rannou, 1970) (D) 2. Lycodes palearis Gilbert, 1896 (B) 3. Lycodes raridens Taranetz and Andriashev, 1937 (B) 4. Lycodes turneri Bean, 1879 (B) Family 53. Stichaeidae 1. Lumpenella longirostris (Evermann and Goldsborough, 1907) (D) Family 54. Pholidae 1. Pholis gunnellus (Linnaeus, 1758) (D)

Family 56. Ammodytidae 1. Ammodytes marinus Raitt, 1934 (D) Family 57. Scombridae 1. Thunnus thynnus (Linnaeus, 1758) (D) Family 58. Pleuronectidae 1. Limanda aspera (Pallas, 1814) (B) 2. Pleuronectes platessa Linnaeus, 1758 (D) 3. Pleuronectes quadrituberculatus Pallas, 1814 (B) 4. Pseudopleuronectes americanus (Walbaum, 1792) (D)

Family 55. Nototheniidae 1. Dissostichus eleginoides Smitt, 1898 (D)



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Ex t r al i mi ta l Spe ci es

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BRACKISH WATER SPECIES Brian W. Coad

A number of species whose life is usually spent entirely in fresh waters may enter brackish waters and have been recorded in estuaries and seashore pools. They are rarely or never found in full sea water (35 ppt), and the records often do not cite salinity levels; they could be quite low in some instances. These species can be identified using the keys in Scott and Crossman (1973), but they are listed here as an advisory.

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Checklists Of Sp ecies

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Family 1. Petromyzontidae 1. Ichthyomyzon unicuspis Hubbs and Trautman, 1937

Family 7. Osmeridae 1. Hypomesus olidus (Pallas, 1814)

Family 2. Acipenseridae 1. Acipenser oxyrinchus Mitchill, 1815

Family 8. Salmonidae 1. Thymallus arcticus (Pallas, 1776)

Family 3. Hiodontidae 1. Hiodon alosoides (Rafinesque, 1819) 2. Hiodon tergisus Lesueur, 1818

Family 9. Percopsidae 1. Percopsis omiscomaycus (Walbaum, 1792) Family 10. Gasterosteidae 1. Culaea inconstans (Kirtland, 1840)

Family 4. Cyprinidae 1. Chrosomus eos Cope, 1862 2. Chrosomus neogaeus (Cope, 1867) 3. Couesius plumbeus (Agassiz, 1850) 4. Cyprinus carpio Linnaeus, 1758 5. Margariscus margarita (Cope, 1868) 6. Notropis atherinoides Rafinesque, 1818 7. Notropis hudsonius (Clinton, 1824) 8. Pimephales promelas Rafinesque, 1820 9. Platygobio gracilis (Richardson, 1836) 10. Rhinichthys atratulus (Hermann, 1804) 11. Rhinichthys cataractae (Valenciennes, 1842) 12. Semotilus corporalis (Mitchill, 1817)

Family 11. Cottidae 1. Cottus bairdii Girard, 1850 2. Cottus cognatus Richardson, 1836 3. Cottus ricei (Nelson, 1876) Family 12. Percidae 1. Etheostoma nigrum Rafinesque, 1820 2. Perca flavescens (Mitchill, 1814) 3. Percina caprodes (Rafinesque, 1818) 4. Percina shumardi (Girard, 1859) 5. Sander canadensis (Griffith and Smith, 1834) 6. Sander vitreus (Mitchill, 1818)

Family 5. Catostomidae 1. Catostomus catostomus (Forster, 1773) 2. Catostomus commersonii (Lacepède, 1803) 3. Moxostoma macrolepidotum (Lesueur, 1817)

Family 13. Sciaenidae 1. Aplodinotus grunniens Rafinesque, 1819

Family 6. Esocidae 1. Esox lucius Linnaeus, 1758



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B r a cki s h Wat e r Spe ci es

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Keys

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INTRODUCTION Brian W. Coad

The following keys are based on characters found in adult fishes. Most juveniles may also be identified here, but very young fish and larvae may not have these characters fully developed. Only species recorded from Arctic Canada are included in the keys. Specimens that do not key out and do not agree with the descriptions may be new to Arctic Canada; they may be identified using the works listed in the bibliography.

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Readers unfamiliar with technical terms or usage of common words as applied to fishes (e.g., nape, saddle) should consult the glossary for definitions.

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spines second dorsal fin

first dorsal fin

spiracle

nostril

heterocercal caudal fin

mouth

gill slits

pectoral fin

clasper anus

Shark external anatomy.

lateral line cheek

snout

nostril

pelvic axillary process

operculum

gill opening

anus pectoral fin

dorsal fin

pelvic fin (abdominal)

anal fin adipose fin

caudal peduncle

caudal fin

Bony fish external anatomy (soft-rayed fish).



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Int r o duc t i on

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spines

operculum

dorsal fins

cheek

soft rays caudal fin

nostrils

mouth

preoperculum

soft rays

gill opening pectoral fin spine

pelvic fin (thoracic)

soft rays spines vent

caudal peduncle

anal fin

Bony fish external anatomy (spiny-rayed fish).

teeth on head of vomer palatine teeth teeth on shaft of vomer maxillary teeth

basibranchial or hyoid teeth

lower jaw or mandible with teeth

Teeth in mouth.

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tongue with teeth

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FAMILIES Brian W. Coad

Families with only a single Canadian Arctic species will key out here, and that species is illustrated. In families with more than one species a selected species serves as an example for that family. Separate keys to families with two or more species follow this family key. The common name of a family follows its scientific name, and both are shown in bold and capitalized. Species names are in italics. Some other common names of recognizable groups of fishes are given but are neither in bold nor capitalized, because the name encompasses more than one family, for example, sharks. Distributional information is given for each species as it keys out. This is a broad summary of current knowledge, and some species may eventually be found to have a wider distribution. The distribution maps should be consulted, and major gaps noted, which may be real or artifacts of collecting. Many species are known only from Davis Strait, for example, which reduces the number of species likely to be keyed out in Beaufort Sea samples.



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The main areas of distribution are given from east to west and are summarized in the introduction to the “Family and Species Accounts” section. Fishes restricted to any one or two of these areas are so named in the keys; others are summarized as, for example, “Baffin Bay to Hudson Bay (which includes presence in the intermediate Davis Strait and Hudson Strait).” Distributions tend to be highly localized (as currently known), widespread (the whole Arctic), or mostly east or west of the Boothia Peninsula. Characters in brackets are additional identification features but are not unique to that family, that species, or subsequent families and species in the key.

F am i l i e s

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1. A single, median nostril; no paired fins; no jaws (Hagfishes and Lampreys) ➞ 2

Nostrils lateral and usually paired, rarely a single opening; 1–2 paired fins (the pectorals and pelvics); jaws present (sharks, Skates, chimaeras, and bony fishes) ➞ 3

2. One gill opening on each side of head; long barbels on snout; mouth not a sucking disc; nostril at the tip of the head; Davis and Hudson Straits = Myxinidae, Hagfishes (Myxine glutinosa)

Seven gill openings on each side; no snout barbels; mouth a sucking disc with teeth; nostril on top of head; Beaufort Sea = Petromyzontidae, Lampreys (Lethenteron camtschaticum)

Myxinidae (Myxine glutinosa)

Petromyzontidae (Lethenteron camtschaticum)

Underside of head

3. Five external gill openings; scales placoid; teeth in conveyorbelt rows ➞ 4

Ventral sucking disc

One external gill opening; scales cycloid, ctenoid, or absent; teeth not in conveyor-belt rows ➞ 7

Cycloid and ctenoid scales

Placoid scales

4. Pectoral fins attached to side of head in front of gill openings, forming a disc-like shape to body; gill openings ventral = Rajidae, Skates

Pectoral fins free and mostly behind gill openings; body form usually fusiform; gill openings mostly lateral; sharks ➞ 5

Ventral view, Rajidae, e.g., Amblyraja hyperborea

Sharks, e.g., Centroscyllium fabricii

Rajidae, e.g., Bathyraja spinicauda

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5. Anal fin present; Davis Strait = Scyliorhinidae, Cat Sharks (Apristurus profundorum)

Scyliorhinidae (Apristurus profundorum)

Anal fin absent ➞ 6

Anal fin absent, e.g., Somniosus microcephalus

6. Upper jaw teeth lanceolate, lower jaw teeth with oblique cusp; [dorsal fins often without a spine at origin, or spine small and often concealed by tissue] = Somniosidae, Sleeper Sharks

Teeth in both jaws with central cusp and lateral cusplets; [dorsal fins each with a stout spine at origin]; Davis Strait and Ungava Bay = Etmopteridae, Lantern Sharks (Centroscyllium fabricii)

Upper teeth (left) and lower teeth (right) of Etmopteridae (Centroscyllium fabricii)

Upper teeth (left) and lower teeth (right) of Somniosidae

Somniosidae, e.g., Somniosus pacificus

Etmopteridae (Centroscyllium fabricii)

7. Barbels present in front of mouth on underside of elongate snout; 5 rows of bony scutes along body; [upper lobe of caudal fin larger than lower lobe]; Hudson and James Bays = Acipenseridae, Sturgeons (Acipenser fulvescens)

Barbels present or absent but never in front of mouth on underside of elongate snout; no scutes in 5 rows ➞ 8

Acipenseridae (Acipenser fulvescens)

8. Illicium and esca (a fishing apparatus modified from dorsal fin) present; gill opening behind and below pectoral fin base (anglerfishes) ➞ 9

No fishing apparatus; gill opening in front (above or below) or over pectoral fin base ➞ 13

Fishing apparatus



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9. Head very large and greatly depressed; mouth very large and wide; skin tabs along anterior belly and lower jaw; Baffin Bay = Lophiidae, Goosefishes (Lophius americanus)

Not as across ➞ 10

Lophiidae (Lophius americanus)

10. Mouth vertical to very strongly oblique (or even backwardly oblique); 2–3 caruncles (fleshy knobs) in front of dorsal fin (2 in the only known Arctic Canadian species, minute in fish greater than 40 cm standard length); Davis Strait = Ceratiidae, Seadevils (Ceratias holboelli)

Mouth horizontal to oblique; no caruncles ➞ 11

Ceratiidae (Ceratias holboelli)

11. Fishing apparatus elongate (greater than 70% standard length), emerging from extreme snout tip; body elongate, laterally compressed; caudal peduncle long; Davis Strait = Gigantactinidae, Whipnoses (Gigantactis vanhoeffeni)

Fishing apparatus short (less than 60% standard length), not emerging from extreme snout tip but posteriorly on snout; body usually globose, elongate in some; caudal peduncle short ➞ 12

Gigantactinidae (Gigantactis vanhoeffeni)

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12. Illicium (fishing rod) not retractable; chin and snout covered with wart-like papillae; large and scattered bony plates on body bearing a median spine; Davis Strait = Himantolophidae, Footballfishes (Himantolophus groenlandicus)

Himantolophidae (Himantolophus groenlandicus)

13. Eyes tiny, placed far anterior near snout tip; mouth enormous ➞ 14

Illicium retractable; no chin papillae; body spines absent or numerous, close set, and minute; Davis Strait = Oneirodidae, Dreamers

Oneirodidae, e.g., Chaenophryne longiceps

Not as across ➞ 15

Tiny eyes near snout tip, and enormous mouth, e.g., Saccopharynx ampullaceus

14. Gill openings closer to anus than to snout tip; dorsal fin origin in front of pectoral fin; pectoral fins minute; Davis Strait = Eurypharyngidae, Gulpers (Eurypharynx pelecanoides)

Eurypharyngidae (Eurypharynx pelecanoides)



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Gill openings closer to snout tip than to anus; dorsal fin origin far behind pectoral fin; pectoral fins well developed; Davis Strait = Saccopharyngidae, Swallowers (Saccopharynx ampullaceus)

Saccopharyngidae (Saccopharynx ampullaceus)

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15. Operculum mostly a fold of skin; males with pelvic and head claspers; tooth plates in mouth (chimaeras) ➞ 16

Operculum mostly comprised of bones; no claspers; no tooth plates (bony fishes) ➞ 17

16. Snout short and rounded; claspers of male bifid or trifid; axis of caudal fin horizontal (diphycercal) with fin nearly symmetrical, and upper and lower lobes of equal size; Davis and Hudson Straits = Chimaeridae, Shortnose Chimaeras (Hydrolagus affinis)

Snout long, pointed, and fleshy; claspers of male a single rod; axis of caudal fin slightly raised (i.e., appears slightly heterocercal) with fin asymmetrical, and upper lobe narrower than lower lobe; Davis Strait = Rhinochimaeridae, Longnose Chimaeras

Chimaeridae (Hydrolagus affinis)

17. Eyes on same side of head; head asymmetrical (“twisted”); body a very flattened oval = Pleuronectidae, Righteye Flounders

Rhinochimaeridae, e.g., Rhinochimaera atlantica

Eyes on opposite sides of head; head bilaterally symmetrical; body not a very flattened oval ➞ 18

Pleuronectidae, e.g., Pleuronectes putnami

18. Stomach and mouth highly distensible (halves of the lower jaw can separate); some teeth depressible in front of mouth, and generally teeth overlap lips when mouth closed; [black overall; teeth in both jaws long and sharp]; Davis Strait = Chiasmodontidae, Black Swallowers (Chiasmodon harteli)

Not as across ➞ 19

Chiasmodontidae (Chiasmodon harteli)

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19. Pelvic fins modified into an adhesive disc ➞ 20

Pelvic fins not modified into an adhesive disc (or fins absent) ➞ 21

Adhesive disc

20. Skin with large tubercles = Cyclopteridae, Lumpfishes

Skin with small prickles or naked = Liparidae, Snailfishes (in part)

Large tubercles, e.g., in Cyclopterus lumpus

21. Skin loose with jelly-like tissue underneath; pelvic fins absent = Liparidae, Snailfishes (in part)

Skin without jelly-like tissue underneath; pelvic fins usually present ➞ 22

22. Small tube visible on flank over pectoral fin (connected to a sac under the cleithrum bone of the shoulder region); [photophores and tusks (enlarged anterior premaxillary teeth) often present] = Platytroctidae, Tubeshoulders

Not as across ➞ 23

Platytroctidae, e.g., Normichthys operosus



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23. Gill openings low on body, in form of slits close together, anterior to pectoral fin insertion or on the belly = Synaphobranchidae, Cutthroat Eels

Gill openings as slits, Synaphobranchus kaupii

Gill openings on the side of the head ➞ 24

Gill openings on belly, Simenchelys parasitica

Synaphobranchidae, e.g., Synaphobranchus kaupii

24. Upper and lower jaws diverging; Davis Strait = Nemichthyidae, Snipe Eels (Nemichthys scolopaceus)

Not as across ➞ 25

Nemichthyidae (Nemichthys scolopaceus)

25. Sac-like, toothed outgrowths of gut visible behind last gill arch under operculum; minute, forward-pointing spine in front of dorsal fin (felt by running finger towards dorsal fin, but may be concealed by skin); 17–25 large pores below anterior half of dorsal fin; Davis Strait = Stromateidae, Butterfishes (Peprilus triacanthus)

Not as across ➞ 26

Stromateidae (Peprilus triacanthus)

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26. Triangular teeth on roof of mouth in double row forming an obvious serrated ridge; Davis and Hudson Straits = Serrivomeridae, Sawpalates (Serrivomer beanii)

Not as across ➞ 27

Serrivomeridae (Serrivomer beanii)

27. Body ribbon-like and elongate, extremely thin, and flattened; Davis Strait = Trichiuridae, Cutlassfishes (Aphanopus carbo)

Not as across ➞ 28

Trichiuridae (Aphanopus carbo)

28. Dorsal fin originating on head; high, steep slope to head, below dorsal fin origin; dorsal, anal, and pelvic fins with scaly sheaths; Baffin Bay and Davis Strait = Caristiidae, Manefishes (Caristius fasciatus)

Not as across ➞ 29

Caristiidae (Caristius fasciatus)



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29. Large, diamond-shaped lateral-line scales and belly with a small midline ridge formed from 13–25 modified scales (indistinct in some); [head with large mucous cavities covered by skin and separated by bony crests]; Davis Strait = Trachichthyidae, Slimeheads (Hoplostethus atlanticus)

Not as across ➞ 30

Trachichthyidae (Hoplostethus atlanticus)

30. Head sculptured and ridged; surface of head bones and scales paper-thin and transparent; lateral line of 1–2 pored scales behind operculum; [1–3 weak spines in dorsal fin]; Davis Strait = Melamphaidae, Ridgeheads (Scopeloberyx robustus)

Not as across ➞ 31

Melamphaidae (Scopeloberyx robustus)

31. Dorsal fin composed solely of spines; anal fin with 13 or more spines and 110 or more soft rays; [no true caudal fin, but anal fin may regenerate around a broken-off tail tip] = Notacanthidae, Deep-sea Spiny Eels

Dorsal fin(s) with some soft rays, rarely dorsal fin absent; combination of high count of anal fin spines and soft rays absent; [caudal fin usually present] ➞ 32

Notacanthidae, e.g., Notacanthus chemnitzii

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32. One nostril on each side of head = Zoarcidae, Eelpouts

One pair of nostrils on each side of head ➞ 33

One nostril in Zoarcidae, e.g., Lycodes reticulatus

33. Body hatchet-shaped and very compressed; an unusual blade in front of the dorsal fin, formed from fused fin spines; light organs large, black and silvery, and vertically elongate (characters for Arctic Canadian species); Davis Strait = Sternoptychidae, Marine Hatchetfishes (Argyropelecus gigas)

Not as across ➞ 34

Sternoptychidae (Argyropelecus gigas)

34. Males with an intromittent organ; dorsal, caudal, and anal fins continuous, without separating notch; anterior nostril immediately above upper lip; Davis Strait = Bythitidae, Viviparous Brotulas (Bythites fuscus)

Not as across ➞ 35

Bythitidae (Bythites fuscus)

35. Isolated spines, without membrane connection, preceding soft dorsal fin = Gasterosteidae, Sticklebacks

No isolated spines ➞ 36

Isolated spines, e.g., Gasterosteus aculeatus

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36. Single infra-orbital bone; no gill rakers in adults; chin barbel as long or longer than head (if absent, mouth has no floor or hexagonal scales present) = Stomiidae, Dragonfishes

Not as across ➞ 37

Stomiidae, e.g., Borostomias antarcticus

37. Scales on a short pedicel embedded in the skin, goblet-shaped and ridged; lateral line in a groove partly covered by scales; large, skin-covered mucous cavities on head; Davis Strait = Anoplogastridae, Ogrefishes (Anoplogaster cornuta)

Not as across ➞ 38

Pedicel scales

Anoplogastridae (Anoplogaster cornuta)

38. Body encased in bony plates = Agonidae, Poachers

Body not encased in bony plates ➞ 39

Bony plates in Agonidae

39. Pelvic fins absent ➞ 40

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Pelvic fins present ➞ 41

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40. Snout rounded and blunt; upper and lower incisor teeth conical and obvious; skin not in oblique folds = Anarhichadidae, Wolffishes

Anarhichadidae, e.g., Anarhichas orientalis

Snout pointed and sharp; no obvious incisors; skin in folds = Ammodytidae, Sand Lances

Ammodytidae, e.g., Ammodytes dubius

41. Photophores or luminescent organs present ➞ 42

Photophores or luminescent organs absent ➞ 43

42. Dorsal fin near caudal fin over anal fin; body elongate; eye small = Gonostomatidae, Bristlemouths

Dorsal fin not near caudal fin but at mid-body; body not elongate; eye large = Myctophidae, Lanternfishes

Gonostomatidae, e.g., Cyclothone microdon

43. Adipose fin present; pelvic fins abdominal ➞ 44

Myctophidae, e.g., Benthosema glaciale

No adipose fin; pelvic fins abdominal, thoracic, or jugular ➞ 49

Positions of pelvic fins



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44. All fins reduced in size; lower jaw and palatine teeth alternately fixed and depressible = Paralepididae, Barracudinas

All or most fins of normal size; teeth not as across ➞ 45

Paralepididae, e.g., Arctozenus risso

45. Pectoral fins very long on elongate body, reaching dorsal fin origin; dorsal fin origin behind pelvic fins; Davis Strait = Notosudidae, Waryfishes (Scopelosaurus lepidus)

Not as across ➞ 46

Notosudidae (Scopelosaurus lepidus)

46. Eye very large, almost half head length; gill opening narrow but not a slit, extending less than half way up side of body; gill membranes joined or united; Baffin Bay to Hudson Strait = Microstomatidae, Pencilsmelts (Bathylagus euryops)

Eye much less than half head length; gill opening wide; gill membranes free or separate ➞ 47

Microstomatidae (Bathylagus euryops)

47. Dorsal fin insertion (where rear fin base touches body) at or in front of pelvic fin level; pectoral fin tip below dorsal fin origin (both equal a dorsal fin forward on back); Davis Strait = Argentinidae, Argentines (Argentina silus)

Dorsal fin insertion over or behind pelvic fins ➞ 48

Argentinidae (Argentina silus)

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48. Pelvic axillary process evident; tail skeleton turned up at tail base = Salmonidae, Trouts and Salmons

Pelvic axillary process absent; tail skeleton not obviously turned up = Osmeridae, Smelts

Pelvic axillary process

Salmonidae, e.g., Salmo salar

Osmeridae, e.g., Osmerus dentex

49. Pelvic fins abdominal (mid-body) ➞ 50

Pelvic fins thoracic or jugular (far forward) ➞ 51

50. Dorsal fin short, at mid-point of back; pelvic axillary process present = Clupeidae, Herrings

Dorsal fin long, nearer caudal fin than mid-point of back; pelvic axillary process absent = Alepocephalidae, Slickheads

Clupeidae, e.g., Clupea pallasii

Alepocephalidae, e.g., Alepocephalus agassizii

51. No spines on head; fins all soft and flexible, not spiny, or, if stiffened, branched ➞ 52



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Spines or spine-like rays in some or all of dorsal, anal, and pelvic fins; often spines on head ➞ 55

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52. No caudal fin; body tapers to a point; developed snout usually projects beyond mouth = Macrouridae, Grenadiers

Caudal fin present; body not tapering to a point; snout not projecting markedly beyond mouth ➞ 53

Macrouridae, e.g., Coryphaenoides carapinus

53. Two dorsal fins, the first short and the second long at the base, and deeply indented in some; one anal fin, moderately to deeply indented = Moridae, Codlings

One to three unindented dorsal fins; one to two unindented anal fins ➞ 54

Moridae, e.g., Lepidion eques

54. No snout barbels and no very elongate pelvic rays; [dorsal fins 1–3; anal fins 1–2 (Brosme has 1 dorsal fin and 1 anal fin, Lota has 2 dorsal fins and 1 anal fin, and rest have 3 dorsal fins and 2 anal fins)] = Gadidae, Cods

Gadidae, e.g., Gadus morhua

55. Anal fin-rays not spiny ➞ 56

106

Snout barbels or very elongate pelvic rays present; [dorsal fins 2–3 (in those with 3 the first is composed of a thickened ray and the second of short rays in a fleshy ridge within a groove); anal fin 1] = Phycidae, Phycid Hakes

Phycidae, e.g., Phycis chesteri

Anal fin with 1–5 spines, either at front of fin or anterior to it ➞ 57

K ey s

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56. Dorsal fins separated into obvious spiny and soft-rayed sections; skin not loose = Cottidae, Sculpins

Cottidae, e.g., Myoxocephalus octodecemspinosus

57. Pelvic fin normal sized, composed of 1 spine and 5 soft rays; body not elongate (body depth enters length about 3 times) = Scorpaenidae, Scorpionfishes

Dorsal fin continuous, spiny part small, low, and partially obscured by skin; skin loose in some = Psychrolutidae, Fathead Sculpins

Psychrolutidae, e.g., Cottunculus thomsonii

Pelvic fin small, composed of 1 spine and 1–4 soft rays; body elongate (body depth enters length about 6 times) ➞ 58

Scorpaenidae, e.g., Sebastes mentella

58. Distance from snout to anal fin origin more than distance from anal fin origin to caudal fin base (anal fin relatively short); pelvic fins composed of 1 spine and 1 ray; whole Arctic = Pholidae, Gunnels (Pholis fasciata)

Pholidae (Pholis fasciata)

Stichaeidae, e.g., Stichaeus punctatus



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Distance from snout to anal fin origin equal to or less than distance from anal fin origin to caudal fin base (anal fin relatively long); pelvic fins composed of 1 spine and up to 4 rays = Stichaeidae, Pricklebacks

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SPECIES IN INDIVIDUAL FAMILIES Brian W. Coad Zoarcidae by Peter Rask Møller gadiforms by Brian W. Coad and Claude B. Renaud

Keys to families having more than one species are given in alphabetical order.

Agonidae, Poachers (Family 45) 1. Two dorsal fins; 5 pairs of barbels around mouth; supralateral plate row present; whole Arctic = Leptagonus decagonus

One dorsal fin; 1 pair or no barbels on end of maxilla; supralateral plate row absent ➞ 2

Leptagonus decagonus

2. Dorsal row plates 45–53; pectoral fin-rays 9–11; no barbel on end of maxilla; Davis Strait to Hudson Strait = Aspidophoroides monopterygius

Aspidophoroides monopterygius

Dorsal row plates 33–40; pectoral fin-rays 12–16; a barbel on end of maxilla; whole Arctic = Aspidophoroides olrikii

Aspidophoroides olrikii

Alepocephalidae, Slickheads (Family 20) 1. Scales absent (or only present in lateral line) ➞ 2

108

Scales present (easily lost, but scale pockets evident) ➞ 3

K ey s

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2. Lateral line in a tube supported by ring-like scales; light organs mostly on ventral body; anal fin-rays 20–22; Davis Strait = Rouleina maderensis

Rouleina maderensis

Lateral line not in a tube; small, rounded light organs scattered on skin of head and body (best seen under a microscope); anal fin-rays 26–30; Davis Strait to Hudson Strait = Xenodermichthys copei

Xenodermichthys copei

3. Teeth absent from maxilla, and vomer bones in mouth ➞ 4

Teeth present on maxilla, and/or vomer bones in mouth ➞ 5

4. Anal fin-rays 15–18; dorsal fin-rays 15–18; lateral-line scales 80– 95; Baffin Bay and Davis Strait = Alepocephalus agassizii

Anal fin-rays 20–25; dorsal fin-rays 18–23; lateral-line scales 62–70; Davis Strait = Alepocephalus bairdii

Alepocephalus agassizii

Alepocephalus bairdii

5. Lower jaw symphysis (central tip) with a ventrally directed and prominent knob; anal fin-rays 13–16; Davis Strait = Bajacalifornia megalops

Bajacalifornia megalops

Lower jaw without prominent knob (weak projection may be present); anal fin-rays 9–11; Davis Strait = Bathytroctes sp.

Bathytroctes sp.

Ammodytidae, Sand Lances (Family 55) 1. Dorsal fin-rays 55–69 (usually 61–67); anal fin-rays 26–36 (usually 31–34); vertebrae 64–78 (usually 71–76), population mean high at 71–74 (dissection or X-rays required); skin folds 124–147 (usually 128–138); body slender; usually offshore; whole Arctic = Ammodytes dubius

Ammodytes dubius

Ammodytes hexapterus



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Dorsal fin-rays 51–63 (usually 55–60); anal fin-rays 23–34 (usually 26–31); vertebrae 60–75 (usually 63–70), population mean low at 67–68; skin folds 106–126; body deep; found inshore; whole Arctic = Ammodytes hexapterus

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Anarhichadidae, Wolffishes (Family 53) Note: The teeth should not be examined in live specimens(!). If the fish has been preserved with its strong jaw muscles locked in the closed position, some force will be needed to open the jaws.

1. Teeth on vomer bone in roof of mouth extending back beyond teeth on palatine bones at the sides ➞ 2

Teeth on vomer bone in roof of mouth not extending back beyond teeth on palatine bones at the sides ➞ 3

Vomer teeth (lower central) extending beyond palatine teeth (at sides) or not (left and right, respectively)

2. Flank and dorsal fin with 9–14 bars; dorsal fin spines 69–79; anal fin-rays 42–48; caudal fin, more or less truncate; Davis Strait to Hudson Strait = Anarhichas lupus

Anarhichas lupus

3. Not distinctly spotted; individual palatine teeth longer than vomerine teeth; upper lip thicker than lower and covered with papillae; Baffin Bay to Hudson Strait and sporadically in western Arctic = Anarhichas denticulatus

Anarhichas denticulatus

110

No bars on flank and dorsal fin; dorsal fin spines 80–88; anal fin-rays 50–55; caudal fin rounded; Bathurst Inlet and Coronation Gulf = Anarhichas orientalis

Anarhichas orientalis

Head, upper flank, and dorsal and caudal fins with distinct small to large black-brown spots; individual palatine teeth about equal in length or longer than vomerine teeth; upper lip not thicker than lower, and covered with irregular folds; Baffin Bay to Hudson Strait = Anarhichas minor

Anarhichas minor

K ey s

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Clupeidae, Herrings (Family 16)

1. Vertebrae 51–60 (mostly 55–57), (dissection or X-rays required); keeled scales 11–17, between the pelvic and anal fins; keeled scales more or less developed anterior and posterior to the pelvic fins; vomerine teeth well developed; eastern Arctic, Davis Strait to James Bay = Clupea harengus

Clupea harengus

Vertebrae 49–57 (mostly 52–55); keeled scales 10–14, between the pelvic and anal fins; keeled scales more or less developed only posterior to the pelvic fins; vomerine teeth less well developed; western Arctic, Queen Maud Gulf to Beaufort Sea = Clupea pallasii

Clupea pallasii

Cottidae, Sculpins (Family 44)

1. Palatine teeth present on roof of mouth (see mouth figure above) ➞ 2

2. Upper preopercular spine bifurcate or trifurcate, not strongly hooked upwards; row of spiny plates above lateral line ➞ 3

Palatine teeth absent ➞ 6 (note: Artediellus uncinatus may have no palatine teeth but has the upper preopercular spine pointed – not bifurcated or broadened at the tip – and hooked upwards)

Upper preopercular spine simple in form, pointed, and strongly hooked upwards (see figure under Cottidae, couplet 4); no bony and spiny plates above lateral line ➞ 4

Upper preopercular spine bifurcate or trifurcate, not strongly hooked upwards



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3. Lateral-line scales ending on or before anterior part of caudal peduncle (rarely running to posterior edge of hypural plate, the end of the vertebral column); caudal peduncle scales usually present above and below midline of caudal peduncle; pectoral axillary scales 9–30; row of small scales often present between dorsal scale row and lateral line; row of scales often present above anal fin; male urogenital papilla cylindrical, tip elongate and tapering; tip longer than depth of caudal peduncle; whole Arctic = Icelus bicornis

Lateral-line scales ending on posterior edge of hypural plate; caudal peduncle scales absent or, if present, only above the lateral line; pectoral axillary scales 1–14; no scale rows between dorsal scale row and lateral line; no scale row above anal fin; male urogenital papilla spatulate, with a short curved or hooklike tip; tip shorter than caudal peduncle depth; whole Arctic = Icelus spatula

Urogenital papilla tip is elongate and longer than caudal peduncle depth

Icelus bicornis

4. Cirri and conical tubercles present on nape and anterior back; Gulf of Boothia, Arctic islands, Dease Strait to Beaufort Sea = Artediellus scaber

Icelus spatula

Cirri and conical tubercles absent from nape and anterior back ➞5

Cirri, conical tubercles, and simple, hooked preopercular spine

Artediellus scaber

112

K ey s

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5. Caudal fin-rays 22–28 (includes smallest rays at caudal peduncle and may require dissection to see); two small pores between eyes anteriorly; vomerine teeth 6–24; palatine teeth 4–39 (on roof of mouth); parietal cirri absent; Baffin Bay to Hudson Strait = Artediellus atlanticus

Artediellus atlanticus

Caudal fin-rays 19–23; no pores between eyes anteriorly; vomerine teeth 0–12; palatine teeth 0–15; parietal cirri usually present (but may be abraded); Baffin Bay to Hudson Strait = Artediellus uncinatus

Artediellus uncinatus

6. Upper preopercular spine with spinules (smaller but evident branches); vomer without teeth; whole Arctic = Gymnocanthus tricuspis

Upper preopercular spine without spinules; vomer with teeth ➞7

Gymnocanthus tricuspis

7. Skin below lateral line not in folds; anal fin-rays 17 or less ➞ 8

Skin below lateral line in oblique folds with serrated edges; anal fin-rays 18 or more; lateral-line plates with backward pointing spines ➞ 12

Oblique folds with serrated edges



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8. Four preopercular spines, three together and exposed with the fourth more ventral and covered by skin; usually large roughtopped spines above eye and on occiput in adults; lateral line usually incomplete; mandibular pores difficult to see; whole Arctic = Myoxocephalus quadricornis

Three, rarely four, preopercular spines; no spines above eye and on occiput, but, if present, not large and not rough topped; lateral line usually complete; mandibular pores obviously present ➞ 9

Four preopercular spines and rough-topped spines above eye and on occiput

Myoxocephalus quadricornis

9. Uppermost preopercular spine long, about four times longer than the spine below it; Hudson Strait, Ungava Bay, and eastern Hudson Bay = Myoxocephalus octodecemspinosus

Uppermost preopercular spine shorter, only about twice as long as the spine below it ➞ 10

Myoxocephalus octodecemspinosus

10. Spiny dorsal fin origin anterior to level of posterior tip of operculum, operculum tip usually under third or fourth spine; no pore behind last gill arch; [broad and dark saddle below first 6 dorsal fin spines; 2 smaller saddles below soft dorsal fin; papillae on top of head weak to absent; head spines prominent; lateral line without plate-like scales below]; Hudson Strait and northern Hudson Bay = Myoxocephalus aenaeus

Spiny dorsal fin origin not markedly anterior to posterior tip of operculum; small pore behind last gill arch under operculum ➞ 11

Myoxocephalus aenaeus 114

K ey s

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11. Pectoral fin-rays 14–18, usually 14–16; total count of dorsal, anal, and pectoral fin-rays 35–38; frontal and parietal spines not well developed and lack accessory spines at base; papillae on top of head large and closely spaced; lateral line without plate-like scales below; whole Arctic = Myoxocephalus scorpioides

Myoxocephalus scorpioides

Pectoral fin-rays 14–19, usually 16–18; total count of dorsal, anal, and pectoral fin-rays 39–43; frontal and parietal spines usually have stout accessory spines at base; papillae on top of head small and scattered; row of spiny, plate-like scales above and below lateral line; whole Arctic = Myoxocephalus scorpius

Myoxocephalus scorpius

12. Lower jaw extends beyond upper jaw; breast, sides of abdomen, and peritoneum (dissection required) with dense melanophores; body dark, saddles beneath dorsal fins and on caudal peduncle absent or faint; pectoral fin-rays 20–23; second (middle) soft ray of pelvic fin longest; orbit diameter large, exceeding snout length and equal to or longer than postorbital distance; dorsal row of scutes reduced; whole Arctic = Triglops nybelini

Orbit diameter large

Jaws equal, or upper extends slightly beyond lower; breast, sides of abdomen unpigmented, peritoneum unpigmented or with widely spaced melanophores; body light, 4–5 saddles (1 beneath spiny dorsal fin, 2–3 beneath soft dorsal fin, and 1 on caudal peduncle); pectoral fin-rays 16–21, usually 16–19; third soft pelvic fin-ray longer than second ray (see figure); orbit diameter small, equals snout length, and less than postorbital distance; dorsal row of scutes well developed ➞ 13

Orbit diameter small

Triglops nybelini



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13. Caudal fin with 3–6 narrow, dark bars; second dorsal fin-rays 18–26, usually 19–24; lateral-line plates 44–49; second soft pelvic fin-ray longest in males, third soft pelvic fin-ray longest in females; males with black spot at rear of spiny dorsal fin; Baffin Bay to James Bay = Triglops murrayi

Triglops murrayi

Caudal fin without bars (rarely, a single faint bar, and upper lobe may have black tip); second dorsal fin-rays 20–28, usually 23–26; lateral-line plates 47–51; third soft pelvic fin-ray longest in males; males lacking black spot at rear of spiny dorsal fin; whole Arctic = Triglops pingelii

Triglops pingelii

Cyclopteridae, Lumpfishes (Family 47)

1. Gill opening large, extending below upper edge of pectoral fin base; tubercles in 3 clearly separated rows along whole flank; first dorsal fin covered by a humped crest of skin in adults; Davis Strait to James Bay = Cyclopterus lumpus

Gill opening large

Gill opening small, not extending below upper edge of pectoral fin base; larger tubercles, not in 3 clear rows along whole flank; first dorsal fin not a humped crest but sometimes skin covered ➞2

Gill opening small

Cyclopterus lumpus

2. Few blunt, large tubercles restricted to the anterior half of the body, others on rear half smaller or absent; supplemental pores usually absent ➞ 3

116

Many large, pointed tubercles, each with many spines on head and body; supplemental pores present above lateral line ➞ 4

K ey s

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3. First dorsal fin rounded with rays unequal in length; postorbital tubercle row present, extending onto the body; Frobisher and Ungava Bays, Prince Leopold Island = Cyclopteropsis jordani

Cyclopteropsis jordani

First dorsal fin rectangular with rays equal in length; postorbital tubercle row absent; Baffin Bay to Hudson Strait = Cyclopteropsis mcalpini

Cyclopteropsis mcalpini

4. First dorsal fin-rays covered with skin and tubercles, not visible; pectoral fin base without tubercles; whole Arctic = Eumicrotremus derjugini

First dorsal fin-rays clearly visible even when tubercles present; pectoral fin base with 2–13 tubercles in females but none in some males; whole Arctic = Eumicrotremus spinosus

Eumicrotremus spinosus

Eumicrotremus derjugini

Gadidae, Cods (Family 32)

Brian W. Coad and Claude B. Renaud

1. One dorsal fin and one anal fin, both joined, or nearly joined, to caudal fin, and each separated from it by a notch; Davis Strait = Brosme brosme

Two or three dorsal fins and one or two anal fins, dorsal and anal fins clearly distinct from caudal fin ➞ 2

Brosme brosme



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2. Two dorsal fins and one anal fin; Hudson and James Bays, Coronation Gulf to Beaufort Sea = Lota lota

Three dorsal fins and two anal fins ➞ 3

Lota lota

3. First anal fin much longer than second anal fin, lying under first and second dorsal fins; Davis Strait = Micromesistius poutassou

First anal fin short, about equal in size to second anal fin, lying under second dorsal fin ➞ 4

Micromesistius poutassou

4. Palatine bones in roof of mouth with teeth; [lateral line interrupted along its whole length]; whole Arctic = Arctogadus glacialis

Palatine teeth absent ➞ 5

Arctogadus glacialis

5. Gill rakers 35–47; lateral line very wavy; cycloid scales not imbricate, but scattered; whole Arctic = Boreogadus saida

Gill rakers 28 or less; lateral line not wavy (but arched over pectoral fin); cycloid scales imbricate ➞ 6

Boreogadus saida

118

K ey s

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6. Sensory pit organs present on head canals; last half of the lateral line interrupted; Queen Maud Gulf to Beaufort Sea = Eleginus gracilis

Sensory pores present on head canals; last third or less of the lateral line interrupted ➞ 7

Eleginus gracilis

7. Body with many distinct greenish, brownish, or reddish spots; lateral line white; barbel length less than eye diameter; peritoneum silvery grey with black speckling (dissection required); southern Baffin Bay to Hudson Strait = Gadus morhua

Gadus morhua

Body with dark marbling on a lighter background; lateral line blotched; barbel length equals or exceeds eye diameter; peritoneum black; whole Arctic = Gadus ogac

Gadus ogac

Gasterosteidae, Sticklebacks (Family 42)

1. Short or long dorsal fin spines, 3, in median line; vertically elongate bony scutes covering much of flank; eastern Arctic, Arctic islands (one record), possibly western Arctic = Gasterosteus aculeatus

Pungitius pungitius

Gasterosteus aculeatus



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Short dorsal fin spines 6–13 (usually 9), alternately sloping left and right; small scutes present but not covering whole flank; whole Arctic = Pungitius pungitius

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Gonostomatidae, Bristlemouths (Family 23) 1. Gill rakers 19–23; anal fin-rays 16–20; no white mass in fold of skin at rear of upper jaw; Baffin Bay and Davis Strait = Cyclothone microdon

Cyclothone microdon

Gill rakers 24–28; anal fin-rays 21–26; a white mass in fold of skin at rear of upper jaw; Davis Strait = Sigmops bathyphilus

Sigmops bathyphilus

Liparidae, Snailfishes (Family 48)

1. Adhesive disc present on ventral surface (see figures) ➞ 2

2. Adhesive disc about same diameter as eye; anal fin-rays 41–58, commonly 45–52; single nostril (note: an adjacent pore is not a nostril opening) ➞ 3

Adhesive disc small

3. Lower pectoral fin lobe much longer than head; body deep (equal to or greater than head length); [disc rounded; anus close behind disc; mouth usually horizontal, rarely oblique]; Hudson Strait = Careproctus longipinnis*

Adhesive disc absent ➞ 8

Adhesive disc several times larger than eye diameter in adults (note: some young Liparis fabricii in particular can have a small disc); anal fin-rays 25–42, commonly 25–40; pair of nostrils on each side of head ➞ 5

Adhesive disc large

Lower pectoral fin lobe about equal to or shorter than head; body slender (usually less than head length) ➞ 4

Careproctus longipinnis 120

K ey s

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4. Total pectoral fin-rays 21–26, 11–14 in upper lobe; colour light to dark brown; peritoneum black (dissection required); Davis Strait = Careproctus kidoi

Careproctus kidoi

Total pectoral fin-rays 25–34, 17–21 in upper lobe; colour white, pinkish, or red; peritoneum pale; [disc oval, pear-shaped, or triangular; disc-to-anus length about equal to disc length; mouth oblique]; whole Arctic = Careproctus reinhardti*

Careproctus reinhardti

5. Distinct notch in the anterior portion of dorsal fin; anal finrays 25–29; pectoral fin-rays 25–31; Ungava Bay = Liparis atlanticus

No notch in dorsal fin; anal fin-rays 31–42; pectoral fin-rays 32–45 ➞ 6

Liparis atlanticus

6. Anal fin-rays 35–42, usually 37–40; peritoneum dark; posterior (jaw corner) teeth simple in adults, only anterior teeth trilobed; whole Arctic = Liparis fabricii

Anal fin-rays 31–38, usually 31–36; peritoneum pale, sometimes with a few pigment spots; posterior (jaw corner) teeth trilobed in adults ➞ 7

Liparis fabricii

* Some authors synonymize these two species.



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7. Usually 8–16 pectoral fin-rays opposite gill opening in adults; total pectoral fin-rays 35–45, usually 38–42; posterior teeth with central lobe much longer than lateral lobes; whole Arctic = Liparis gibbus

Usually 0–7 pectoral fin-rays opposite gill opening; total pectoral fin-rays 31–40, usually 32–37; posterior teeth with central lobe about equal in length to lateral lobes; often with distinctive bar(s) on the caudal fin; whole Arctic = Liparis tunicatus

0–7 fin-rays opposite gill opening

Liparis gibbus

8. Lower 3 pectoral fin-rays joined as a filament longer than head and separate from rest of pectoral fin; gill opening large, extending from above pectoral fin base to lower pectoral base; Baffin Bay, Davis Strait, and Beaufort Sea = Rhodichthys regina

Liparis tunicatus

No filament, and several short rays between pectoral fin lobes; gill opening small, at most reaching down to upper pectoral fin base ➞ 9

Rhodichthys regina

9. Body black (note: skin may be lost in damaged specimens, leaving a pale body, but usually fragments of black skin are present); upper lobe pectoral fin-rays about 18–20; Baffin Bay and Davis Strait = Paraliparis bathybius

Body pale; upper lobe pectoral fin-rays 13–17 ➞ 10

Paraliparis bathybius

122

K ey s

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10. Teeth in single row; dorsal fin-rays 59–68; anal fin-rays 53–60; lower snout and mouth brown to black, strongly contrasting with surrounding tissues; Davis and Hudson Straits, Ungava Bay = Paraliparis copei

Paraliparis copei

Teeth in several rows; dorsal fin-rays 54–60; anal fin-rays 49–53: snout and mouth light, similar in colour to surrounding tissues; Davis Strait to Hudson Strait = Paraliparis garmani

Paraliparis garmani

Macrouridae, Grenadiers (Family 29) Brian W. Coad and Claude B. Renaud

1. Second dorsal fin-rays equal to or longer than anal fin-rays; gill rakers elongate; outer gill slit without a fold over upper and lower parts ➞ 2

Second dorsal fin-rays shorter than anal fin-rays; gill rakers are short tubercles or absent; outer gill slit with a fold over upper and lower parts, connecting first gill arch to operculum and narrowing gill slit ➞ 3

Gill rakers flattened and elongate (left) or short tubercles (right)

2. Mouth wide and terminal; snout rounded; scales without spinules; post-temporal fossa absent; no enlarged scutes along dorsal and anal fins; very elongate rays in first dorsal, pectoral, and pelvic fins; Davis Strait = Gadomus longifilis

Gadomus longifilis

Trachyrincus murrayi



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Mouth small and inferior; snout long and pointed; scales with spinules; post-temporal fossa present; row of enlarged scutes along dorsal and anal fins; Davis Strait = Trachyrincus murrayi

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3. Seven branchiostegal rays; anus well anterior to anal fin; Baffin Bay and Davis Strait = Nezumia bairdii

Six branchiostegal rays; anus at anal fin origin ➞ 4

Branchiostegal rays, and anus anterior to anal fin

Nezumia bairdii

4. Origin of second dorsal fin in front of anal fin origin level; no gill rakers on outer side of first gill arch; Baffin Bay to Hudson Strait = Macrourus berglax

Origin of second dorsal fin behind anal fin origin level; gill rakers present on outer side of first gill arch (few and small) ➞ 5

Macrourus berglax

5. Upper jaw not extending back to posterior third of orbit; Davis Strait = Coryphaenoides guentheri

Upper jaw extending back to posterior third of orbit or more ➞ 6

Coryphaenoides guentheri

6. Total inner gill rakers on first arch 17–20; snout rounded and tipped with large blunt scute; northern Baffin Island, Davis and Hudson Straits = Coryphaenoides rupestris

Total inner gill rakers on first arch 7–16; snout blunt or pointed, but without terminal scute ➞ 7

Coryphaenoides rupestris 124

K ey s

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7. Mandibular teeth in 2 or more rows; Davis Strait = Coryphaenoides carapinus

Mandibular teeth in 1 distinct row ➞ 8



Coryphaenoides carapinus

8. Pelvic fin-rays 10–12; premaxillary teeth in 1–2 rows, 1 in large adults; interorbital width 21%–27% of head length; Davis Strait = Coryphaenoides armatus

Coryphaenoides armatus

Pelvic fin-rays 8–9; premaxillary teeth in 3 or more rows; interorbital width 26%–41% of head length; Davis Strait = Coryphaenoides brevibarbis

Coryphaenoides brevibarbis

Moridae, Codlings (Family 30)

Brian W. Coad and Claude B. Renaud

1. Snout with shelf-like projections on each side; upper jaw projecting beyond lower jaw; Baffin Bay and Davis Strait = Antimora rostrata

Snout without shelf-like projections on each side; upper jaw not markedly projecting beyond lower jaw ➞ 2

Snout with shelf-like projections, upper jaw projecting

Antimora rostrata



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2. No chin barbel; second dorsal fin-ray not elongated; vomer without teeth; Davis Strait = Halargyreus johnsonii

Halargyreus johnsonii

Chin barbel present; second dorsal fin-ray very elongated (first ray small and hidden under skin); vomer with a few teeth on its head; Davis Strait = Lepidion eques

Lepidion eques

Myctophidae, Lanternfishes (Family 28)

Simplified distribution of photophores (see also family account)



1. AO row continuous; PLO below upper base of pectoral fin; eyes semi-telescopic (directed dorsally); Davis Strait = Protomyctophum arcticum

AO row subdivided into an anterior (AOa) and a posterior group (see figure showing simplified distribution of photophores); PLO more than its diameter above level of upper base of pectoral fin; eyes not semi-telescopic ➞ 2

Protomyctophum arcticum

2. Second PVO (PVO2) well above upper base of pectoral fin; dorsal fin-rays 20–23; Davis and Hudson Straits = Notoscopelus kroyeri

Second PVO at or below upper base of pectoral fin; dorsal finrays 12–16 ➞ 3

Notoscopelus kroyeri

126

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3. Two Prc; Dn minute ➞ 4

Four Prc; Dn absent ➞ 6

4. Anal fin-rays 17–19; PVO horizontal or almost horizontal; PVO1 not more than its diameter below PVO2; Vn small, obscure; VO2 elevated; Baffin Bay to Hudson Strait, Beaufort Sea = Benthosema glaciale

Anal fin-rays 20–23; PVO inclined; PVO1 more than its diameter below PVO2; Vn larger, well defined; VO2 level ➞ 5

Benthosema glaciale

5. SAO in a straight or slightly angled line; first SAO behind third VO; Davis Strait = Myctophum punctatum

Myctophum punctatum

SAO strongly angled; first SAO in advance of third VO; Davis Strait and Ungava Bay = Symbolophorus veranyi

Symbolophorus veranyi

6. Gill rakers 21 or more; [AOa photophores 6–8]; Davis Strait to Hudson Strait = Lampanyctus macdonaldi

Gill rakers 18 or less ➞ 7



Lampanyctus macdonaldi

7. Pectoral fins shorter, not reaching anal fin origin; AOa photophores 5–8, usually 5–7; Ungava Bay = Lampanyctus crocodilus

Lampanyctus crocodilus

Lampanyctus intricarius

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Pectoral fins very long, extending beyond anal fin origin; AOa photophores 8–10; Davis Strait = Lampanyctus intricarius

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Notacanthidae, Deep-sea Spiny Eels (Family 10) 1. Dorsal fin spines 5–15; anal fin spines 13–21; total gill rakers 12– 17; Baffin Bay and Davis Strait = Notacanthus chemnitzii



Notacanthus chemnitzii

Dorsal fin spines 26–36; anal fin spines 37–42; total gill rakers 22–28; Davis Strait = Polyacanthonotus rissoanus

Polyacanthonotus rissoanus

Oneirodidae, Dreamers (Family 36)

1. Sphenotic spines absent; bones highly cancellous; pectoral finrays usually 18 or more; Davis Strait = Chaenophryne longiceps

Sphenotic spines present; bones not highly cancellous; pectoral fin-rays usually 17 or less ➞ 2

Chaenophryne longiceps

2. No close-set skin spines; body globose; Davis Strait = Oneirodes sp.

Oneirodes sp.

128

Close-set skin spines covering body and fins; body elongate; Davis Strait = Spiniphryne gladisfenae

Spiniphryne gladisfenae

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Osmeridae, Smelts (Family 21)

1. Scale size very small; lateral-line scales 170–220; anal fin-rays 16–26; whole Arctic = Mallotus villosus

Scale size moderate; lateral-line scales 51–73; anal fin-rays 11–18 ➞ 2

Mallotus villosus

2. Vertebrae usually 64–67 (range 63–68); parietal bones of skull more or less contiguous; Bathurst Inlet to Beaufort Sea = Osmerus dentex

Osmerus dentex

Vertebrae usually 61–63 (range 59–65); parietal bones not in contact with each other; Hudson and James Bays = Osmerus mordax

Osmerus mordax

Paralepididae, Barracudinas (Family 27)

1. Dorsal fin absent; chin with projection; anal fin-rays 12–17; Hudson Strait = Anotopterus pharao

Dorsal fin present; chin without projection; anal fin-rays 20–34 ➞ 2

Anotopterus pharao

2. Pelvic fin origin at or behind level of dorsal fin insertion (end of dorsal fin at base); pectoral fin-rays 9–13; anal fin-rays 28– 35; Davis and Hudson Straits = Arctozenus risso

Pelvic fin origin in front of or below dorsal fin; pectoral finrays 14–18; anal fin-rays 20–26 ➞ 3

Arctozenus risso



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3. Rear of premaxilla (rear end of upper jaw) clearly behind nostril; longest gill raker much less than twice as long as next adjacent raker (rakers are in groups of three or more on each base); Davis Strait = Magnisudis atlantica

Magnisudis atlantica

Rear of premaxilla under nostril; longest gill raker more than twice as long as next adjacent raker; Davis Strait = Paralepis coregonoides

Paralepis coregonoides

Phycidae, Phycid Hakes (Family 31) Brian W. Coad and Claude B. Renaud

1. Two dorsal fins, rays (after very elongate third dorsal fin-ray) in first dorsal fin progressively shorter but clearly visible; first and second pelvic rays greatly elongate (the second almost to caudal fin); no barbels on snout; southern Baffin Bay to Hudson Strait = Phycis chesteri

Phycis chesteri

2. Three snout barbels (one short and median, and two long and lateral); end of dorsal and anal fins and lower caudal fin lobe with black pigment blotches; Davis and Hudson Straits = Enchelyopus cimbrius

Three dorsal fins, first fin an elongate ray, second fin with very short rays, similar in size, hair-like and inconspicuous; first and second pelvic rays not markedly elongate; snout barbels present ➞ 2

Rays in second dorsal fin are very short

Two lateral snout barbels; no blotches as above ➞ 3

Enchelyopus cimbrius

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3. Elongated first dorsal fin-ray relatively short, not reaching third dorsal fin, 3.5%–8.8% of total body length; interorbital width less than eye diameter; Baffin Bay to Hudson Strait = Gaidropsarus argentatus

Gaidropsarus argentatus

Elongated first dorsal fin-ray relatively long, reaching third dorsal fin, 11.3%–25.9% of total body length; interorbital width equal to eye diameter; Baffin Bay to Hudson Strait = Gaidropsarus ensis

Gaidropsarus ensis

Platytroctidae, Tubeshoulders (Family 19)

1. Pelvic fins absent; body very compressed and deep (3.0 times or less in standard length); about half of dorsal and ventral body margins with a sharp keel and non-muscular; predorsal margin sharp and one scale wide; Davis Strait = Platytroctes apus

Pelvic fins present; body compressed to rounded and shallow to deep (3.5 times or more in standard length); dorsal and ventral body margins extended by muscular or non-muscular tissue; predorsal margin more than one scale wide and rounded (except in Normichthys) ➞ 2

Platytroctes apus

2. Photophores distinct, absent in young; pelvic fin-rays usually 9; cleithral symphysis without a spine; Davis Strait = Holtbyrnia anomala

Photophores absent or weakly developed; pelvic fin-rays usually 6–8; cleithral symphysis produced as a spine (see figure of Platytroctes apus for spine) ➞ 3

Holtbyrnia anomala

3. Upper jaw extending back to about behind pupil level; frontal bones widest over posterior eye level or behind eye level; one to several large, scale-sized openings in scale pockets behind shoulder girdle; Davis Strait = Normichthys operosus

Upper jaw extending far back beyond eye level (to rear of eye in smaller fish); frontal bones widest over middle of eye level, abruptly narrower before and behind; pores in scale pockets much smaller than scales ➞ 4

Normichthys operosus



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4. Lateral line with papillae or unmarked; photophores present; body deep (3.5–4.0 times in standard length) and snout short (3.8–4.8 times in head length); shoulder pit opening several scale-rows wide behind supracleithrum; premaxilla with 4–8 teeth behind tusks (enlarged anterior premaxillary teeth); Davis Strait = Maulisia mauli

Maulisia mauli

Lateral line with enlarged and modified scales; photophores absent; body relatively shallow (4.3–5.2 times in standard length) and snout long (3.2–3.8 times in head length); no enlarged pit behind supracleithrum; premaxilla with 7–15 teeth behind tusks; Davis Strait = Maulisia microlepis

Maulisia microlepis

Pleuronectidae, Righteye Flounders (Family 58)

1. Numerous (up to 16) large, round, mucous pits on blind side of head; right pectoral fin darkly pigmented distally in adults; Davis Strait = Glyptocephalus cynoglossus

Not as across ➞ 2

Glyptocephalus cynoglossus

2. Dorsal and anal fins with 4–9 distinctive dark bands; large, rough, star-shaped scales on eyed side, and also on bases of dorsal and anal fins; Melville Sound to Beaufort Sea = Platichthys stellatus (note: almost all Canadian Arctic P. stellatus (> 99%) are left eyed)

No distinctive bands or stellate scales; right eyed ➞ 3

Platichthys stellatus 132

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3. Mouth large, extending to or past mid-eye on eyed side; teeth pointed, about equal in extent in both jaw halves ➞ 4

Mouth small, extending almost to below anterior part of eye on eyed side; teeth blunt and conical or incisor like, extent greater on blind side of jaws ➞ 7

Small mouth

Large mouth and eye in notch

4. Uppermost eye lying in a notch on the upper head margin (see previous figure); blind side of body dark in adults; Smith Sound to northern Hudson Bay, Fury and Hecla Strait, Arctic islands, western Banks Island = Reinhardtius hippoglossoides

Uppermost eye not lying in a notch but on side of head; blind side of body white in adults ➞ 5

Reinhardtius hippoglossoides

5. Lateral line on eyed side strongly arched above pectoral fin; scales cycloid on eyed side; caudal fin obviously emarginate (middle rays shortest); Baffin Bay and Davis Strait = Hippoglossus hippoglossus

Arched lateral line

Lateral line on eyed side straight, or gently bent above pectoral fin; scales ctenoid on eyed side; caudal fin not emarginate (middle rays longest) ➞ 6

Straight lateral line

Caudal fin not emarginate, middle rays longest Hippoglossus hippoglossus

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6. Dorsal fin-rays 76–101, usually more than 80; anal fin-rays 60–79; branchiostegal rays 8; Baffin Bay to Hudson Bay = Hippoglossoides platessoides

Hippoglossoides platessoides

7. Head margin above eye strongly concave; lateral line steeply curved over pectoral fin; dorsal fin-rays 61–77; anal fin-rays 45–58; Melville Sound, Dolphin and Union Strait, western Banks Island = Limanda proboscidea

Dorsal fin-rays 66–80, usually less than 80; anal fin-rays 51– 64; branchiostegal rays 7; Melville Sound and Bathurst Inlet = Hippoglossoides robustus

Hippoglossoides robustus

Head not strongly concave above eye; lateral line almost straight over pectoral fin; dorsal fin-rays 64 or less; anal finrays 46 or less ➞ 8

Limanda proboscidea

8. Queen Maud Gulf to Beaufort Sea = Pleuronectes glacialis*

Pleuronectes glacialis

Frobisher Bay = Pleuronectes putnami*

Pleuronectes putnami

* These two species are very similar and cannot easily be separated as adults except by distribution. Characters used in previous studies for adults (head length, pectoral-fin length, and vertebral counts) overlap based on 100 specimens examined for this book. Eggs of P. glacialis are pelagic and large (1.54–1.70 mm), while those of P. putnami are demersal and small (1.1–1.4 mm), and pre-larvae and larvae of the two species differ in the rate of morphological transformation, indicating specific distinction. Mitochondrial DNA data have proved promising for separating these species.

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Psychrolutidae, Fathead Sculpins (Family 46)

1. Spines absent from head and preoperculum; interorbital width greater than twice exposed eye diameter; Beaufort Sea = Psychrolutes phrictus

Spines present but may be blunt; interorbital width less than twice exposed-eye diameter ➞ 2

Psychrolutes phrictus

2. Skin rough with many obvious spines in groups close together; body with bands of pigment; eyes relatively small (bony orbit diameter usually less than bony interorbital width); anal finrays 9–11; lateral-line pores 11–15; Baffin Bay to Hudson Strait, and Beaufort Sea = Cottunculus microps

Cottunculus microps

Skin smooth with only scattered spines (area covered by spines less than naked area); no bands on body; eyes relatively large (bony orbit diameter equal to or greater than bony interorbital width); anal fin-rays 12–14; lateral-line pores about 18–19; Baffin Bay and Davis Strait = Cottunculus thomsonii

Cottunculus thomsonii

Rajidae, Skates (Family 8) One species, in the genus Bathyraja, is known only from egg cases and a juvenile caught in Franklin Bay, Cape Bathurst area of western Amundsen Gulf, and is not in this key.

1. Anterior pectoral fin-rays extending almost to snout tip (may be seen best by shining a light through the flesh); [rostral bar soft and flexible; a thorn between dorsal fins (absent in young); no thorns on disc; thorns on tail midline 21–26; upper-jaw tooth rows 30–34; tail short, 0.9 times body length]; Baffin Bay to Hudson Strait = Bathyraja spinicauda

Bathyraja spinicauda

Anterior pectoral fin-rays extending almost to snout tip, or not



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Anterior pectoral fin-rays not extending almost to snout tip; [rostral bar usually stiff (felt through the skin as a rod)] ➞ 2

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2. No thorns (large spines) present on disc behind shoulder and on much of tail; fine spinules (minute spines) present over whole dorsal surface and underside of tail; [dorsal fins connected, no thorn between them; rostral bar flexible; tooth rows 54–63; tail long, 1.3 times body length]; Davis Strait = Malacoraja spinacidermis

Thorns present variably on disc, always on tail; dorsal surface often smooth but may have spinules ➞ 3

Malacoraja spinacidermis

3. Thorns on body and tail in 2–5 parallel rows, median row much smaller or only in young; tail long (1.3–1.6 times body length); [large thorns on nuchal-shoulder region 20–30; snout very short; tooth rows 30–38; lower surface of disc and tail bare]; Baffin Bay and Davis Strait = Rajella fyllae

Thorns on body and tail in a median row, parallel rows absent or much smaller; tail shorter (1.4 times or less of body length, and often less than body length) ➞ 4

Rajella fyllae

4. Large median thorns from nape to first dorsal fin 11–20; [upper surface covered with thornlets and very rough to touch; tooth rows 27–49; dorsal fins separated by distinct gap, without intervening thorn; tail, 1.0–1.1 times body length]; Baffin Bay to northern Hudson Bay = Amblyraja radiata

Large median thorns from nape to first dorsal fin, 22–51 ➞ 5

Amblyraja radiata

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5. Body thick, flabby; ventral disc darkly blotched with spots or bands in adults, white in young; upper body surface with a bluish-grey to bluish-purple cast, and mottled; [thorns from nape to first dorsal fin 22–32; tooth rows 33–48; dorsal fins separated by narrow but distinct gap, no intervening thorn (but wide in some young, with 1–2 thorns); tail very short, 0.7– 0.8 times body length]; Smith Sound to Hudson Strait, and Beaufort Sea = Amblyraja hyperborea

Body not thick, solid; ventral disc and pelvic fins white to grey ➞6

Amblyraja hyperborea dorsal and ventral views

6. Large spines from nape to first dorsal fin in midline 24–31; tooth rows 56–66; distance from rear orbit margin to pelvic axil usually shorter than pelvic axil to first dorsal fin origin; [dorsal fins separated by narrow but distinct gap, no intervening thorn (except in young with wide gap and 1–2 thorns); tail short, 0.9–1.0 times body length]; Baffin Bay to Hudson Strait = Amblyraja jenseni

Large spines from nape to first dorsal fin in midline 31 or more (33 or more in adults); tooth rows 34–50; distance from rear orbit margin to pelvic axil usually longer than pelvic axil to first dorsal fin origin ➞ 7

Amblyraja jenseni



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7. Lower surface of disc, pelvic fins, and tail uniform dark-brown in young, darker than grey-brown on upper surface (but lower surface is uniform white in adults, upper surface grey white); tooth rows 34–42; large spines from nape to first dorsal fin in midline 31–44 (33 or more in adults); tail long, 1.4 times body length; [dorsal fins connected, no intervening thorn]; Baffin Bay = Rajella bathyphila

Lower surface of disc and pelvic fins whitish with broad grey margin from wing tips posteriorly, median underside of tail grey, grey blotch on each side of vent, lower surface paler than upper surface, which is fawn grey to uniform cream; tooth rows 47–51; large spines from nape to first dorsal fin in midline 40–51 (about 35–36 on tail); tail 0.9–1.0 body length; [large thorns white in contrast to disc; dorsal fins connected or with very small gap, no intervening thorn]; Baffin Bay and Davis Strait = Rajella lintea

Rajella lintea ventral view to show grey blotches

Rajella bathyphila

Rajella lintea

Rhinochimaeridae, Longnose Chimaeras (Family 3)

1. Denticles or knobs absent from upper margin of caudal fin; dental plates ridged and knobby; Davis Strait = Harriotta raleighana

Harriotta raleighana

138

Denticles or knobs present on upper margin of caudal fin; dental plates mostly smooth; Davis Strait = Rhinochimaera atlantica

Rhinochimaera atlantica

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Salmonidae, Trouts and Salmons (Family 22) Salmonids in the sea have a strong silvery colour, obscuring the pigment characters given here. However, careful examination will reveal them.

1. Scales large, 13 or less from dorsal fin origin to lateral line, 115 or less in lateral line (often 110 or less); teeth in lower jaw absent or weak and brush-like (except in Stenodus leucichthys); lower jaw short, not extending back to mid-eye (except in Stenodus leucichthys) ➞ 2

2. Mouth large, maxilla extending to rear margin of pupil, lower jaw obviously projecting beyond upper jaw; teeth in roof of mouth in broad bands; head length enters usually less than 4 times in standard length; body pike-like; Beaufort Sea = Stenodus leucichthys

Scales small, 19 or more from dorsal fin origin to lateral line, 105 or more in the lateral line (often 115 or more); teeth in lower jaw strong and conical; lower jaw long, extending back to or past mid-eye ➞ 9

Mouth small, maxilla not extending to rear margin of pupil, lower jaw not projecting markedly beyond upper jaw; teeth in roof of mouth few or absent; head length enters usually 4 times or more in standard length; body not pike-like ➞ 3

Stenodus leucichthys

3. Single flap between nostrils; body rounded in cross section; gill rakers 14–21; young with parr marks; Ungava Bay to James Bay, and Bathurst Inlet to Beaufort Sea = Prosopium cylindraceum

Single nostril flap

Two flaps between nostrils; body compressed in cross section; gill rakers 18–64; young without parr marks ➞ 4

Double nostril flaps

Young Prosopium cylindraceum with parr marks

Prosopium cylindraceum



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4. Total gill rakers 33 or less; profile of upper lip vertical or overhanging ➞ 5

Total gill rakers 35 or more; profile of upper lip slopes backwards in line with the forehead ➞ 6

Profile of upper lip sloping backwards and vertical or overhanging

5. Forehead concave in profile; adult usually with a hump behind head; maxilla length more than twice its breadth; length of longest gill raker more than one-fifth of interorbital width (1.32%–1.97% of standard length); gill rakers usually 24–33 total (but as low as 19); whole Arctic = Coregonus clupeaformis

6.

Forehead rounded in profile; adult without a hump behind head; maxilla length less than twice its breadth; length of longest gill raker shorter than one-fifth of interorbital width (0.85%–1.35% of standard length); gill rakers 18–25 total; Queen Maud Gulf to Beaufort Sea = Coregonus nasus

Coregonus clupeaformis

Coregonus nasus

Gape of mouth vertical or nearly so, mouth superior; pelvic fins dusky or black in adults [total gill rakers 48–53]; Chantrey Inlet to Beaufort Sea = Coregonus sardinella

Gape of mouth not vertical, mouth not superior; pelvic fins usually pale in adults [total gill rakers usually 35–50] ➞ 7

Coregonus sardinella

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7. Scales between dorsal fin origin and lateral line 7–10; chin speckled or dark (pelvic fin tips dark in some populations); tip of lower jaw is equal to or projects slightly beyond upper jaw; [total gill rakers 36–64, usually 40–50]; whole Arctic = Coregonus artedi

Scales between dorsal fin origin and lateral line 11–12; chin and pelvic fin tips pale; tip of lower jaw not projecting beyond upper jaw; [total gill rakers less than 49] ➞ 8

Coregonus artedi

8. Gill rakers on lower limb of arch 26–31 (total 41–48); body without black spots, fins without white spots; Chantrey Inlet to Beaufort Sea = Coregonus autumnalis

Gill rakers on lower limb of arch 21–25 (total 35–40); black spots with faint halos on body, or white spots on fins, or both; not known in Canadian Arctic marine waters = Coregonus laurettae

Coregonus laurettae

Coregonus autumnalis

9. Total anal rays including small ones at origin 8–13; gill rakers 27 or less; dorsal fin usually with dark or light spots (except Salvelinus alpinus) ➞ 10

Total anal rays including small ones at origin 12–19; gill rakers 16–44; dorsal fin without spots ➞ 14

10. Flank spots dark brown or black and X-shaped on a light background (except sea-run fish, which are silvery); pelvic and anal fins without leading white edges; teeth on head and shaft of vomer bone in roof of mouth; scales along flank obvious, 109–124; Davis Strait to Hudson Bay = Salmo salar

Flank spots light (red, pink, orange, yellow, grey, or whitish) on dark background (except sea-run fish, which are silvery); pelvic and anal fins with leading white edges; teeth on head of vomer only; scales along flank inconspicuous, about 150–250 ➞ 11

Salmo salar



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11. Caudal fin deeply forked; irregular whitish (rarely orange) spots on flank; pyloric caeca about 81–200 (dissection required); parr marks irregular and narrow; rarely in the sea, usually in fresh water; whole Arctic = Salvelinus namaycush

Caudal fin emarginate or square when spread out (can be quite forked in young); orange, pink, or red spots on flank (sea-run fish silvery, but spots still visible); pyloric caeca 13–75 ➞ 12

Pyloric caeca

Salvelinus namaycush

12. Dorsal and caudal fins with vermiculations (dark wavy lines); pelvic and anal fins with snow-white leading edge set off by black behind; tip of lower jaw and roof of mouth black; basibranchial teeth absent on floor of mouth between gills; regularly arranged parr marks along flank 6–12; Ungava Bay to James Bay = Salvelinus fontinalis

Dorsal and caudal fins without vermiculations; white leading edge not usually set off by black behind; tip of lower jaw and roof of mouth whitish; basibranchial teeth present; parr marks irregular and not clearly defined ➞ 13

Salvelinus fontinalis

13. Gill rakers on upper arch usually 7–13, on lower arch 12–19 (mostly about 13); pyloric caeca usually 13–75 (mostly about 45; dissection required); spots usually large (eye diameter) and not numerous; whole Arctic = Salvelinus alpinus

Salvelinus alpinus

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Gill rakers on upper arch usually 3–13, on lower arch 8–15 (mostly about 10); pyloric caeca usually 13–47 (means are 32 or less); spots small (smaller than eye diameter) and numerous; Coronation Gulf to Beaufort Sea = Salvelinus malma

Salvelinus malma

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14. Caudal fin with black spots at least as large as pupil ➞ 15

Caudal fin speckled black or unspotted ➞ 17

15. Black spots on back and caudal fin, the largest as big as eye; scales in first row above and paralleling lateral line 169–229; gill rakers 24–35; Hudson Bay (probably not surviving) and Beaufort Sea = Oncorhynchus gorbuscha

Black spots on back and caudal fin, the largest only as big as pupil; scales in first row above lateral line 140–153; gill rakers 16–30 ➞ 16

Oncorhynchus gorbuscha

16. Small black spots on upper caudal fin lobe only, when present; gums around lower jaw teeth pale; pyloric caeca 45–114; gill rakers 18–25; Beaufort Sea = Oncorhynchus kisutch

Oncorhynchus kisutch

Oncorhynchus tshawytscha

17. Gill rakers short and widely spaced, 16–28; pyloric caeca 140– 249; Hudson and James Bays (not surviving), Amundsen Gulf, and Beaufort Sea = Oncorhynchus keta

Oncorhynchus keta

Gill rakers long and crowded together, 28–44; pyloric caeca 45–115; Bathurst Inlet to Beaufort Sea = Oncorhynchus nerka

Oncorhynchus nerka



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Small black spots on both lobes of caudal fin; gums around lower jaw teeth black; pyloric caeca 90–240 (dissection required); gill rakers 16–30; Coronation Gulf to Beaufort Sea = Oncorhynchus tshawytscha

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Scorpaenidae, Scorpionfishes (Family 43)

1. Anal fin-rays 6–8, usually 7; second dorsal fin-rays 12– 15, usually 13–14; commonest anal and dorsal fin-ray combinations 7 and 13 or 7 and 14; body orange-red with green-black blotches; [knob at chin tip long and sharp]; southern Davis Strait and Hudson Strait = Sebastes fasciatus

Anal fin-rays 7–11, usually 8 or more; second dorsal fin-rays 12–17, usually 14–15; body scarlet red or orange to yellow, usually without green-black blotches ➞ 2

Sebastes fasciatus

2. Knob at chin tip long and sharp; lower unbranched pectoral fin-rays usually 8 or less (rarely 9); 1 or 2 muscle heads attaching to gas bladder with posterior tendon not branched (dissection required); bright red to scarlet but silvery between origin of pectorals and operculum, on cheeks, and in pelvic region; Baffin Bay to Hudson Strait = Sebastes mentella

Sebastes mentella

Knob at chin tip weak and blunt; lower unbranched pectoral fin-rays usually 9 or more (rarely 8); 3 or 4 muscle heads attaching to gas bladder with posterior tendon having 6 branches usually; orange, orange yellow, or golden yellow; Baffin Bay to Hudson Strait = Sebastes norvegicus

Sebastes norvegicus

Somniosidae, Sleeper Sharks (Family 7)

1. Each dorsal fin with a small spine at origin (often concealed by skin); size, medium to 1.2 m; Davis Strait = Centroscymnus coelolepis

No dorsal fin spines; very large, to 4.4 m or more ➞ 2

Centroscymnus coelolepis

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2. Origin of first dorsal fin nearer to the tip of the snout (< 45% of total length) than to tip of caudal fin; space between dorsal fins equal to length from snout tip to first gill opening; Lancaster Sound, northern Baffin Island to James Bay, possibly western Arctic = Somniosus microcephalus

Somniosus microcephalus

Origin of first dorsal fin almost equidistant (> 45% of total length) between tip of snout and tip of caudal fin; space between dorsal fins about 70% of length from snout tip to first gill opening; Beaufort Sea = Somniosus pacificus

Somniosus pacificus

Stichaeidae, Pricklebacks (Family 51)

1. Large cirrus on head above eye, smaller cirrus in front; [dorsal fin spines 50–54; anal fin soft rays 35–40; note that finray counts in combination in this family can help separate species]; Hudson Strait = Chirolophis ascanii

No cirri ➞ 2

Chirolophis ascanii

2. Lower 5 pectoral fin-rays extended and finger like; [dorsal fin spines 57–64; anal fin soft rays 34–44]; whole Arctic = Leptoclinus maculatus

Lower pectoral fin-rays not extended ➞ 3

Leptoclinus maculatus

3. Four lateral lines, a complete mid-flank line, and 1 above and 2 below; last 3 anal fin-rays spine like; [dorsal fin spines 47– 50; anal fin soft rays 30–35]; whole Arctic = Eumesogrammus praecisus

One or two lateral lines ➞ 4

Four lateral lines

Eumesogrammus praecisus

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4. Two lateral lines, a complete one along mid-flank, and one above it ending under anterior half of dorsal fin; an oval blotch between spines 5–6 and 8–10 of dorsal fin; [dorsal fin spines 43–44; anal fin soft rays 30–31]; Davis Strait = Ulvaria subbifurcata

One lateral line; no oval dorsal fin blotch ➞ 5

Ulvaria subbifurcata

5. Lateral line incomplete, ending near mid-body; 4–9 large, dark spots edged posteriorly in white on dorsal fin; [dorsal fin spines 46–51; anal fin soft rays 32–38]; whole Arctic, rare in west = Stichaeus punctatus

Lateral line complete; white-edged, dark spots absent from dorsal fin; [dorsal fin spines 58–85; anal fin soft rays 37–62] ➞6

Stichaeus punctatus

6. Dorsal fin low anteriorly, height rising gradually; snout with frenum (tissue joining lip to upper jaw); [dorsal fin spines 59– 76; anal fin soft rays 41–48]; Beaufort Sea = Acantholumpenus mackayi

Frenum present

Dorsal fin high anteriorly, height rising abruptly; snout without frenum ➞ 7

Frenum absent

Acantholumpenus mackayi

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K ey s

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7. Dorsal fin spines 68–85, usually 71–85; anal fin soft rays 47–62; Davis Strait to Hudson Strait = Lumpenus lampretaeformis

Dorsal fin spines 67 or less; anal fin soft rays 37–45 ➞ 8

Lumpenus lampretaeformis

8. Dorsal fin spines 58–63; pectoral fin-rays 13–15, usually 13– 14; anal fin low anteriorly, higher posteriorly; whole Arctic = Anisarchus medius

Anisarchus medius

Dorsal fin spines 60–67, usually 63–66; pectoral fin-rays 14–17; anal fin of uniform height; whole Arctic = Lumpenus fabricii

Lumpenus fabricii

Stomiidae, Dragonfishes (Family 25)

1. Scales present, or, if lost, body marked by scale pattern ➞ 2

2. Dorsal fin origin far in front of pelvic fin base, behind head; teeth gigantic fangs; premaxillaries not protractile; Baffin Bay to Hudson Strait = Chauliodus sloani

Chauliodus sloani

Scales absent ➞ 3

Dorsal fin origin far behind pelvic fin base, in front of tail; teeth evident but not as across; premaxillaries protractile; Baffin Bay to Hudson Strait = Stomias boa

Stomias boa

3. No floor to mouth; dorsal fin origin over anal fin at rear of body; no chin barbel; Davis Strait and eastern Hudson Strait = Malacosteus niger

Mouth with floor; dorsal fin origin near centre of body, well before anal fin origin; chin barbel present (can be damaged or lost) ➞ 4

Malacosteus niger



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4. Teeth on rear part of upper jaw close together and slanting rearward; OV photophores (upper row on flank between opercle and pelvic fin insertion) 19 or less (14–15 in richardsoni); Davis Strait = Astronesthes cf. richardsoni

Teeth on rear part of upper jaw clearly separate and not slanting rearward; OV photophores 22–25 ➞ 5

Astronesthes cf. richardsoni

5. Dorsal adipose fin present; lower caudal fin-rays not elongate; no line of luminous tissue along flank; Davis Strait = Borostomias antarcticus

Borostomias antarcticus

Dorsal adipose fin absent; lower caudal fin-rays elongated; line of luminous tissue along flank; Davis Strait = Rhadinesthes decimus

Rhadinesthes decimus

Synaphobranchidae, Cutthroat Eels (Family 11)

1. Snout very blunt and rounded; mouth very small, clearly not reaching back to eye; slimy; gill slits tiny, below and anterior to pectoral fin insertion; Davis Strait = Simenchelys parasitica

Snout pointed and elongate; mouth extending beyond eye level; not slimy; gill slits adjacent on throat; Davis Strait = Synaphobranchus kaupii

Gill slits adjacent on throat in Synaphobranchus kaupii

Blunt snout, small mouth, and anterior gill slits in Simenchelys parasitica

Simenchelys parasitica

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Synaphobranchus kaupii

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Zoarcidae, Eelpouts (Family 50) Peter Rask Møller

1. Dorsal fin with a notch, about 16–24 spiny rays long, at rear of fin; dorsal fin origin in front of pectoral fin base level; Davis Strait to Hudson Strait = Zoarces americanus

No such notch; dorsal fin origin over or behind pectoral fin ➞2

Zoarces americanus

2. Pelvic fins absent; pectoral fin-rays 15 or less (range 9–15); scales absent (in Gymnelinae) ➞ 3

Pelvic fins present; pectoral fin-rays 13–24 (13–15 counts in Lycenchelys and Lycodonus, 16 or more, rarely 15, in Lycodes); scales present or absent (in Lycodinae) ➞ 8

Pelvic fins present

3. Gill opening pore like; pectoral fin-rays 6–8(9); flesh gelatinous; Davis Strait = Melanostigma atlanticum

Gill opening not pore-like, extending to middle of pectoral fin base; pectoral fin-rays 9–15; flesh firm (Gymnelus) ➞ 4

Melanostigma atlanticum

4. Origin of dorsal fin well behind end of pectoral fin; predorsal distance 23% or more than standard length; Davis Strait to northern Hudson Bay, Arctic islands = Gymnelus retrodorsalis

Origin of dorsal fin above pectoral fin; predorsal distance 22% or less than standard length ➞ 5

Gymnelus retrodorsalis



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5. Occipital head pores 0–2; pectoral fin-rays 9–12 ➞ 6

Occipital head pores 3; pectoral fin-rays 12–15 ➞ 7

6. Eye diameter 3.5% or more than standard length; bars on body usually not below mid-body; Dease Strait to Beaufort Sea = Gymnelus knipowitschi

Eye diameter 3.4% or less than standard length; bars on body absent or reaching below mid-body; whole Arctic = Gymnelus viridis

Gymnelus knipowitschi

7. Colour uniformly dark brown or with a few irregular light spots; Ungava Bay = Gymnelus barsukovi

Gymnelus barsukovi

8. Body very elongate and rounded; head pores very large; lower jaws without cartilage crests (Lycodonus and Lycenchelys) ➞ 9

Gymnelus viridis

Colour pattern of 8–11 dark, wide bands with a light cellular pattern on a light background; Beaufort Sea = Gymnelus bilabrus

Gymnelus bilabrus

Body moderately elongate and laterally compressed; head pores minute or moderate; lower jaws with cartilage crests (Lycodes) ➞ 13

Lower jaw cartilage crests

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9. Dorsal and anal fin-rays with bony plates at base (6–10 free plates in front of dorsal fin); branchiostegal rays 5; Baffin Bay and Davis Strait = Lycodonus mirabilis

Dorsal and anal fin-rays without bony plates at base; branchiostegal rays 6 (Lycenchelys) ➞ 10

Lycodonus mirabilis

10. Abdomen without scales; dorsal fin with dark and white blotches; Nares Strait to Ungava Bay, and Amundsen Gulf = Lycenchelys kolthoffi

Abdomen scaled; dorsal fin uniformly dark brown ➞ 11

Lycenchelys kolthoffi

11. Predorsal area naked, length less than 19% of standard length (15.1%–18.2%); pectoral fins light with dark dorsal margin; Baffin Bay to Hudson Strait = Lycenchelys muraena

Predorsal area scaled, length greater than 17% of standard length (17.1%–23.1%); colour of pectoral fins different ➞ 12

Lycenchelys muraena

12. Suborbital head pores 7 (rarely 6); pectoral fins usually uniformly dark; total vertebrae 127–137; Davis Strait = Lycenchelys paxillus

Lycenchelys paxillus

Lycenchelys sarsii

13. Pre-anal length usually 45% or more than standard length (45%–61%, rarely 42%–44%) (“short-tailed”); lateral-line system a single medio-lateral one (see figure under couplet 20 of Zoarcidae); otolith with a filled sulcus (dissection required) ➞ 14



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Suborbital head pores 6 (rarely 7); pectoral fin usually light anteriorly, dark posteriorly; total vertebrae 121–126; Davis Strait = Lycenchelys sarsii

Pre-anal length usually 45% or less than standard length (34%– 45%, rarely 48%–50%) (“long-tailed”); lateral-line system a single ventral, double, ventral + half medio-lateral or ventromedio-lateral one; otolith with a deep sulcus ➞ 20

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14. Body naked or with very few scattered scales ➞ 15

Body scaled ➞ 17

15. Teeth on jaws, vomer, and palatines blunt, robust, and blond; Spence Bay to Beaufort Sea = Lycodes jugoricus

Teeth on jaws, vomer, and palatines pointed ➞ 16

Lycodes jugoricus

16. Body with 3–7 triangular white marks, one horseshoe-shaped band on neck (sometimes reduced in large specimens); whole Arctic = Lycodes mucosus

Lycodes mucosus

17. Pectoral fin-rays 22–24; colour more or less pink with 6–8 white bars dorsally; juveniles grey brown with pink pectoral fins; Baffin Bay to Hudson Strait = Lycodes luetkenii

Body with 7–13 narrow white bands dorsally, one straight white band on neck; whole Arctic = Lycodes polaris

Lycodes polaris

Pectoral fin-rays 17–22; colour not pink ➞ 18

Lycodes luetkenii

18. Margin of dorsal and anal fin with uninterrupted black-andlight horizontal bands in adults; neck scaled; vertebrae, 99–105; Frobisher Bay to Hudson Strait = Lycodes lavalaei

Margin of dorsal and anal fin not with uninterrupted black-and-light horizontal bands; neck naked; vertebrae, 99 or less ➞ 19

Lycodes lavalaei

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19. Peritoneum not pigmented (dissection required); anterior scales begin at 28%–34% of standard length from tip of snout; snout convex; lower pectoral fin-rays moderately free; whole Arctic = Lycodes reticulatus

Lycodes reticulatus

Peritoneum weakly pigmented; anterior scales begin at 30%– 56% of standard length from tip of snout; snout concave; lower pectoral fin-rays very free; whole Arctic = Lycodes seminudus

Lycodes seminudus

20. Lateral-line system a ventro-medio-lateral one ➞ 21

Lateral-line system a single ventral, double, or ventral + half, medio-lateral one ➞ 23

Lateral-line types: 1 = medio-lateral, 2 = ventro-medio-lateral, 3 = ventral/half medio-lateral, 4 = double, 5 = single ventral

21. Pectoral fin-rays 22–23; isthmus width more than 5% of standard length (5.1%–7.1%); scale rows above anal fin origin 28–34; Baffin Bay to Hudson Strait = Lycodes mcallisteri

Pectoral fin-rays 16–18; isthmus width less than 5% of standard length (2.4%–4.9%); scale rows above anal fin origin less than 28 ➞ 22

Lycodes mcallisteri



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22. Scale rows above anal fin origin to dorsal fin 9–21; peritoneum speckled (dissection required); body with or without light vertical bars; bend of lateral line behind anal fin origin; Dease Strait to Beaufort Sea, Prince Patrick Island = Lycodes marisalbi

Lycodes marisalbi

Scale rows above anal fin origin to dorsal fin 21–27; peritoneum dark brown; body uniformly brown; bend in lateral line above anal fin origin; Beaufort Sea = Lycodes sagittarius

Lycodes sagittarius

23. Lateral-line system a single ventral one (medio-lateral rows of neuromasts absent) (see figure under couplet 20) ➞ 24

Lateral-line system a double or ventral + half, medio-lateral one ➞ 26

24. Dorsal part of body grey brown with irregular dark stripes; teeth in jaws and palatines blunt; postorbital head pores 4, occipital head pores 3; Davis Strait to Hudson Strait = Lycodes vahlii

Body uniformly grey, brown, or black; teeth in jaws and palatines pointed; postorbital head pores 1–2, occipital head pores 0–2 ➞ 25

Lycodes vahlii

25. Anterior half of body (45%–58% of standard length) without scales; pectoral fin-rays 16–19; precaudal vertebrae 19–21 (dissection required); Baffin Bay to Ungava Bay, Beaufort Sea = Lycodes adolfi

Lycodes adolfi

26. Medio-lateral neuromasts on trunk and tail; vertebrae 105–124 ➞ 27

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Entire body scaled; pectoral fin-rays 19–23; precaudal vertebrae 21–24; northwest of Ellesmere Island = Lycodes frigidus

Lycodes frigidus

Medio-lateral neuromasts on tail only; vertebrae 93–107 ➞ 29

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27. Pectoral fins more or less emarginate, base without scales; number of neuromasts in medio-lateral row greater than 65; length of premaxillary teeth row equal to palatine teeth row; Nares Strait to Hudson Strait, Beaufort Sea = Lycodes eudipleurostictus

Pectoral fin rounded, base scaled; number of neuromasts in medio-lateral row less than 65; length of premaxillary teeth row usually longer than palatine teeth row ➞ 28

Lycodes eudipleurostictus

28. Dorsal part of body dark with 4–9 white marks; predorsal row of neuromasts present; palatine teeth length greater than 0.32% of standard length (0.33%–0.43%); palatine teeth 5–13; Davis Strait to Hudson Strait = Lycodes esmarkii

Lycodes esmarkii

Body uniformly light to dark brown; predorsal row of neuromasts absent; palatine teeth length less than 0.32% of standard length (0.21%–0.31%); palatine teeth 2–6; Davis Strait = Lycodes terraenovae

Lycodes terraenovae

29. Nostril tubes short (1.0% of standard length or less); Baffin Bay to Hudson Strait = Lycodes paamiuti

Nostril tubes long (0.8% of standard length or more) ➞ 30

Lycodes paamiuti

30. Abdomen and predorsal area without scales; Davis Strait to Dease Strait = Lycodes pallidus

Lycodes pallidus

Lycodes squamiventer

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Abdomen and predorsal area scaled; Beaufort Sea = Lycodes squamiventer

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Family and Species Accounts

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INTRODUCTION Brian W. Coad

Following is an explanation of the format of the family and species accounts.

Family Account The family accounts follow the arrangement in the American Fisheries Society (AFS) list (see the bibliography herein) with species given alphabetically within each family. Some deep-water families are not covered in the AFS list, and Eschmeyer’s Catalog of Fishes (accessible online) is followed for these families. Each family account has the scientific, English, and French names of the family. The family account serves to place the Arctic species in a world and Canadian setting through information on distribution and number of species. General features of anatomy, biology, and importance are laid out and are meant to avoid redundancy and repetition in the species accounts.

Species Account The species accounts have a standard format as follows.

Scientific and Common Names The scientific name comprising the genus and species, with the author and date, heads each species account and is taken primarily from the AFS list (Page et al., 2013). See also the section “Checklists of Species” for further information on the sources of scientific names. The scientific name is followed by the English and French common names. These names have been advanced by the AFS (Page et al., 2013) in an effort to standardize usage, as some species have many common names. However, the AFS list does not cover all the species dealt with here. Fish found in deep waters, seen only by research scientists, do not have true common names similar to those that are familiar to local people, anglers, and naturalists. They may have names that have been especially coined for them, so-called

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book names. Coad, with Waszczuk and Labignan (1995) and then FishBase (Froese & Pauly, 2013) are the main sources for common names not in the AFS list. English names of fishes are capitalized, and French names are not, following usage by the AFS.

Illustration A black-and-white line drawing shows the general features of the species and is usually based on an Arctic specimen or has been checked against specimens. Some further anatomical details can be found, illustrating the keys.

Common Names Common names may also require some explanation, and this too is given. Throughout the text generally, the common names of fishes, birds, and mammals are capitalized in order to avoid confusion; for example, Arctic Sculpins (more than one Myoxocephalus scorpioides) has a different meaning from Arctic sculpins (more than one species of Sculpins found in Arctic waters). Common names of families are also capitalized for clarity, but some names, for example, sharks, are not as they comprise several families. This section of “Family and Species Accounts” contains various local or Indigenous names as well as other common names in English and French. Many species have no Indigenous name, being from deep waters and seen only by scientists. Some sources give local or Indigenous names without specifying the language or dialect. Some species may have many commonly used names in Indigenous languages, but only the most generally used ones are given. Variant names result from many causes: local spelling or pronunciation differences; translation to Roman orthography; single, dual, and plural forms; regional dialects; and/or nuances regarding the fish themselves (e.g., reference to different life stages, place of capture, or method of capture). As yet, no standardization or comprehensive research has occurred regarding Indigenous names of all marine fishes; accordingly care should be exercised when using these names. Local names in Indigenous languages or dialects (such as Inuktitut, Inuvialuktun, Gwich’in, and their dialects) may be found in traditional knowledge reports developed by resource co-management

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boards in the north and also in many environmental impact assessments conducted to assess resource developments in the various areas – for example, various online sources of the Naonaiyaotit Traditional Knowledge Project; Kitikmeot Inuit Association; Graves & Hall (1985); T. Williamson (1997); Haszard & Shaw (2000); the Inuktitut Living Dictionary (accessed 27 August 2014 at www.livingdictionary.com); Inuvialuit Renewable Resources Committee (2003) at www.jointsecretariat.ca/pdf/js/IHS10yrDataMethodsReport.pdf; Papik, Marschke, & Ayles (2003); Hart & Amos (2004); Inuuvik Community Corporation, Tuktuuyaqtuuq Community Corporation, & Aklarvik Community Corporation (2006); Thompson & Millar (2007); and Gwich’in Renewable Resource Board (2009) at www.grrb.nt.ca/pdf/GHS/Harvest_Study_Report_FINAL09Web. pdf, etc. McAllister, Legendre, and Hunter (1987) recorded Indigenous fish names from Arctic Canada under the heading “Inuktitut” in the broad sense, which then included Inuvialuktun and various dialects. Local names without a language specifically given in parentheses after the fish name are then Inuktitut in this broad sense. Berkes and Mackenzie (1978) give Cree fish names from James Bay. Greenlandic common names have been formed for all species occurring in waters around Greenland, mostly based on Danish names (Møller et al., 2010), and these are indicated by Danish/ Greenlandic. Some are true local names in Kalaallisut and are indicated by Greenlandic.

Taxonomy This part of “Family and Species Accounts” gives information on the meaning of the Latin- or Greek-derived genus and species names. Not all name origins are evident, not being given by the original describer, and they are open to more than one interpretation. Where there are two or more species in a genus, the explanation for the genus name is given where it first appears and is not repeated under each species. Scientific names are not static and may change as the understanding of relationships develops. The species may appear in different works under more than one genus or species name, and these names are mentioned here to aid retrieval of information. Authors may disagree on the genus to which the species belongs. Synonyms from marine waters of Arctic Canada and synonyms from elsewhere that are often used in Arctic Canadian literature are listed. Some species have a worldwide distribution in deeper waters, and their synonyms may be listed as potentially relevant. Other synonyms for freshwater salmonids and other freshwater species in the far northern Arctic are also given as a guide to older literature. Complete synonymies of some species are quite extensive and may include names that are never used in the Arctic, Canada, or indeed North America, and so are not included here. Readers are referred to Eschmeyer (1998; also online as Catalog of Fishes at the California Academy of Sciences) for more references on synonymies, type localities, and conflicting viewpoints. Opinions vary on the relative validity of species and subspecies names, and these too are summarized in Eschmeyer (1998, and online). Some ichthyologists do not favour subspecies and maintain that any diagnosable taxon should be accorded species rank



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(the phylogenetic species concept; see Kottelat (1997) for discussion). Few subspecies in Canadian Arctic waters have found general acceptance.

Description This section gives a series of characters that describe the species. Some species have a unique character that distinguishes them from all other fish species in Arctic Canadian waters, and others can only be identified by a combination of characters. The “Keys” section should be referred to for any species without a unique character. Species identification should, in any case, depend on a series of characters. Characters included are mostly externally visible, important ones, especially uniquely distinguishing character(s). General family characters, found in all species of a family, will be addressed under the family account that precedes the species accounts. Some critical identification characters are illustrated in the “Keys” section. The glossary is another source for explanations of structures. One important type of character is the ranges in counts of such structures as fin-rays, scales, and gill rakers that are not apparent on an illustration of a single specimen. The range given here is the known range for the species, not just for the Arctic specimens. Few Arctic species have been studied in detail using material solely from the Canadian Arctic, and restricting counts to the known data would be misleading. New material could easily fall outside the range from a few known specimens. However, for species widely distributed outside the Canadian Arctic, the ranges in countable characters may well be much wider than those encountered in Canadian specimens. Colour in life is given, but some species are known mostly from preserved and faded museum specimens, and the description is then of pigment patterns that nonetheless may be distinctive. Size is given as either total or standard length, being the maximum size for the species, not just Canadian Arctic fish. Maximum weight may be given for larger or commercial species. Lengths and weights for Arctic populations are given in the “Biology” section of the account whenever studies have been undertaken.

Habitat Environmental parameters are described in this section, including temperature, salinity, depth range, association with ice, movements, migrations, and bottom conditions. Depth ranges for some deep-sea species may include the minimum and maximum depths at which a net was hauled, and the specimen(s) caught somewhere in this range.

Biology The biology presented is a summary, rather than an extensive critical review of knowledge. It is a synthesis for those species that have been studied intensively. Other species are poorly known, and information on their biology is mostly anecdotal, being chance observations on a few specimens. Some migratory species may have detailed and numerous accounts of biology on the freshwater phase and little on the marine phase. Freshwater biology is not given here other than to mention its importance. Where there is little or no information on biology from Canadian Arctic waters, data are

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summarized from studies in the Alaskan Beaufort Sea, the northwestern Atlantic Ocean (Labrador-Greenland), and the Russian or European Arctic. In some cases the only available information comes from remoter areas. Information without Canadian Arctic localities can be considered as a general summary, indicative of species biology but needing more work in Canadian waters to confirm its validity there. The type of data presented here includes ecological significance, age and growth, reproduction, food, predators, significant infestations, and unusual behaviours. Parasites are not listed extensively, because summary works on parasites of Canadian fishes are readily available (Margolis & Arthur, 1979; Margolis & Kabata, 1984–96; T. McDonald & Margolis, 1995).

Importance There is little commercial fishing in Canadian Arctic waters, but catches in weight and value are given where known and potential uses are suggested. Some species are used locally as a food for humans and, in the past, for dogs. The use of marine catches for dog food is in decline as machines replace the more traditional dog sled for transport. Importance elsewhere is briefly summarized. Many smaller species are important in the ecology of Arctic waters as food for other fishes, for mammals, and for birds, but this is mentioned in the “Biology” section. All species form an integral part of the ecosystem, so an indication of “no importance” is more a measure of our ignorance.

Distribution Distribution is outlined both by a map and by text. The text gives the world and Canadian distribution outside Arctic waters for each species as well as a summary of the distribution in the Canadian Arctic. Some distributional information from literature sources is not specific enough to warrant a spot on a distribution map but can be cited here. Unusual or problematic distributions may also be discussed here. The source for rare records is given and may include a museum catalogue number for the collection; for example, sources may include ARC (Atlantic Reference Centre, St Andrews, New Brunswick); CMNFI (Canadian Museum of Nature, Ottawa); NRM (Naturhistoriska riksmuseet, Stockholm); and ZMB (Museum für Naturkunde, formerly Zoologischen Museum, and Humboldt-Universität, Berlin). The main areas of distribution are given from east to west. They begin with Nares Strait and Smith Sound in the north between Ellesmere Island and Greenland, including localities and hamlets where nearby collections have been made, such as Aujuittuq (Grise Fiord) on southern Ellesmere Island, Nirjutiqavvik (Coburg Island) off southeast Ellesmere Island, and Ikpiarjuk (Arctic Bay) and Mittimatalik (Pond Inlet) on northern Baffin Island; then southward through Baffin Bay and Davis Strait (with two main inlets on Baffin Island – Cumberland Sound and Frobisher Bay); then westward through Hudson Strait leading into Ungava, Hudson, and James Bays; northward into Foxe Basin and the Gulf of Boothia, the Arctic islands (including the northernmost group – the Queen Elizabeth Islands – and also Somerset, Prince of Wales, Victoria and Banks

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islands), along with the hamlet of Qausuittuq (Resolute) and Resolute Bay; and along the Arctic shore of the North American land mass including Queen Maud Gulf, Dease Strait, Melville Sound, Bathurst Inlet, Coronation Gulf, Dolphin and Union Strait, Amundsen Gulf, and the Beaufort Sea. These named areas reflect access and collecting effort. Older scientific literature uses English place names. Many of these places now have traditional names, which are given with the English name in parentheses. Some localities lie on a bay, an inlet, or a sound that seems to retain the English name; for example, Iqaluit, the capital of Nunavut, was once called Frobisher Bay and lies at the inner end of Frobisher Bay. Iqaluit means “place of fish.” Distributions off Greenland follow Møller et al. (2010) where northwest ranges from around 68°30' N northwards in Davis Strait and Baffin Bay to the northernmost point of Greenland; southwest covers from about 68°30' N to the southernmost tip of Greenland (both northwest and southwest extend out to the midline between Canada and Greenland); and southeast and northeast refer to the eastern side of Greenland, more remote from Canadian Arctic waters. The spot distribution map is based mainly on specimens held in the Canadian Museum of Nature, Ottawa, with a smattering of holdings in other museums, data from fisheries surveys and exploratory cruises (cited as “cruise data”), and an analysis of literature. Literature records and cruise data, unlike museum specimens, are unverifiable (except by revisiting the locality). If there is considerable doubt in the accuracy of literature records, as to either locality or species identity, the record is omitted. Some dubious records are included but are commented on in the text. Generally, literature records fill in gaps between verified records. Each distribution spot may represent more than one collection or, given the map scale for widely distributed species, several adjacent collections. Some localities are brackish waters near or in river mouths. Marine species from such localities are mapped because this may be the only record for the area, but the fishes listed in this book as brackish-water (primarily freshwater) species are not mapped (see “Brackish Water Species” in the “Checklists of Species” section). Species with few records (five or less) are documented in the text. More numerous records are available in a database held at the Canadian Museum of Nature and at Fisheries and Oceans Canada, Winnipeg. Map records range from one for such deep-sea species as Argyropelecus gigas to 18,354 records for Reinhardtius hippoglossoides, a commercially sought species. The total number of records analysed was 112,552 (including rejected records). Mapping reflects both rarity and commonness of the species but also accessibility. There will be fewer records of species that are found in the deep sea, are large and fast moving, under ice, in coastal areas away from population centres, in open waters between land masses, in areas not surveyed for fisheries purposes, non-commercial, or naturally rare. In the following map that shows all map points some spots may represent a single specimen and species and so are not a thorough survey of fish diversity at this locality. The most intense collecting efforts have been carried out in Baffin Bay, Davis Strait, Hudson Strait, Ungava Bay, and the Beaufort Sea as part of commercial fish

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Distribution of the marine fishes of Arctic Canada

and shrimp surveys and/or environmental assessments. Collections are sparse in the High Arctic with its ice cover and inaccessibility and generally in the open waters between islands, which are often ice covered.

Sources Sources are the literature on which the family and species accounts are based, in addition to direct observation of specimens in the field and as preserved material. To avoid repetition, general works



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covering many species are listed in the bibliography and are not cited under each family and species. Literature cited under the family account is generally not repeated in the species accounts. Literature for deep-sea or rare fishes is often limited to taxonomic descriptions, and commoner species have extensive literature sources, for which see the online bibliography (Coad & Reist, 2016). This latter is a bibliography on Arctic fishes, their biology, and distributional information for mapping, and not all works therein are cited in this text because of duplication of content.

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Family Myxinidae Hagfishes, Myxines

Claude B. Renaud

in tide pools. Their body fluids are isosmotic with sea water. They are morphologically hermaphroditic (male and female), but only one of the two gonads is functional in a given individual. Eggs are large (about 20 mm long) and yolky with hooked tufts on each end and can number up to 30 per functional female. There is no metamorphosis (development is direct). Their commercial importance is primarily of a negative kind, due to the fouling of nets with slime when they attack commercially important fishes and destroy the fishes themselves. There is also a small market for their skin, which is transformed into expensive “leather” goods.

sources: B  rodal & Fänge (1963); Wisner & McMillan (1995);

Fernholm, in J. Jørgensen, Lomholt, Weber, & Malte (1998); Martini, in Jørgensen et al. (1998); Mok, Saavedra-Diaz, & Acero (2001); Department of Fisheries and Oceans (ed. S. Rowe) (2010).

Collectively Hagfishes are also known as Hags or Slime Eels. Use of the latter name is discouraged because the only feature that the distantly related Hagfishes and eels share is their general body shape. Hagfishes are generally exclusively marine (some species, however, have been reported entering estuaries) and are distributed in the temperate zones of both hemispheres, as well as the Gulf of Panama and off the Caribbean coast of Colombia. There are over 70 species (3 in Canada, but only 1 in the Arctic) in seven genera and two subfamilies (Myxininae having one pair of gill openings, and Eptatretinae with multiple pairs of gill openings). Some authors elevate these subfamilies to families. The maximum total length is about 110.0 cm. Their body is elongate and naked (scaleless). Species vary in the number of external gill openings they possess, from 1 to 16 pairs. A fin fold extends continuously from the ventral body surface around the tail and onto the dorsal surface. This fin fold is not differentiated into anal, caudal, and dorsal fins, nor does it contain any fin-rays. There are no paired fins. The eyes are covered by skin. Two pairs of barbels surround the single terminal nostril, and two pairs of barbels flank the mouth. The nostril leads directly to the pharynx. They have no jaws, but the oral cavity possesses a single palatine tooth and two pairs of lingual plates bearing teeth. The teeth are made of keratin. The tooth arrangement on the tonguelike piston constitutes a biting apparatus that can remove chunks of flesh. Slime glands produce large amounts of mucus that is excreted through 57–200 pores, depending on the species, aligned ventrally on either side of the body. Hagfishes exhibit a knotting behaviour whereby their body is thrown into a knot that gradually slips along the body to rid them of excess slime or to enable them to bite flesh off their prey. They feed as scavengers on dead or dying invertebrates, fishes, and marine mammals. Many species generally live in close association with soft-bottom substrates (sand or mud), but some species are primarily found on hard bottoms where burying is not possible. Hagfishes are known to inhabit depths as great as 5,000 m, and some species have been seen swimming in surface waters or have been collected

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Myxine glutinosa Linnaeus, 1758

Atlantic Hagfish, myxine du nord

Myxine glutinosa, ventral view

common names: A local name is Ivik (Greenlandic). Other common names are Borer, Common Hag, Northern Hagfish, and Slime Eel. taxonomy:  The genus comes from the Greek myxa (mucus, slime) and the Latin -inus (pertaining to), and the species name comes from the Latin glutinosus (viscous, sticky), referring to the slime production. It belongs to the subfamily Myxininae. Myxine glutinosa Linnaeus, 1758, was based on more than one species from various localities, including eastern North Atlantic material. Apparently no lectotype has been designated. Myxine limosa Girard, 1859, and

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Myxine glutinosa

M. atlantica Regan, 1913, are synonyms based on western North Atlantic material (the first from the Bay of Fundy, New Brunswick, and the second from off Nova Scotia). There is a debate as to whether the Atlantic Hagfish on both sides of the North Atlantic Ocean represents distinct species (i.e., M. limosa in the western North Atlantic and M. glutinosa in the eastern North Atlantic) or a single one as treated here.

habitat:  This species is bottom living, occurring in coastal waters at depths of 20–1,200 m in or within a few meters of soft, muddy bottoms in waters below 12°C and in salinities of 30‰ or more. Capture depths are 409–1,156 m in Davis and Hudson Straits. They inhabit shallow transient burrows that parallel the sediment surface. They are non-migratory.

description:  This species is distinguished by the absence of paired fins, the long barbels on the snout, and the nostril at the tip of the head. The body is naked (without scales), elongate, nearly cylindrical anteriorly, and compressed posteriorly. There is a single terminal nostril surrounded by a pair of anteriorly directed barbels on each side. A pair of barbels is directed posteriorly on each side of the inferior jawless mouth. There is a single nasal sinus papilla visible on the undersurface of the nostril, one keratinous palatine tooth, and 28–40 teeth in total on the four lingual tooth plates (internal character). Only the anterior-most tooth on each lingual plate is bicuspid, the rest being unicuspid. The eyes are covered with skin and muscle. Usually there are six internal gill pouches on each side of the body (more rarely five or seven) emptying into a single external gill opening on either side of the body (larger one on the left side), anterior to the ventral fin fold. There are no true fins, but a continuous fin fold unsupported by fin-rays begins dorsally about two-thirds of the distance from the head, runs around the tail and ventrally to about one-third of the distance from the head. Total slime pores along the lower aspect of the body on each side number 85–118. Body colour in live, active specimens is bright pink or pink grey and in resting or preserved specimens is pale grey, deep slate grey, reddish brown, or dark purple; the head colour is either pale or dark, usually with a narrow whitish streak extending along the dorsal midline from the fin fold anteriorly. This species attains 95.0 cm total length, but usually under 51.0 cm.

biology:  This species is a scavenger, feeding on dead or moribund invertebrates (hermit crabs, limpets, nemerteans, polychaetes, priapulid worms, shrimps), but also on fishes (Atlantic Cod, Atlantic Hagfish eggs, Atlantic Herring, Atlantic Mackerel, European Ling, European Sprat, Haddock, hakes, Pollock, Porbeagle, Spiny Dogfish, Sturgeons), marine mammals (whales) of various species, and even bird remains lying on the sea floor. However, in areas where their densities are high, it has been suggested that they are primarily predators and secondarily scavengers. It does not possess jaws but has four laterally biting tooth plates that have sharp cusps. When feeding on larger and tougher tissue, these teeth in conjunction with the body-knotting behaviour – whereby the tail loops to make a simple knot that proceeds anteriorly through muscular contractions – can tear flesh to the extent that the hagfish can completely bury itself inside the animal it consumes. It may enter large fish through the mouth or anus. Atlantic Hagfish eggs are preyed upon by conspecifics, Atlantic Cod, and halibut (species not specified), and Atlantic Hagfish individuals are preyed upon by Atlantic Cod, Spiny Dogfish, White Hake, Harbor Porpoise, and Harbor Seal. There is no larval stage. Growth rate is unknown, and there is no known way to age them. Although individuals greater than 40 cm total length generally possess both sexes, only one of these is functional in a given individual. There are 1–30 oval eggs per female, egg size varies from 16 mm to 27 mm in length and 5 mm to 10 mm in width, and mature eggs are whitish and possess a tuft of anchortipped filaments at both ends, used for attaching to other eggs. The female may spawn throughout the year, based on the fact that mature females are found year round. No spawning grounds have yet been confirmed, but spawning is believed to occur at depths of 90–270 m. Adults can remain buried in mud for long periods with only the tip of the snout projecting and are as active at night as during the day. They can be locally abundant, especially when attracted by food, which is detected by scent. Hundreds of individuals can converge on a food fall within minutes of its appearance. The highest density reported is 500,000 individuals per square kilometer (sq km), and the average is 59,700 individuals per sq km. The swimming speed is 0.25 meters per second (m/s), with bursts of speed of 1 m/s, and they can swim backwards almost as easily as forwards. The large amount of slime they produce when encountering a threat is used for defence against predators, and, when feeding, to discourage competitors. Since the slime is also lethal to the hagfish that secretes

Myxine glutinosa, ventral head



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it, the knotting behaviour is used to remove excess slime and prevent asphyxiation. There are no known parasites of Atlantic Hagfish.

importance:  Atlantic Hagfish are known to opportunistic-

ally feed on commercially important fishes (Atlantic Cod, Atlantic Herring, Atlantic Mackerel, Haddock, and Spiny Dogfish) that are caught with fishing gear, including bottom longlines and bottom set gill-nets. Since the 1990s Canadian and American fishermen have harvested Atlantic Hagfish in the Gulf of Maine and sent them frozen to South Korea, where the hagfish skin is manufactured into expensive “leather” goods. Between 1991 and 1999 over 250 million Atlantic Hagfish greater than 50.0 cm in total length were shipped to South Korea. They have been used in heart research (as they possess several hearts with diffused pacemakers) and in immunology (wounds heal very slowly but do not become infected).

distribution:  The Atlantic Hagfish is found in Davis and Hudson Straits, on both sides of the North Atlantic Ocean, and just into the Barents Sea. In the eastern North Atlantic it occurs in the North, western Baltic, and Mediterranean Seas. In the western North Atlantic it occurs from Davis Strait (63°55' N on the Canadian side and 66°53' N on the Greenland side), on the east side of Greenland (65°23' N), and south to Campeche Bank, Gulf of Mexico (24°25' N). Møller, Feld, et al. (2005) suggested that records of M. glutinosa from depths greater than 800 m on the Canadian side of Davis Strait may belong to Myxine jespersenae Møller, Feld, Poulsen, Thomsen and Thormar, 2005 (described from off West Greenland, eastern Davis Strait), but this has not been confirmed.

Distribution of Myxine glutinosa

sources:  Karrer (1973); Hudon (1990a); Møller, Feld, Poulsen, Thomsen, & Thormar (2005).

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Family Petromyzontidae Lampreys, Lamproies

Claude B. Renaud

The Northern Hemisphere Lampreys, as they are also collectively called, comprise 36 species (11 in Canada, only 1 in Arctic marine waters) variously grouped under four to eight genera depending on authors. They were previously treated as a subfamily of a more inclusive group that included the Southern Hemisphere Lampreys. They are distributed between 18° N and 72° N. The maximum total length of larvae is about 25 cm, and of adults is 120 cm. The body is elongate and naked (scaleless). A single median nostril on top of the head ends in a blind sac. There are seven branchial openings on each side of the body. Lampreys have a caudal fin and either one or two dorsal fins depending on the genus. There are no anal or paired fins. The eyes are covered by skin in the larva and exposed in the adult. They have no jaws but possess at the adult stage a round suctorial oral disc lined internally with keratinous teeth – some as single teeth and others grouped on plates (laminae). Three keratinous tooth plates are also found on the tongue-like piston. It is the action of the tooth-bearing, tongue-like piston that enables the adult lamprey to feed on its host. At the junction of the oesophagus (above) and the branchial tube (below) in the adult is the velar apparatus, which bears forwardly directed tentacles. The sexes are separate. All lampreys spawn in fresh water and die a few days after spawning. The adults use their oral disc to arrange gravel into a more or less circular nest or redd into which they deposit the eggs and milt. Fertilization is external, but the two sexes intertwine their bodies so that the urogenital papillae are in close proximity during the release of the sexual products. Eggs are about 1 mm in diameter, and fecundity can vary between 500 and 300,000 eggs depending on the species. A number of years are spent as a filter-feeding larva, called an ammocoete, living in the soft bottom of streams and rivers. After metamorphosis, in which the horseshoe-shaped oral hood of the ammocoete transforms into the round oral disc of the adult, species lead either a “parasitic” mode (15 species) or a non-parasitic mode (21 species) of life. Non-parasitic species are confined to fresh water and do not feed during an adult life that lasts less than a year. “Parasitic” species live up to 3.5 years

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as adults and either are restricted to fresh water or are anadromous, and feed predominantly on either the blood (those are parasites) or the flesh (more properly these are predators) of fishes and marine mammals. A pair of bean-shaped buccal glands, just behind the oral disc, secretes lamphredin, an anti-coagulant that keeps the blood of the host flowing during feeding. The commercial importance of Lampreys is viewed both positively and negatively. They are primarily recognized as harmful to commercially important fishes, especially in the Laurentian Great Lakes. Their impact in marine waters is, however, most probably negligible. Some anadromous species on their upstream spawning migration are harvested for human consumption, notably in Alaska, but not in Arctic Canada.

sources:  Berg (1962); Vladykov & Kott (1979); Holčík (1986); Renaud (1997, 2011); H. Gill, Renaud, Chapleau, Mayden, & Potter (2003).

Lethenteron camtschaticum (Tilesius, 1811)

Arctic Lamprey, lamproie arctique

common names:  Local names are Nemeryaq and Nû-mugû-shûk (Alaska). The older, and occasionally the recent, Russian literature uses the common name Pacific Lamprey for this species, creating confusion because that common name has a long-standing use for another lamprey species, Entosphenus tridentatus (Gairdner in Richardson, 1836), which also occurs in North America and Asia. taxonomy:  The genus comes from the Greek lethe (oblivion) and enteron (intestine), referring to the degeneration of the intestine. The species name comes from the Russian Kamchatka and the Latin –icus (pertaining to), referring to the region from which this species was originally described. It belongs to the subfamily Lampetrinae. Originally described under the genus Petromyzon, it has also been treated under the genera Lampetra and Entosphenus. An important synonym is Petromyzon japonicus von Martens, 1868, described from Japan. It has been suggested that this species consisted of three subspecies, japonica, septentrionalis, and kessleri. However, the subspecies septentrionalis is no longer recognized, and the subspecies kessleri is now considered a distinct species. Other synonyms include Petromyzon fluvialis Richardson, 1823, and Petromyzon borealis Girard, 1858 (both based on material from Great Slave Lake, Northwest Territories) and Ammocoetes aureus Bean, 1881, described from Anvik, Yukon River, Alaska. Some authors consider the species to contain both a parasitic and a non-parasitic form. Treatment of the species is restricted to the parasitic form, and the non-parasitic form is considered a distinct species (Lethenteron alaskense Vladykov and Kott, 1978, described from West Creek, a tributary of Brooks Lake, Alaska) until the question is settled. Additionally, it has been considered that there are two anadromous forms: a larger-sized, later-maturing form and a smaller-sized, earlier-maturing one, called praecox. The existence of these two forms requires further study, and they are treated together here. description:  The Arctic Lamprey is distinguished by the absence of paired fins, the seven gill openings on each side, the nostril on top of the head, and the mouth as a sucking disc with teeth. This species has two dorsal fins that are well separated in immature adults and touching at the base in mature adults. The second dorsal fin merges with the spade-shaped caudal fin. Oral disc length, as a percentage of branchial length in adults of 13.0–46.0 cm in total length, is 45%–76%. At the periphery of the oral disc are 12–22 oral papillae, and internal to these are 87–112 oral fimbriae. Dentition of the oral disc includes a single row of marginal teeth; a broad supraoral plate (lamina) with a unicuspid (rarely bicuspid) tooth on either end; typically three bicuspid endolateral teeth

Lethenteron camtschaticum, oral disc

Lethenteron camtschaticum



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on each side of the mouth; an infraoral plate with 5–10 teeth, the lateral-most ones usually bicuspid, the intermediate ones unicuspid; three rows of anterial teeth; no exolateral teeth; a single row of 12–28 posterial teeth; a transverse lingual plate that is scoop-shaped (U-shaped) and bears 13–18 unicuspid teeth, the median one greatly enlarged; and longitudinal lingual laminae with 10–14 teeth each. Trunk myomeres number 65–77, with strong modes at 68–71. Velar tentacles number 5–7 (usually 7), and one on each side is folded back into a wing. The dorsal surface of adults in fresh water is dark brown to blue black in colour, and the ventral surface is pale. In marine waters the dorsal surface of adults is steel blue, the ventral surface is pale to silvery, there is a black blotch at the apex of the second dorsal fin, and the caudal fin is usually weakly to moderately pigmented. Larvae are brown to grey dorsally, paler ventrally, and can reach 22.0 cm in total length. Adults can vary between 11.0 cm and 62.5 cm in total length, but are usually 30.0–45.0 cm, and attain a weight of 237.0 g.

habitat:  Some populations are anadromous (adults feed in the

coastal marine environment down to a depth of 50 m and spawn in fresh waters), and others are permanent freshwater residents (upper Mackenzie, Slave, and Hay Rivers, Great Slave and Artillery Lakes, Northwest Territories). The larvae live in the soft silty substrate of streams, rivers, and lakes where the current is slight. Spawning occurs on a pebble-sand substrate in rivers and streams at water temperatures of 12.2°C–15.0°C and currents of 0.2–0.3 m/s.

biology:  The larvae (called ammocoetes) filter feed on microscopic algae and detritus. As an adult the species feeds by removing large chunks of flesh from a variety of fishes including Burbot, Cisco, Inconnu, Lake Trout, Lake Whitefish, Longnose Sucker, Pygmy Whitefish, Rainbow Trout, Sockeye Salmon, and Threespine Stickleback in fresh water and Arctic Char, Arctic Cisco, Atlantic Herring, Atlantic Salmon, Broad Whitefish, Chinook Salmon, Chum Salmon, Dolly Varden, Inconnu, Least Cisco, Pacific Herring, Pink Salmon, Rainbow Smelt, Saffron Cod, and Starry Flounder in the sea. It is fed upon in fresh water by Burbot, Inconnu, Northern Pike, and Walleye, as well as gulls and American White Pelican. Larvae are believed to live four years and adults between one and three years, giving a total lifespan of 5–7 years. Age classes of ammocoetes attain the following total lengths: 35 mm in the first year; 30–65 mm in the second year; 60–155 mm in the third year; 150–220 mm in the fourth year. Metamorphosis into the adult stage occurs in August and September at total lengths of 140 mm to more than 184 mm. Downstream (feeding) and upstream (spawning) migrations vary from 200 km to 2,100 km. Downstream migrations either to the sea (anadromous populations) or to Great Slave Lake (freshwater-resident populations) occur from May to November. In the Yukon River, Alaska, upstream migration begins in November under ice. Upstream migrants are believed to travel as much as 27 km/day. Spawning occurs from late May to early July. Nests made of gravel are constructed by both sexes and are 15–25 cm in diameter and up to 7.5 cm deep at water depths of 8–20 cm. The male:female ratio is about 1:1 on the spawning grounds, and a female may spawn with more than one male. Fecundity ranges from 63,000 to 119,000 eggs per female in the anadromous form and from 9,790 to 29,780 eggs per female in the freshwater-resident form (upper Great Slave Lake basin). Eggs are dark bluish, adhesive, and slightly ellipsoidal, measuring about 1.0 mm by 0.9 mm. The incubation period is about one month, and the emerging prolarva measures about 7 mm in total length. Internal parasites include acanthocephalans, cestodes, nematodes, protozoans, and trematodes. importance:  The Arctic Lamprey is consumed by humans in

Lethenteron camtschaticum, oral disc

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Japan and was consumed in Russia at least into the twentieth century. It was consumed by Indigenous people, and especially their dogs, in Alaska into the twentieth century. Recently there has been a renewed interest in starting a commercial fishery for upstream migrants in the Yukon River, Alaska, for the Asian market in the United States and abroad to South Korea, in addition to the traditional subsistence harvest. The 2003 quota was set at 20,000 kg. The

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taste has been compared to that of sardine because of a high lipid content that reaches 38% of the body weight. Indigenous Alaskans prepare them by either baking or pressure-cooking.

distribution:  In Canada this species occurs in the Beaufort

Sea during its marine phase. It is the most widely distributed of the Northern Hemisphere Lampreys and is found at the highest latitude (72° N in Alaska). It occurs on both sides of the North Pacific Ocean basin (Asia and North America) as well as in the Arctic Ocean basin (Europe, Asia, and North America).

Distribution of Lethenteron camtschaticum

sources:  Evermann & Goldsborough (1907a); Berg (1931); Wal-

ters (1955); Nikolskii (1956); Birman (1960); Berg (1962); McPhail & Lindsey (1970); Nursall & Buchwald (1972); Vladykov & Kott (1978); Hunter (1981); Holčík in Holčík (1986); Kottelat (1997); Sandstrom, Lemieux, & Reist (1997); Khidir & Renaud (2003); Renaud, Gill, & Potter (2009).



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Family Rhinochimaeridae Longnose Chimaeras, Chimères à long nez

James D. Reist

Rhinochimaerids are small- to medium-sized fish (to 1.4 m in length) comprising three genera with about eight species. There are three species in Canadian waters, two occurring in the Arctic. Members of this family are found in the Atlantic, Indian, and Pacific Oceans, typically in deeper areas (200–2,000 m) along continental and island slopes (i.e., bathybenthic). One species from the Harriotinae, Harriotta haeckeli Karrer 1972 (Haeckel’s Chimaera, chimère d’Haeckel), occurs in southern Greenlandic waters. It is a rare benthic fish (2,020 m) from Davis Strait and so possibly present in our area. This family, and the related Shortnose Chimaeras, have cartilaginous skeletons; naked skin in most adults (no scales, but the young have dermal denticles); two dorsal fins, the first being short and erectile with a venomous spine at the front, and the second being longer and lower than the first; pre-pelvic and head claspers in the male, used to grasp the female during copulation (which is aided by pelvic claspers); an inferior mouth with grinding tooth plates that may protrude (hence ratfishes or rabbitfishes as common names); internal fertilization and egg case production; four gills overlaid by a gill cover, so there is one gill opening on each side of the head as in bony fishes; separate anal and urogenital openings; no stomach, spiracle, or ribs; and no vertebrae. They are related to sharks and rays. Longnose Chimaeras are easily distinguished from related Shortnose Chimaeras by the possession of a long, fleshy pointed snout that is not hook-like. In addition, the claspers of the male are a single rod, and the axis of the caudal fin is slightly raised (i.e., appears slightly heterocercal) with the fin asymmetrical, the upper lobe being narrower than the lower lobe. The snout length varies among the species. The mouth has tooth plates of varying thickness, two pairs in the upper jaw and one pair below. Water is breathed in through the nostrils rather than the mouth. Within the family the caudal fin is variably distinct from or confluent with the anal fin. The pectoral fins are large and wing-like and situated immediately behind the head and far forward. Eyes are moderately large and prominent.

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Two subfamilies are recognized: Harriotinae with thick tooth plates, and Rhinochimaerinae with smooth, thin tooth plates. Each subfamily has a single species in the Canadian Arctic. All species in the family are poor swimmers, epibenthic on muddy bottoms in general ecology, and appear to consume mostly crustacean and shellfish prey, although they also likely consume carrion. Males possess a bulbous head clasper (tenaculum), which with the pre-pelvic tenacula is used to hold the female for copulation. The rod-like pelvic claspers aid internal fertilization. The family is oviparous, producing ovoid, pear-shaped, or spindle-shaped egg cases that have a ribbed lateral web. Only one egg capsule develops in each ovary. The long snout characteristic of this family has both chemical and electrical sensory organs, likely used to detect prey or mates. They have no economic importance in Canada, although some are undoubtedly captured as by-catch in deep-water fisheries and used as fish-meal.

sources:  Bigelow & Schroeder (1964); Karrer (1972); Compa-

gno, Stehmann, & Ebert (1990); González, Teruel, López, & Paz (2007); www.iucnredlist.org, accessed 26 August 2010; FishBase, www.fishbase.org.

Harriotta raleighana Goode and Bean, 1895

Longnose Chimaera, chimère-spatule

common names:  A local name is Smalsnudet Havmus (Danish/Greenlandic). Other common names are Bent-nose Chimaera, Bentnose Rabbitfish, Bigspine Spookfish, Narrow-nose Chimaera, Spear-nose Chimaera, and chimère à nez rigide. The common name Pacific Longnose Chimaera from FishBase is not used, because the species was described from the Atlantic Ocean.

taxonomy:  The genus comes from Thomas Harriott, an English naturalist in Raleigh’s Roanoke Virginia colony in 1585 who first published on American natural history in English. The species name refers to Sir Walter Raleigh, leader of the colony and by whom Thomas Harriott was sent to the New World. description:  The Longnose Chimaera is distinguished from its relative by the lack of denticles or knobs on the upper margin of the caudal fin and by the presence of ridged or knobbed dental plates. This species is a large-sized fish characterized by a long, flattened, and pointed fleshy snout. The body is soft and moderately elongate, with a smallish head that has a slightly arched profile. The body is compressed and tapers evenly towards the tail. The small, inferior mouth is located under the large eyes and has moderately fleshy lips. Tooth plates are broad, with blunt edges, and are ridged or knobbed. The spine preceding the first dorsal fin is long and straight with serrations. The pectoral fins are large and broadly rounded and extend to the origin of the pelvic fins. The pelvic fins are small with pointed or rounded tips and form slender rod-like claspers in males. The caudal fin is lanceolate, tapering to a long slender filament (often lost in captured specimens), and the anal fin is usually continuous with the caudal. Tubercles (denticles) occur on the snout in adults. The species is a brownish colour overall with lighter fins edged by brown. Although the pelvic fins are black, paler areas may be present on females. The iris is a pale green. The total length attained is 1.2 m. habitat:  Th  is deep-water species is likely associated with soft or

silty bottoms on continental slopes or abyssal areas at depths of primarily 360–2,603 m, although some individuals have been captured at both shallower and deeper depths (to 3,100 m). It is associated with cold waters (primarily 2.5°C–10.9°C), and higher salinities (34.1–35.6 psu). Similar to others in the group, Longnose Chimaeras are likely sluggish swimmers that glide on the large pectoral fins. Adults and juveniles likely occupy different habitats.

biology:  Similar to that of others in the family, the biology is

poorly known. The diet includes bottom invertebrates such as small

Harriotta raleighana

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shellfish, amphipods, and polychaetes, i.e., organisms usually associated as infauna in soft sediments. Squid beaks in one individual and some fish remains in others suggest active predation on epibenthic or pelagic organisms. Information on growth and age is virtually non-existent. Sexes are dimorphic in size (females larger) and in secondary sexual characters (males with head and pelvic claspers). Fertilization is internal, females are oviparous, and egg cases are black elongate cylinders with wide, thin, laterally ribbed flanges (16.5 cm long). Reproductive output is likely low. Young hatch in summer. Abundance is unknown; however, it is inferred to be relatively common due to wide-spread occurrence and frequent capture of this species in deep-water research trawls and some commercial fisheries.

importance:  The Longnose Chimaera is captured occasionally

as by-catch in deep-water trawl fisheries and used as fish-meal and, more frequently, in research surveys. It is likely to be an important epibenthic predator and scavenger in deep-water habitats. It has been globally assessed as of “Least Concern.”

distribution:  Th  is species is found in Davis Strait, and in southeast Greenland south to Nova Scotia and Chesapeake Bay. Occurrence is probably wider than indicated. It is wide spread and has been documented from most of the global oceans, although numbers in the western Pacific and north Atlantic are the highest.

Rhinochimaera atlantica Holt and Byrne, 1909

Knifenose Chimaera, chimère-couteau

common names: A local name is Spydnæset Havmus (Danish/ Greenlandic). Other common names include Longnose Chimaera, Spearnose Chimaera, Straightnose Rabbitfish, and chimère à nez mou. taxonomy:  The genus comes from the Greek rhin (snout) and chimaira (a fabulous monster). The species name comes from the Latin atlanticus (of the Atlantic). The name Harriotta raleighana is mis-applied in some early literature. description:  This species is distinguished from its relative the Longnose Chimaera by the smoothness of its tooth plates, versus knobbed or ridged ones, and the upper edge of the caudal fin is typically a series of 19–33 (rarely to 43) knobs or denticles along the margin, versus being smooth. The knobs are more prominent in males. The body tends to be soft and elongate, with a moderately large head that is distinguished by a long, fleshy snout tapering to a point. The snout is somewhat flattened in dorsal view. The snout and head have a network of lateral line canals that may aid in sensing food items. The profile of the head is flat, and the mouth is located anteriorly to the eyes and has prominent fleshy lips. The tooth plates are smooth, sharp-edged, and narrow. The dorsal spine is moderately long and smooth in adults; however, fine serrations along its length may be present in juveniles. The pectoral fins are long and narrow. The caudal fin lobes are asymmetrical, with the lower being large and more obvious. Juveniles have a long, terminal caudal filament, which becomes quite short in adults. The anal fin is absent. The overall colour is chocolate brown, although whitish overtones may be present, and the fins are darker in colour. The gill opening is dark. This species attains about 140.0 cm in total length. habitat:  This is primarily a benthic or epibenthic species found

at the bottom of continental slopes out to deeper waters (i.e., > 200 m to 1,500 m). Canadian Arctic captures have been at 915–1,098 m. Although the captures are relatively few and widely spaced, this is likely an artefact of sampling effort and difficulty in deep waters. Abundance and distribution are likely greater than presently documented. It has been captured in association with deep, cold-water coral reefs where it may be a significant component of the benthic ecosystem. It is likely a sluggish swimmer that uses the pectoral fins for gliding, with locomotion by gentle undulations of the body.

Distribution of Harriotta raleighana

sources:  Jordan & Evermann (1896–1900); Bigelow & Schroeder (1953b); Mauchline & Gordon (1983); WoRMS (2013); Encyclopaedia of Life (2013).



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biology:  The biology of this species, like most deep-water taxa,

is poorly known. The diet includes benthic crustaceans (e.g., crabs, shrimps, and possibly sea spiders) and perhaps deep-water corals. It also likely consumes dead fishes and other carrion from the sea floor but is unlikely to actively pursue live fishes. Information on growth

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Rhinochimaera atlantica

is poor, but total lengths recorded in Canadian waters reach 1.2 m, and specimens of up to 1.4 m in total length have been recorded from elsewhere. Secondary sexual characteristics are well developed at maturity, with males possessing pelvic and head claspers and having more prominent caudal fin nodules. Fertilization is internal, with the males using the claspers to hold the females during copulation. The females are oviparous and produce few (perhaps as low as two – one per ovary), large (to 15 cm long), tadpole-shaped egg cases that have narrow anteriors, elongate posteriors, and wide lateral membranes with obvious cross-ribbing. Egg cases are likely laid on the substrate. Reproductive potential is likely low, and so susceptibility to impacts (e.g., by-catch in fisheries) may be high.

importance:  This species may be moderately abundant along

continental slopes and in deeper areas, although no quantitative survey data are available. Captured primarily as by-catch in deepwater trawl and longline fisheries, it is most likely discarded. It is possibly an important predator in deep-water benthic ecosystems. This species is globally assessed as “Least Concern” due to habitat occupancy beyond the depth limit of most commercial fishing and a suspected wide distribution.

distribution:  Th  is species has been found in Davis Strait (62°22' N, 60°39' W) as a single record from cruise data, and in southeast Greenland, south to the Gulf of Mexico, in the western Atlantic. This species may occur further north in Canadian (and perhaps western Greenland) waters than indicated on the distribution map. Originally described from the eastern North Atlantic near Ireland, this species appears to have a wide-spread distribution in suitable habitats (colder, deep waters) throughout the North Atlantic, including Iceland, northeastern Atlantic waters off the British Isles and Faroe Islands, as well as amphitropically in colder South Atlantic waters. It does not appear to range northwards to Nordic waters (but this may be an artefact of sampling effort). It has been reported off the coast of tropical West Africa, but these identifications are suspect, and from southwestern African waters off Namibia.

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Distribution of Rhinochimaera atlantica

sources:  Bigelow & Schroeder (1953b); Carpenter (1969); Compagno et al. (1990); Dagit & Compagno (2006); Hall-Spencer, Allain, & Fossa (2002); Møller, Kullberg, & Jørgensen (2004).

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internal fertilization. Two egg capsules (one per ovary) each contain one embryo, develop (oviparous), and are deposited on the substrate. Once laid, embryos take 9–12 months to develop. Head pores in the open lateral line grooves contain electro-sensory organs analogous to those of sharks. They are generally caught incidentally during other fisheries and have no or very limited economic importance in Canada. Directed fisheries do occur in other areas; for example, Iceland fisheries took approximately 500 t of Chimaera monstrosa, a northeastern Atlantic species, in 1991, but catches declined to about 10 t in 1997. They have been used for fish-meal, and the liver produces a fine-quality, machine-grade oil.

Family Chimaeridae Shortnose Chimaeras, Chimères

James D. Reist

source:  Bigelow & Schroeder (1953b).

Shortnose Chimaeras or Ratfishes are small to medium-sized fish (to 1.5 m) comprising about 22 species in the Atlantic, Indian, and Pacific oceans (two in Canada, of which one is in Arctic waters). One additional species, Hydrolagus pallidus Hardy and Stehmann, 1990 (Small-eyed Rabbitfish or Ghost Shark), is recorded at 1,336 m depth from nearby southwestern Greenlandic waters. The Latin root of the family name refers to the odd appearance that appears to encompass parts of several animals and was called a chimaera (monster) in mythology; this monster had a lion’s head, a goat’s body, and a serpent’s tail. General features of this family and the related Longnose Chimaeras are given under the latter’s family account. The snout is rounded and short; the mouth has chisel-like tooth plates; the tail is internally and externally symmetrical (diphycercal); the pre-pelvic tenacula are paired; toothed cartilaginous clasping structures are enclosed in dermal pockets that are partially covered by a flap of skin; egg cases are tadpole shaped and may have a filament at one end, but with little or no lateral web; the poison gland at the base of the dorsal spine produces a painful venom; water is breathed in through the nostrils rather than the mouth; the lateral line is an open or closed groove (the latter has pores at intervals); the eyes are large and prominent; the caudal peduncle is tapering, with a “leaflike” or filamentous caudal fin; the anal fin is present or absent and may be confluent with the tail (Hydrolagus), or separate (Chimaera, not in Arctic Canada); and the pectoral fins are large, wing-like, and situated far forward. Shortnose Chimaeras are benthic in habit and swim poorly. They occur from coastal areas (rarely) to the deeper waters of the continental margins along slopes and at relatively great depths (bathybenthic); however, about 3,000 m is the postulated limit. Food includes invertebrates and small fishes. The ecological importance is unknown but could be significant in mid- to deep-water benthic and epibenthic habitats. Similar to rhinochimaerids, males use the head clasper and pre-pelvic claspers to hold the female during copulation. The bi- or tri-furcate pelvic claspers facilitate



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Hydrolagus affinis

(de Brito Capello, 1868) Deepwater Chimaera, chimère de profondeur

common names: A local name is Småøjet Havmus (Danish/

Greenlandic). Other common names include Small-eyed Rabbitfish and Ratfish (and variants) and chimère à petits.

taxonomy:  The genus comes from the Greek hydro (water) and lagus (hare or rabbit; hence the variants of the English common names) in reference to the lips and teeth. The species name comes from the Latin affinis (related to) in reference to another chimaerid, Chimaera monstrosa. It was originally described as Chimaera affinis from deep water off Portugal. Chimaera plumbea Gill, 1878, described from southeast of LeHave Bank (42°40' N, 63°23' W), and C. abbreviata Gill, 1883, described from the northwestern Atlantic (40°16'50" N, 66°58' W), are synonyms. The Deepwater Chimaera has been suggested by some to be con-specific with South African and/or Japanese taxa, but it likely represents a valid taxon confined to the North Atlantic. description:  This species is distinguished from the related Longnose Chimaeras by having a snout that is short and rounded (versus elongate and pointed), and claspers that are bifid or trifid (versus a single rod). The soft body is elongate, the head is large, and the trunk is deepest anteriorly just behind the gill opening, tapering to a pointed tail; the head is short with a blunt, short, conical snout. There is a small inferior mouth, and the eyes are large and oval. The tall first dorsal fin has a free outer edge preceded by a shorter stout spine, and the long second dorsal fin is without a notch. The small lanceolate caudal fin has a short filament (longer in juveniles), and its lower lobe continues forward as a fleshy ridge. The anal fin is small and indistinct, not separated from the caudal, and considered absent

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Hydrolagus affinis

by some. The abdominal pelvic fins have pointed tips. The pectoral fins are large and wing-like with pointed tips, are located ventrally below the first dorsal fin, and reach the origin of the pelvic fins when depressed, but do not overlap them. There are no scales, but the smooth skin is pockmarked with rounded depressions. There is a prominent lateral line with several branches on the head. The trunk is lead coloured or in shades of brown above (including purplish overtones) and is generally lighter or white below; whitish spots may be present dorsally. The snout is grey. Fins tend to be lighter with dark margins especially along the posterior edges. Tooth plates are greyish, and the tongue is purple. Eyes are reflective as a shiny blue. The Deepwater Chimaera reaches 147.0 cm in total length and over 8.0 kg in weight.

habitat:  This is primarily an epibenthic species, occupying

waters from coastal margins (rare) between depths of 300 m to 2,900–3,000 m, although it may also occur off the bottom benthopelagically. Canadian Arctic records are from 1,271–1,472 m. It appears to be rare to moderately common on continental slopes down to deep-sea plains. It is a sluggish swimmer, using the pectoral fins for locomotion with gentle undulations of the body. Recent studies associate this species with deep, cold-water coral reefs, a poorly studied deep-sea benthic ecosystem.

biology:  The biology of this species is generally very poorly

known. The reported diet consists of benthic and epibenthic invertebrates including sea pens, hermit crabs, and fishes, which implies some degree of active hunting behaviour. It is possibly nocturnal, given the large eyes, although this may be an adaptation to depths. Information on growth is generally poor and lacking in Canadian waters; the largest male is reported to have been 84 cm in total length, and the largest female 103 cm in total length – smaller than elsewhere. It likely does not mature sexually until at least 60 cm body length. Reproduction involves internal fertilization of the

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female. Females are oviparous, with the embryo being enclosed in a brown, horny, tadpole-shaped capsule (about 20 cm long in related species), which is likely laid upon the substrate (egg capsules have been observed to be set upright intertidally in the related Hydrolagus colliei or Spotted Ratfish of Canadian Pacific waters). Only one egg capsule develops in each ovary; at most two are laid at one time. Development takes about a year in H. colliei. This low reproductive potential, and possibly low survival of the developing embryo, presumably results in increased susceptibility to impacts especially exploitation as by-catch in longline or trawl fisheries.

importance:  This species was (and perhaps still is) plentiful

along the slopes of banks fronting Nova Scotia as it was frequently reported from longline catches below about 600 m in around 1875. Once this fishery was discontinued, reports of the species decreased. Present abundance is unknown. As noted in the family account, species in the past have been used commercially as fish-meal and also to produce high-quality machine oil from the liver. No present fisheries target the species, although incidental catch likely occurs in trawl and/or longline fisheries. It is globally assessed as “Least Concern” due to wide-spread geographic occurrence especially at depths beyond most commercial fishing areas.

distribution:  The Deepwater Chimaera is found in Davis and Hudson Straits, based on cruise data and ARC 8602861, ZMB 22680, and ZMB 23704, and is common and spawning in southwest and southeast Greenland waters. The species likely is more wide spread in our area than is indicated on the distribution map. This northern Atlantic species is present from Iceland to mid-latitudes (about 37° N) but may also extend further south (e.g., to about 15° N at Cape Verde Islands) in the eastern Atlantic. In the western Atlantic it occurs from Greenland and Canadian Arctic waters (about 65° N) south to the Cape Cod area (about 41° N).

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Family Scyliorhinidae Cat Sharks, Roussettes

Brian W. Coad

Distribution of Hydrolagus affinis

sources:  Jordan & Evermann (1896–1900); Bigelow & Schroeder (1953b); Hardy & Stehmann (1990); Gunnarsson, Jonsson, & Palsson (1998); Møller, Kullberg, & Jørgensen (2004); Priede et al. (2006); Møller, Nielsen, Knudsen, et al. (2010); Priede & Froese (2013).

The Cat Sharks make up the largest family of sharks, with over 113 species and more than 25 undescribed. They are found worldwide in polar to tropical waters, with eight species in Canada, one of which is in the eastern Arctic. These are small sharks, seldom exceeding one meter in total length. The first dorsal fin base lies opposite or behind the level of the pelvic fin base rather than in front as in other sharks. Nictitating eyelids are weakly developed; the eyes are cat-like; spiracles are present and well developed; the last pair of gill slits lies over the pectoral fin base; fin spines and precaudal pits are absent; the lower caudal fin lobe is weakly developed; the intestine has a spiral valve with 5–22 turns; and several rows of very small teeth are functional at once. Egg cases are broader at the posterior end. These sharks are found from intertidal waters down to over 2,000 m, usually on the bottom. They are relatively poor swimmers and do not migrate significant distances. Individuals of some species sleep together in rocky crevices during the day and emerge at night to feed separately. Tough egg cases may be produced that are attached to the substrate for almost a year as the embryo develops. Others hatch in less than a month after a long internal development in the mother, a strategy to reduce the chance of being eaten. Yet other species of this family are ovoviviparous. Food items are mainly invertebrates and small fishes. Deep-water Cat Sharks are black overall, but those in shallow waters may have complex patterns on their body. Some species are used as human food, as fish-meal, for oil, and in aquaria, but Canadian and Arctic species are too rare or too deep to be commercially important.

sources:  Springer (1979); Compagno (1984).



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Apristurus profundorum (Goode and Bean, 1896)

Deepsea Cat Shark, roussette de profondeur

habitat:  This species lives on or near the bottom, generally at

1,300–1,830 m, some as shallow as 256 m. Specimens in Davis Strait have been caught at 1,149–1,417 m.

biology:  Unknown.

common names: Other common names are Deep-water Catshark and holbiche papoila.

importance:  This species is only caught in deep exploratory

taxonomy:  The genus comes from the Greek a (without) plus

distribution:  The Deepsea Cat Shark is found in the northern Atlantic Ocean from Davis Strait southwards in the western part, with some possible records in the southern Atlantic.

pristis (saw) and oura (tail), referring to the supposed absence of modified saw-like scales on the upper edge of the caudal fin. The species name comes from the Latin profundum (depth) and -or (that which), meaning of the depths, the type being caught at 816 fathoms. This genus of Cat Sharks is poorly understood taxonomically. The correct names for species in the Davis Strait and the northwestern Atlantic Ocean are disputable. Fishes caught off Newfoundland and in Davis Strait are often referred to A. profundorum, and a Davis Strait specimen was identified as this species by DNA bar-coding. Other names used for these fishes are A. laurussonii (Saemundsson, 1922) and A. manis (Springer, 1979), although the Davis Strait specimen was distinguished from A. manis in the bar-coding.

description:  The presence of an anal fin is distinctive among Arctic sharks, and the caudal fin has a crest of enlarged denticles. The body is slender, not tapering strongly to the head. The snout is longer than the mouth is wide. There are no crests over the eyes. Teeth are tricuspid with smooth edges. Nostril openings are elongate ovals. Flank denticles are nearly overlapping and erect, giving the skin a fuzzy texture. The first dorsal fin is nearly the size (twothirds) or the same size as the second dorsal fin. The dorsal fin origin is usually about opposite the pelvic fin mid-bases. The space between the dorsal fins is longer than the first dorsal fin base. The spiral valve count is 8–12. The labial furrows are very long, and the upper labial furrow is equal to or shorter than the lower. The overall colour is grey brown to dark brown or black. The Deepsea Cat Shark attains at least 62.5 cm in total length.

trawls and has no commercial importance.

Distribution of Apristurus profundorum

sources:  Jørgensen, Hvingel, Møller, & Treble (2005); Ebert, Fowler, & Compagno (2013).

Apristurus profundorum

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Family Etmopteridae

sources:  Jordan & Evermann (1896–1900); Nielsen, Bertelsen, & Nystrøm (1992); Compagno (1984, 2002); Compagno, Dando, & Fowler (2005); Priede et al. (2006); www.iucnredlist.org, accessed 2 September 2010; FishBase, www.fishbase.org.

Lantern Sharks, Requins-lanternes

James D. Reist Centroscyllium fabricii (Reinhardt, 1825)

Black Dogfish, aiguillat noir

common names: Local names are Eqalussuaq Qernertoq and Kukilik (Greenlandic). taxonomy:  The genus comes from the Greek kentron (spine),

Lantern Sharks have previously been considered as a subfamily with several others within the Family Dalatiidae in the wide sense. In the strict sense used here and followed recently by specialists, the Lantern Sharks are a family of about 50 species in 5 genera found in most oceans, typically in deeper waters. There are three species in Canada, one of which is in the Arctic. Lantern Sharks are characterized by their dwarf to small size (10– 107 cm in total length); cylindrical or slightly compressed bodies; photophores that are either inconspicuous or forming black marks along the belly, lower sides, and tail; and two dorsal fins with strong grooved spines preceding them, the second dorsal fin being usually larger than the first, and both having straight or concave margins. The anal fin is absent; there are no lateral keels on the caudal peduncle; and the caudal fin has a subterminal notch. The tooth morphology is fairly diagnostic among genera. In the Canadian Arctic species, for example, the upper teeth have strong cusps, and the lower teeth are laterally widened, blade-like, and larger than the upper, and cusplets are present on teeth in both jaws. Species are usually dark coloured with conspicuous black markings and luminescent organs on the lower surface (abdomen, caudal peduncle, pelvic and anal fin bases). The family name comes from the Greek etmo (cut) and pterus (fin), in reference to the original type described by Rafinesque in 1810 that had frayed fins. Most species are bottom dwelling, often at great depths off continental shelves (e.g., 200–2,200+ m). Like almost all demersal chondrichthyans, occurrences deeper than 3,000 m are rare. They feed on small fishes, cephalopods, crustaceans, and other benthos. Reproduction is ovoviparous, with 6–20 pups in a litter. In general, as a result of their abyssal habit, these sharks are poorly known, many are endemic and geographically restricted in distribution, and new species are regularly described. Most specimens are in poor condition as a result of capture in fishing gear, and great care is needed to ensure correct identification. Although poorly known, most species are likely common where they occur, and many are likely taken as by-catch in various fisheries conducted at depth.



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referring to the dorsal fin spines, and skyllo (to rend or tear to pieces) and skylion (dogfish or small shark). The species is named for Otto Fabricius (1744–1822), a Danish naturalist and missionary who studied the fishes of Greenland and authored Fauna Groenlandica. The species was originally described from Greenland waters as Spinax fabricii.

description:  This species is distinguished from other Arctic sharks in Canada by the lack of an anal fin and the presence of dorsal fins that each have a stout spine at the origin, and of teeth in both jaws with central cusp and lateral cusplets. It is also separated from other genera in the family by these tooth characteristics, by the first dorsal spine being well behind the tips of the pectorals when the latter are laid back, and by the space between the second dorsal and the caudal fins being about the same length as the eye to the first gill opening (although keys to the species of this genus are provisional at this time). Black Dogfish bear a superficial resemblance to a northern species, the Rough Sagre or Great Lanternshark (sagre rude; Etmopterus princeps Collett, 1904), which is not reported from our area but is present in southeast Greenland and is the only likely co-occurring similar shark. However, the Black Dogfish can be distinguished in that the Great Lanternshark has upper and lower teeth that are distinctly different, with the lower teeth being unicuspid. Although the Black Dogfish also superficially resembles the Spiny Dogfish (aiguillat commun; Squalus acanthias Linnaeus, 1758; Squalidae), which also has not yet been reported from our area (though it too occurs rarely in southwest and northwest Greenland waters), it is easily distinguished by tooth characters (a continuous cutting edge, unlike the cusps of the Spiny Dogfish); it lacks the lateral grooves that are on the dorsal spines of the Spiny Dogfish; and it has a notched caudal fin with a large dorsal lobe and a small ventral lobe. The body is short but robust, compressed, with a thick, fleshy, moderately long snout, long stout abdomen, and short caudal peduncle; lateral keels along the caudal peduncle are absent, and the head is long, with the mouth originating below the eye, and

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Centroscyllium fabricii

moderately arched. The first dorsal fin originates far forward and is low, the second dorsal fin originates over or just behind the pelvic fins and is larger than the first, each fin having a white spine that possess lateral grooves and extends about half the height of the corresponding fin. Each eye is large and oval with a spiracle immediately behind. Upper and lower teeth each have 3–5 cusps and are very similar, the middle cusp being the longest. The pectoral fins are small, with square tips, and originate behind the last (fifth) gill slit, which is small. The pelvic fins are about the same size as the second dorsal fin, with pointed tips. The skin has widely spaced, thorn-shaped denticles and is mostly bare except on the trunk. This species is sexually dimorphic, with males being smaller than females and possessing pelvic claspers used for internal fertilization. The colour of adults is uniformly dark brown to black, with no conspicuous black markings on the ventral surfaces or sides of the tail. Juveniles have white fin margins. As with most other members of the family, there are luminescent organs in the skin that are seen as epidermal thickenings or dots. The species is usually 60–90 cm in length, although the maximum recorded size is 107.0 cm in total length.

habitat:  This species is found in deep water and is primarily benthic, usually from 180 m to 2,250 m on outer edges of continental shelves and slopes. It is reported at 575–614 m in Davis Strait, and in Canadian waters off Baffin Island at 619–1,404 m. An Ungava Bay record is from 187 m. In northern parts of the range it is reported to occur near the surface especially during colder, or perhaps darker, times of the year. These fish have been caught through the ice at the surface and may approach shallow waters in the Arctic. They are typically captured in water temperatures of 3.5ºC–4.6ºC, although reported too from 1ºC waters. Some evidence (e.g., lack of capture in 2.5ºC water versus numerous captures from 3.1ºC water on the slopes of Davis Strait Ridge) suggests that this is primarily a northern Atlantic cool-water species, rather than an Arctic taxon. This is partially verified in that recent winter longline fisheries in cold waters of Cumberland Sound did not capture this species as by-catch, although trawls in warmer waters in southern Davis Strait did. It is reported to be a schooling species although segregation by sex and

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age (size) appears to be typical. Such schooling may account for some of the high catches reported occasionally.

biology:  Its biology is poorly known generally. The diet consists

of pelagic and/or benthopelagic invertebrates including cephalopods, crustaceans, and jellyfish, as well as smaller benthic fish of several species including redfishes. Fish increase in importance in the diet as the sharks grow larger. The role of carrion in the diet of this species is unknown but could be significant, especially in association with discards of offal or by-catch from trawlers. Ages and longevity are unknown for this species; however, unverified estimates for other genera in the family range to 60 years. The growth rate is poorly known, but sexually mature females have been reported at 58–70 cm total length in the southern portions of the Canadian range. Half of the males from Iceland were sexually mature at 28 cm total length, whereas half the females were mature at 39 cm, with 100% maturity shown at 64 cm and 70 cm, respectively. Growth is likely slow and, combined with late age of maturity and a low reproductive rate, suggests that the species may be susceptible to anthropogenic impacts. In Icelandic waters both sexes show similar length-weight relationships, with an individual of the mean length of 60 cm averaging 1 kg in weight. The few males captured were 80 cm in length and weighed about 2.5 kg, whereas females at this size were 3 kg. The largest reported Canadian individual was 84 cm in total length. Habitat segregation by sex and size appears to occur at least in some areas, with males being more numerous in shallower Icelandic waters (< 1,000 m depth) and females more numerous in deeper areas. However, the reverse trend is reported for west Greenlandic waters: small immature individuals have been found at 500–800 m depth, and larger individuals at more than 800 m, with females being more numerous in deeper waters. Individuals found in deeper waters also tend to be shorter, and so are likely younger than those in shallower waters. Reproduction is ovoviviparous, with a few fertilized eggs developing within the female (e.g., young to 17 cm in length have been found in some females captured from west Greenland waters in February). The litter size averages about 16 (range 4–40). The smallest free-living individuals observed were about 165–175 mm in total

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length, indicating that birth occurs around a size of 175 mm in west Greenland waters. Mature, seasonally reproductive individuals were observed in both summer and fall; thus, annual reproduction appears to not be seasonally defined.

importance:  This species is of minimal importance to Canadian

fishers at present although it is likely one of the more abundant shark species in deep water, especially along the shelf drop-off. There are occasional reports of relatively large catches in benthic trawls, especially from scientific work. It is likely taken regularly in baited longline and bottom-trawl fisheries in suitable areas in the south, although most are likely discarded as by-catch. Observer survey data from foreign fleets fishing in NAFO areas outside the Canadian EEZ indicate that this species is one of the few sharks retained from the by-catch of fisheries for Greenland Halibut. In other areas, it is simply discarded. Biomass indices estimate substantial standing crops, for example, up to 49,000 t in some areas. If retained, they are likely used as fish-meal. The ecological role or importance is unknown but, given the apparent abundance, could be quite significant in benthic trophic webs. The frequency in tows and catch per tow has increased in Canadian Atlantic waters from the 1970s to the mid-1990s. This species has been globally assessed as of “Least Concern” due to wide depth distribution, and therefore its presence being outside the areas of significant fishing pressure, and because population trends in the northwest Atlantic appear stable. Significant by-catch from fisheries in the northeast Atlantic throughout much of the geographic and depth range indicates a regional assessment of “Near Threatened.”

distribution:   The Black Dogfish is found off Baffin Island in Davis Strait and south to Labrador, Newfoundland, Scotian Shelf, and Georges Bank south to perhaps Florida (although the extreme southern records require verification); it is especially abundant in the Laurentian Channel of the Gulf of St Lawrence. A record from eastern Ungava Bay is based on cruise data and was captured at 187 m, the shallower end of the species depth range, and needs verification. It is also present in southwest and southeast Greenland, and the northwards distribution in eastern Davis Strait along the western coast of Greenland appears to be further north than that observed across Davis Strait in Canadian waters. This is likely due to the warmer nature of the waters along the Greenland side of Davis Strait and Baffin Bay. Globally the species is confined to the Atlantic Ocean between about 66º N and 40º S, from Iceland and northern Europe south to western Africa in the eastern areas (although tropical records are uncertain).



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Distribution of Centroscyllium fabricii

sources:  Jordan & Evermann (1896–1900); Bigelow & Sch­roeder (1948); Jensen (1948); Templeman (1963); Compagno (1984); Nielsen, Bertelsen, & Nystrøm (1992); Yano (1995); S.K. Brown, Mahon, et al. (1996); Gascon (1996); Jakobsdottir (1998, 2001); Mahon et al. (1998); Castro, Woodley, & Brudek (1999); Musick (1999); Compagno et al. (2005); Ebert et al. (2013).

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Family Somniosidae Sleeper Sharks, Somniosidés

James D. Reist

Sleeper Sharks were previously considered to be a subfamily with several others, including the Etmopteridae, within the Family Dalatiidae in the wide sense. In the strict sense used here, and followed recently by specialists, the Sleeper Sharks are a distinct family of about 19 species in 7 genera. Representatives are found globally in most oceans including the Arctic and Antarctic Oceans; thus, this group appears to contain the only truly polar shark species. There are three species in Canada, including two to three species in the Canadian Arctic. Sleeper Sharks are characterized by their small to very large size (40 cm to > 7 m in total length); a cylindrical to slightly compressed body; a broad head with a flat snout; a short transverse mouth, with small needle-like teeth in the upper jaw and larger, blade-like teeth in the lower jaw; a large spiracle immediately behind each eye; lateral abdominal ridges; a caudal peduncle, also with lateral ridges in Somniosus; pectoral fins that are low, angular, or rounded, with short, rounded rear tips; pelvic fins that are equal to or smaller than the dorsal fins; two small dorsal fins, with the origin of the first being in front of the pelvic fins, distance between the two being larger than the fin base, and the second dorsal fin being smaller or equal to the first; no anal fin; spines that are present in some but absent in Somniosus; a heterocercal caudal fin with a strong subterminal notch; and no photophores. Species of this family tend to be benthic at great depths off slopes (i.e., > 200 m to 3,675 m). The exception appears to be Somniosus spp., which in northern latitudes occur on continental shelves, enter intertidal areas, and may occur at the surface. However, they are found at depth in southerly non-Arctic parts of the range, presumably avoiding warmer surface waters. Similar to that of most sharks, distribution below 3,000 m is likely limited. Aggregations appear not to be formed. All appear to be somewhat sluggish as scavengers or active predators and consume diverse prey including fishes, scavenged carrion of seals and whales, and invertebrates. They are important components of deep-water benthic communities. Reproduction is ovoviparous, with 4–59 pups per litter. None are known to attack humans, although anecdotal reports exist of

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Greenland Sharks stalking humans on ice, and they are attracted by blood in the water. These fish are generally caught as by-catch in fisheries using bottom trawls or long lines, although directed fisheries for some do occur. Baited long lines appear particularly attractive, and some reports of very high by-catches are known (with a concomitant loss of the intended catch such as halibut). Some directed fisheries occur for some species in this family; for example, Icelandic fisheries reportedly took about 80 t of Greenland Shark annually in 1982–97. Uses of the latter include liver oil (squalene), dog food, and fishmeal, as well as human consumption (in dried form). However, the flesh, like that of most sharks, contains urea, which is toxic and must be removed by washing, cooking, and rinsing, or drying. Its status is generally unknown, but, as a result of suspected low reproduction and slow growth, the potential for over-exploitation is high (see also Family Rajidae).

sources:  Nielsen, Bertelsen, & Nystrøm (1992); Gunnarsson et al. (1998); Musick (1999); Carpenter (2002); Compagno (1984); Compagno et al. (2005); Priede et al. (2006).

Centroscymnus coelolepis

Barbosa du Bocage and de Brito Capello, 1864 Portuguese Shark, pailona

common names: A local name is Portugisisk Fløjlshaj (Danish/ Greenlandic). Another common name is Portuguese Dogfish. taxonomy:  The genus comes from the Greek kentron (sting or

spine) and skymnos (an ancient shark name meaning “pup or lion whelp”). The species name comes from the Greek koilos (hollow) and lepis (scale) in reference to the concave dermal denticles. The species was described from the eastern Atlantic near Portugal where there is a deep-water fishery, hence its English common name. Originally aligned with squalids (Dogfish Sharks), Centroscymnus is now one of seven genera within Family Somniosidae and thus is related to the Sleeper Sharks that it resembles. Some specimens captured in the eastern Atlantic in the late 1800s and early 1900s were also identified as Centrophorus coelolepis (a generic synonym in part), but the species-level taxonomy has remained stable. The species is one of two presently aligned with this genus, although many generic realignments of this and closely related species have occurred in these poorly known deep-water sharks. Scymnodon melas Bigelow, Schroeder, and Springer, 1953, described from off Georges Bank, 40°00' N, 68°52' W, in the northwestern Atlantic is a synonym.

description:  This species is distinguished from other Arctic sharks by the lack of an anal fin, the upper jaw teeth are lanceolate

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Centroscymnus coelolepis

and the lower jaw teeth have an oblique cusp, and each dorsal fin has a small spine at the origin (often concealed by skin). It is a medium-sized shark with a robust, rounded body, somewhat ventrally flattened, and with a small head. Two dorsal fins are present, the second being larger than the first and situated near to the tail. Each pectoral fin originates immediately behind the last gill slit. The eyes are large, oval, and laterally positioned. A reflective layer (tapetum lucidum) results in eyeshine and likely enhances vision at depth. Similar to those of other Sleeper Sharks, the relatively large pelvic fins are situated far to the rear. The short, broad tail has a notch separating the upper and lower lobes. The large mouth is slightly arched, with thin lips. The denticles are large and flat. The upper teeth are slender and lanceolate with a single central cusp. The lower teeth are markedly different, being short and broad with a single oblique cusp and overlapping bases, forming a continuous cutting edge. The upper-jaw teeth number 43–68, the lower-jaw teeth 29–42. The species is dark brown (adults) or bluish black (partly grown individuals), a coloration that superficially resembles a small Greenland Shark. It reaches 1.22 m in total length.

habitat:  All reports suggest that this is a reasonably common

deep-water demersal species of the continental slopes mostly below 400 m depths and down to abyssal areas to 3,675 m (among the deepest known records for any shark species). A Canadian Arctic specimen was recorded at 1,301 m. Occasionally it is captured in shallower waters (about 128 m). Water temperatures are 5°C–13°C. Some evidence suggests segregation of sexes and sizes by depth, pregnant females being in shallower areas and juveniles in deeper.

biology:  Visual acuity to bioluminescence, and studies of its

diet, suggest that this species is an active predator, consuming fishes including other sharks, cephalopods (especially squids and gastropods), and whale carcasses. It can take bites out of live prey. Age and longevity are unknown, although generation time is estimated to be about 20 years. Reproduction is by aplacental vivipary (ovoviviparity), with eggs retained internally but nourished from their yolk sacs until birth. Young are born live at about 23–30 cm length in



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Atlantic waters. Typical litter size is 12 but ranges from 1 to 29. Both sexes appear to mature at about the same size (90–100 cm for males, 85–115 cm for females), although variation occurs throughout the range. Recent genetic studies suggest wide dispersal in the eastern Atlantic (i.e., low levels of population differentiation), stable population sizes (i.e., reasonably high genetic variation within sites), and spatial segregation by size and maturity. Long-distance migrations through the reproductive cycle appear likely, and distinct nursery areas are suspected.

importance:  This species is frequently taken in trawl, longline-hook, and gill-net fisheries both as a targeted species and as by-catch. Its primary use is oil (squalene) produced from livers, which may be nearly 50% oil by weight (oil-rich livers aid in neutral buoyancy at depth). Flesh may also be used (fresh, salted) or processed into fish-meal. Harvests (e.g., 300–900 t annually landed in some fisheries), limited abundance, demersal habit, likely low productivity, and long generation time all suggest that this is a sensitive species. The global status assigned is “Near Threatened.” Size, habitat, and behaviour all indicate that this shark poses no threat to humans. distribution:  C  entroscymnus coelolepis has been found in Davis Strait as two records from cruise data (67.51° N, 59.22° W; and 61.971° N, 60.258° W), and also in northwest and southeast Greenland. Globally it is found in suitable habitats throughout both sides of the Atlantic from the Arctic to southern Africa, including the Mediterranean Sea in the east; it is present from northern Canada to northern South America in the western Atlantic, and in the Pacific in Japanese waters and off South Australia. Thus, the species appears to have an amphiboreal distribution possibly associated with waters of suitable temperatures.

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sidae; however, taxonomic rearrangements are occurring at both the family and the subfamily levels, and new species are being described. Five (possibly six) species occur in the genus worldwide, organized into two subgenera: large sized (Somniosus), with three species distributed anti-polarly; and small sized (Rhinoscymnus), with two species. Despite the uncertainty of the validity of these and various synonymizations, the three Somniosus species appear to be valid and distinct from each other. Their large size (> 4 m length), the absence of spines preceding both small dorsal fins, the first dorsal fin being posterior to the pectoral fin base, and the caudal fin having almost equally sized lobes distinguish this subgenus from others in the family. The short, rounded snout, heavy body, oblique cusps on teeth in the lower jaw, and hook-shaped denticles further differentiate this subgenus.

Distribution of Centroscymnus coelolepis

sources:  Jordan & Evermann (1896–1900); Bigelow & Schroeder (1953); Verissimo, McDowell, & Graves (2011); Ebert et al. (2013); WoRMS (2013).

Somniosus microcephalus (Bloch and Schneider, 1801)

Greenland Shark, laimargue atlantique

common names:  Local names are Ekaludjuaq, Eqalukjuaq,

Iqalujjuaq and Iqalukuak (Inuktitut); Eqalussuaq and Niialingaq (Greenlandic); and Ekalugssûp Piarâ for a young shark (Greenlandic). Also it is known as Skalugsuak in Inuit legend. Other common names are Grey Shark, Ground Shark, Gurry Shark, Large Sleeper, Sleeper Shark, and requin dormeur.

taxonomy:  The genus comes from the Latin somniosus (sleepy) as, based on its small fins, the shark was thought to be slow, and the species name comes from the Greek micros (small) and cephalus (head), presumably in reference to the short snout. It was originally described from the Arctic Ocean as Squalus microcephalus. Synonyms include Somniosus brevipinna Lesueur, 1818, described from off Marblehead, Massachusetts; Scymnus glacialis Faber, 1829, described from Iceland; and Scymnus micropterus Valenciennes, 1832, described from the Arctic Ocean. Somniosus is one of seven genera within the Family Somnio-

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description:  This species is distinguished from other Arctic sharks by the lack of both an anal fin and spines at the front of the dorsal fins; the upper-jaw teeth are lanceolate, and the lower-jaw teeth have an oblique cusp; the origin of the first dorsal fin is nearer to the tip of the snout (< 45% of the total length) than to the tip of the caudal fin; and the space between dorsal fins is equal to the length from the snout tip to the first gill opening. The Greenland Shark is a very large shark with a heavy cylindrical but variably shaped body and a small to moderate-sized head. The snout tends to be short, rounded, and thick. Lateral keels are present in the caudal area. The skin is rough with large, strong, hook-like denticles. The mouth is of moderate size. The upper teeth are slightly recurved, thorn-like, and taper to the tip. The lower teeth are rectangular, their breadth being about half their height, and the cusps are smooth and notched to fit together to form a saw-like continuous edge. The upper teeth number 48–52, the lower teeth 50–52. The eye is small and circular and positioned towards the front of the mouth and is often occluded by an ectoparasite. All fins are small. The first dorsal fin originates well posterior to the pectoral fin base. The second dorsal fin originates somewhat posterior to, or over, the pelvic fin insertion. The pectoral fins originate immediately behind the last gill slit and are small with rounded tips. The pelvic fins are small and well posterior. The caudal fin is asymmetrical and variable in profile, with a short dorsal lobe having a small but noticeable subterminal notch, and with a well-developed ventral lobe. The body colour is grey to blue grey, coffee brown to black, occasionally with darker bands, mottling, or smudges on the sides. Albinos have been reported from Greenland. This species attains perhaps 7.3 m and perhaps 1,020 kg, although most are less than 5.0 m and proportionately lighter. habitat:  Its habitat use varies from near shores and estuaries, and

shelves to upper slopes (at least to 2,647 m), and perhaps deeper in southern locales. It has been reported to 388 m in Hudson Strait, to 1067 m in Cumberland Sound, at 420.5–635.0 m and 0.5°C–1.2°C in Davis Strait, and also to 1,160 m in Davis Strait. The Greenland Shark prefers colder waters (−1.8°C to 12°C) and thus may be inshore in winter and offshore in summer. Limited by-catch statistics suggest greater occupancy of slope habitats. Depth association is also

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Somniosus microcephalus, with upper and lower teeth (bottom)

latitudinally dependent, the depths being shallower in the north and deeper in southerly locations. The species appears to be tolerant of a range of salinities, often being found in estuaries, fiords, and nearshore areas, especially during summer sea-ice melt. Although it is generally considered a benthic species, ultrasonic tracking suggests that more time is spent well above the bottom than was previously thought. It is one of three common elasmobranchs in NAFO Subareas 0 and 1 (with Skates Amblyraja hyperborea and A. radiata).

biology:  This species feeds as a scavenger upon carrion and offal or gurry (hence Gurry Shark) discarded from whaling and fishing operations, often coming inshore to do so. Similarly it feeds upon carcasses discarded in the water. One entire reindeer has been found in a Greenland Shark’s intestine. Although reported as “sluggish,” it appears also to be an active predator, feeding upon seals especially in ice-covered areas, whales caught in nets, many demersal fishes, jellyfishes, cephalopods, gastropods, crustaceans, and sea birds. It is able to remove large chunks of flesh from a whale carcass with slow and deliberate bites. This shark has been reported as stalking a human walking on pack ice and also divers in the Gulf of St Lawrence. At Mittimatalik (Pond Inlet) one of these sharks was found to have eaten another shark that had eaten a third. In Cumberland Sound this species feeds on and derives much of its energy from Greenland Halibut, and in Norway it relies heavily on pelagic teleosts and seals. Cumberland Sound sharks feed on invertebrates such as the whelk Buccinum cyaneum, the sea urchin Strongylocentrotus droebachiensis, the basket star Gorgonocephalus arcticus, squids and amphipods, the fishes Amblyraja hyperborea, Reinhardtius hippoglossoides, Myoxocephalus scorpius, Lycodes reticulatus, skate eggs and lumpfish, and Ringed Seals. Tracking studies indicate that individuals exhibit irregular “up-and-down” oscillatory patterns of behaviour, which are likely associated with active foraging (see also Pacific Sleeper Shark). Evidence exists for site fidelity over shorter time frames



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as well as extensive movements over several years (i.e., > 1,100 km over 16 years). Although it is a predator, limited results from stable isotope studies have suggested a somewhat lower trophic position than would be fulfilled by a top carnivore. Carbon stable isotopes suggest a greater degree of pelagic, rather than strictly benthic, feeding. Thus, omnivory across various trophic levels appears to be the norm for this species. Growth is poorly known; adults are usually 2.4–4.3 m in total length, with a confirmed maximum size of 6.4 m total length and unconfirmed of 7.3 m total length. Females are usually larger than males by about one meter. This species attains a weight of 775 kg, although likely can be heavier. It is likely long lived (e.g., some estimates suggest about 200 years or even greater) and may be locally abundant (at least in the past in Greenlandic waters). Ages are unreported, growth is likely very slow, maturity is likely late (e.g., about 260 cm for males but variable, possibly 400 cm for females, and at 100 years), and reproductive rate is low, so it may be highly susceptible to anthropogenic impacts. It reproduces by ovovivipary with up to 10 pups in a litter, those being 38 cm in total length. Female fecundity appears variable, but several hundred eggs have been found in some individuals, with some ova being large (i.e., to 6 cm in diameter). Greenland Sharks are host to an ectoparasite (Ommatokoita elongata, Copepoda, Lernaeopodidae), the large females of which attach to the cornea. These parasites appear to infect most individuals (e.g., 99% out of 1,505 examined), cause eye lesions, and possibly render individual sharks blind. However, well-developed olfaction likely aids feeding in the low-light (seasonally and at depth) habitats occupied; thus, impacts on the populations appear to be minimal. It has also been suggested that as these parasites are very obvious and possibly luminescent, they may act as lures to attract prey, but this is unproven. Arctic Char from shark stomachs were reported to be complete except for the tail, suggesting that the char were attracted by the parasite and taken head first by the shark.

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Somniosus microcephalus

importance:  At present this species is of minimal importance to

fisheries in Canadian waters. It is regularly captured as by-catch on baited long lines, deep-set gill-nets, and trawl fisheries especially for Greenland Halibut (Reinhardtius hippoglossoides). There have been experiments with Greenland Halibut long-line gear, investigating reduced gangion-breaking strength, in order to reduce the by-catch in Cumberland Sound. Shark flesh from the Cumberland Sound halibut fishery was too high in mercury for use as food. Fisheries for this species have been documented since the thirteenth (Norway) and fourteenth (Iceland) centuries. It can be caught through holes in ice-covered waters. These sharks were accidentally caught in nets set by missionaries that were meant for seals, and oil was sent to England from Killiniq (Port Burwell) in the early-twentieth century. Fisheries in Greenlandic waters have historically been fairly high (e.g., 32,000 fish annually to about 1935), primarily to render the livers for oil. Whether such levels are sustainable is unknown. Over 447 liters of oil have been taken from one shark. Diet information suggests that this species is a significant component of northern deep-water ecosystems as a predator and scavenger and may also have a substantive natural influence on marine mammals (especially seal) populations. Population dynamic parameters suggest sensitivity to overfishing. Thus, even low levels of by-catch may be having

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significant population effects. It is globally assessed as “Near Threatened” on this basis. The Greenland Shark is an important source of income and sustenance for Indigenous and northern communities and is part of Indigenous culture. Dried or boiled flesh was also used to feed dogs in the past. Fermented meat is considered a treat by devotees in Iceland. As with most sharks, urea storage in the flesh renders it toxic in the fresh state, although this appears to vary with the diet of the individual shark. At Mittimatalik (Pond Inlet) dogs ate fresh shark meat without effect, but in Greenland dogs became “shark drunk” (drowsy and giddy), and some died. Ravens and Fulmars scavenging shark carcasses have become too ill to fly and stumble around as if intoxicated. Inuit legend tells of an old woman washing her hair in urine, then drying it with a cloth; the urine-soaked cloth blew into the sea and became Skalugsuak, the first Greenland Shark – an obvious reference to the urea stored in the flesh. Other historical uses include rough skin as sandpaper, tanned skins for leather (e.g., boots), and teeth rows as knives and saws.

distribution:  This species is found in Davis and Hudson Straits; Ungava, Hudson, and James Bays; and inshore areas in the eastern Arctic Archipelago including northern Baffin Island inlets,

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Lancaster Sound, and Resolute Bay. Hudson and James Bay reports are vague or anecdotal. Distribution is likely much wider than indicated by point distributions, but knowledge is hampered by a lack of suitable effort, especially in ice-covered areas. Its westernmost known location is Resolute Bay. It has not yet been reported from the western Canadian Arctic (but may occur there in deeper offshore Atlantic water masses), or from the Arctic Ocean northwest of the archipelago (although this may represent a lack of effort; see also S. pacificus). Globally it occurs in the boreal Atlantic south from the Canadian Arctic and southwest Greenland to New England and Georgia through Atlantic Canada. It is found in the North Sea northwards to the Barents Sea and the Arctic Ocean in the east and eastwards to Novaya Zemlya, although its extent in the Arctic Ocean proper is unknown. The Greenland Shark occurs in more southerly areas at greater depths (e.g., Georgia, USA, at about 2,200 m; and Portugal, although this may be a different species).

Somniosus pacificus

Bigelow and Schroeder, 1944 Pacific Sleeper Shark, laimargue du Pacifique

common names: Another common name is laimargue dormeur. taxonomy:  The species name comes from the Latin pacificus (peaceful) and -icus (belonging to), after the Pacific Ocean. General family and subgeneric taxonomy is as given previously for S. microcephalus. Specimens from Alaskan waters were referred to as S. microcephalus in the 1800s and early 1900s; thus, northeast Pacific references for this species are mixed with Greenland Shark. description:  This species is distinguished from other Arctic sharks by the lack of an anal fin and of spines at the front of the dorsal fins; the upper-jaw teeth are lanceolate, and the lower-jaw teeth have an oblique cusp; the origin of the first dorsal fin is almost equidistant (> 45% of the total length) between the tip of the snout and the tip of the caudal fin (i.e., the first dorsal fin is nearer to the tail than to the snout); and the space between dorsal fins is about 70% of the length from the snout tip to the first gill opening. This species is very large in size with a heavy cylindrical body, a short rounded snout, and no dorsal fin spines, all characteristics shared with the Greenland Shark. Lateral keels are usually present in the caudal area. The caudal fin has a large lower lobe. The skin is rough with denticles having strong hook-like cusps. The mouth, teeth, fins, and colour are all similar to those of the Greenland Shark. The body and fins are uniform grey to slate green or blackish brown. The size is more than 6.0 m in total length, with unsubstantiated reported estimates of 7–8 m, but most appear to be 3.7–4.3 m. habitat:  Its habitat use appears to be very similar to that of

the Greenland Shark, from nearshore to upper slopes, and similarly variable depending upon the local circumstances. It has been reported from a tide pool. This species likely occurs to at least 2,000 m depth, with depth correlated with latitude (shallower in the north). It is common in lower shelf and slope waters.

Distribution of Somniosus microcephalus

sources:  Jordan & Evermann (1896–1900); Bernier (1910); Bigelow & Schroeder (1948); Jensen (1948); Beck & Mansfield (1969); Polar Gas (1977–8); Crawford (1990); Hudon (1990a); D.B. Stewart, Dunbar, & Bernier (1993); Borucinska, Benz, & Whiteley (1998); Anonymous (2000f); Fisk, Tittlemier, Pranschke, & Norstrom (2002); Skomal & Benz (2004); Yano, Stevens, & Compagno (2004, 2007); Kyne, Sherrill-Mix, & Burgess (2006); Department of Fisheries and Oceans (2007); Idrobo (2008); Benjamins, Kulka, & Lawson (2010); Idrobo & Berkes (2012); MacNeil et al. (2012); McMeans, Arts, & Fisk (2012); Munden, Grant, & Hedges (2012); Ebert et al. (2013); McMeans, Arts, & Lydersen (2013); Nielsen, Hedeholm, Simon, & Steffensen (2013).



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biology:  This shark feeds on the bottom and in the water column

on several species of fishes, many of which are fast swimming, and on cephalopods, larger invertebrates, and seals, with importance being seasonally dependent. The head and mouth cavity are long, and the mouth small, suggesting that it may feed by suction, pulling in prey with a sudden inrush of water to the mouth cavity. It scavenges offal and carcasses especially those of larger marine mammals, although active consumption also appears to occur (i.e., biting off chunks of live animals). Tagged individuals in the Gulf of Alaska have exhibited almost continuous vertical movements consisting of various patterns. Movements were below the photic zone during daylight and closer to the surface at night. Oscillatory patterns with depth, exhibited by most individuals, suggest a distinctive active

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Somniosus pacificus

foraging pattern that, coupled with ambush predation, appears to enhance the capture success of active prey. Depths occupied range from 2 m to 724 m, and temperatures range from 4.4°C to 11.8°C. It is likely long lived and locally abundant. Ages are unreported, but growth is likely slow, maturity late, and reproductive rate low, and therefore it is susceptible to anthropogenic impacts. It is likely ovoviviparous with up to 300 eggs. In northwestern Bering Sea samples, females ranged in total length from 88 to 190 cm and in weight from 6.2 to 59.0 kg, whereas males ranged from 84 to 242 cm and from 4.7 to 133.0 kg, all of which were immature. Maturity appears to occur at a total length of more than 2.5 m. A 4.23 m mature female from the Kuril Islands south of Kamchatka weighed an estimated 819 kg. Similarly to the Greenland Shark, copepod ectoparasitic infections on the eyes are common.

be expended to capture and properly identify large sharks in the western Arctic. Reports of this species in the southern hemisphere represent the distinct species S. antarcticus Whitley, 1939, the Southern Sleeper Shark.

?

importance:  This species appears to be of limited fishery

importance, with some older, small-scale commercial harvests having occurred in southern areas (presumably for oil rendered from the liver). Population dynamic properties suggest that it may be sensitive to overfishing. Diet and behavioural information suggests that this species is an important component of shelf and slope ecosystems as a predator and scavenger. It may take salmon and halibut from long lines. It is globally assessed as “Data Deficient” due to paucity of information.

distribution:  The easternmost confirmed location along the north slope of Alaska is Point Hope. However, it is expected to occur further east in Alaskan waters and likely also into the western Canadian Beaufort Sea. Although it is not confirmed from Canadian waters, an anecdotal report by an Inuvialuit fisher (2008) of a large shark near to Tuktoyaktuk was possibly referring to this species. It is present in North Pacific coastal waters from Taiwan, Japan, Hawaii, and Baja California northwards, throughout Alaskan and eastern Asian sub-Arctic waters. It is found north of the Bering Strait in Chukchi and western Beaufort Seas. Its westernmost extent in Chukotka is unknown. Given the presence of Atlantic water masses in the western Canadian Arctic below 200 m (i.e., west of Banks Island), there is the possibility that Greenland Sharks may also (or alternatively) occur in the western Arctic. Accordingly, effort should

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Distribution of Somniosus pacificus

sources:  Musick (1999); Orlov (1999); Benz, Hocking, Kowunna, Bullard, & George (2004); Glubokov (2004); Hulbert, Sigler, & Lunsford (2006); Sigler, Hulbert, et al. (2006); Ebert, Fowler, & Compagno (2013).

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Family Rajidae Skates, Raies

James D. Reist

Skates are a highly diverse family of marine cartilaginous fishes present in tropical to polar seas and occurring from shallow to deep waters. About 26 genera and about 260 species likely exist. However, more are being described as deeper waters around the globe are probed. There are about 28 species in Canada, including 9 species in the Arctic. Skates are dorso-ventrally flattened with greatly enlarged, winglike pectoral fins attached to the side of the head anterior to the ventral gill openings. As a result of the greatly enlarged pectoral fins, in dorsal view the body is almost circular, and the snout may be more or less pointed. The eyes and spiracles are on the upper surface. The spiracle enables water to pass over the gills for respiration and out through the five gill openings on the underside without the gills becoming clogged by sediment. Anal fins are absent. Jaws are protrusible, and teeth form pavement-like plates. Both sexes may have pointed teeth, or both sexes may have rounded teeth, but males develop sharp, conical teeth just before maturing. The male uses his jaws to hold the female during copulation. Skates are characterized by a slender tail with a variably well-developed, reduced, or absent caudal fin; 0–3, usually 2, dorsal fins present well posterior; and prickles usually present and often developed as a midline row on the back (which may be further developed and enlarged as thorns). Malar thorns are those close to the edge of the disc in front of the eyes; alar thorns are close to the edge of the pectoral fin tips; nuchal thorns are on the nape on the midline behind the eyes; and scapular thorns are on the shoulder region behind the eyes on each side of the midline. The anterior edge of the neurocranium or skull is extended as a rostral bar. This structure may be slender and soft (Softnose Skates such as Bathyraja species) or thick and stiff (Hardnose Skates such as Amblyraja and Rajella species). The total length may be to 2.5 m, although most are much smaller. Taxonomic revisions within Family Rajidae have been progressively occurring since about 1970, with particular emphasis on the North Atlantic. At that time most existing Skates were realigned within a number of subgenera with Raja. In addition to descriptions



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of new species and their alignment with existing subgenera, two developments affected Skate taxonomy. First, a phylogenetic approach has been applied to relationships within the family (i.e., shared derived characters used to delineate taxa and support relationships). Second, as relationships became clearer, subgenera were elevated to generic status. Regional works and some older literature often do not reflect these changes. More details relevant to our area are developed in this family account and in the individual species’ descriptions. Two subfamilies, Arhynchobatinae (Softnose Skates) and Rajinae (Hardnose Skates), are presently differentiated although some authorities elevate these to the family level. Nine (and perhaps as many as twelve) representatives from these subfamilies are documented from our area: Arhynchobatinae with Bathyraja (two species, one of which is only known from an egg case and young in the western Arctic); and Rajinae with Amblyraja (three species), Malacoraja (one species), and Rajella (three species). The family is poorly documented in western Canadian and Alaskan Arctic waters, but whether this is due to low diversity or poor sampling is unknown. Regardless, the family is well known in eastern Canadian Arctic and Greenland waters as well as throughout the North Atlantic. Skates are likely more widely dispersed in Arctic waters than presently documented, especially in deeper waters. One additional species occurs immediately extralimitally in eastern Arctic waters and thus may eventually be encountered in our area: Rajella bigelowi Stehmann 1978 (Bigelow’s Skate, raie de Bigelow), a rare species from southwest Greenland. Two, and perhaps three, additional species have been identified recently from eastern Arctic waters. However, confirmed identifications and specimens are lacking, and these are not included herein. No extralimital rajids are known from the waters of Alaska immediately west of our area, but deeper areas here and in the Arctic Ocean north of Canada are poorly surveyed. All species are benthic in habitat, typically over soft substrates, and most are found in deeper water (to 3,000 m); however, some range in depth from nearshore shallows to several thousand meters. Abundance in our area ranges from very rare to common, but this may partially represent sampling artefacts. Skates swim by flapping or undulating their pectoral fins, but some have elongate pelvic finrays and “walk” over the sea bed. Many (perhaps most) Skates appear to hunt as ambush predators by agitating soft sediments, letting these settle and partially cover the body, and then laying in wait for prey to approach. Some species may also more actively prey on benthic food such as crustaceans. Crustaceans (especially decapods), fishes, marine worms, and other invertebrates are important diet items. Studies of trophic level suggest that Skates are higher level predators than generally considered (i.e., trophic levels approaching 4.0); thus, they are positioned slightly below many sharks and similar higher-level predators. Abundance and wide-spread occurrence argue for the importance of Skates in deep-water benthic communities. Sexual dimorphism is prominent, with males exhibiting pelvic claspers and alar spines on the tips of the pectoral fins, which are used to embrace the female during copulation. Females are usually larger than males. Leathery egg cases (“mermaid’s purses”) are more or less square or rectangular and may contain one to several eggs. The four elongate horns or filaments on the corners likely aid

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attachment or anchoring of the egg case. Studies suggest embryonic development may be quite long. Predators include gastropods (upon egg cases), larger fishes (e.g., other Skates and some sharks), and marine mammals such as seals. Skates are generally captured as by-catch in fisheries for other deep-water species that use bottom trawls or baited long lines. Generally, Skates are not targeted in Canadian fisheries (e.g., about 820 t in 1988 reported in total; 716 t in 2011 for the Atlantic fishery, worth $240,000). Elsewhere they are used for fish-meal, pet food, and human consumption. Discs made from the pectoral fins or “wings” are occasionally marketed as “scallops.” By-catch discards may be large in some fisheries, and thus the incidental effects of bottom fisheries are significant; these may become more so as new areas (i.e., northern and deeper waters) are fished. Similar to sharks, Skates appear to mature late in life, are slow growing and likely fairly long lived, are large bodied, and have relatively low fecundity (i.e., so-called K-selected species). Such species are inherently susceptible to anthropogenic perturbations, especially exploitation. Accordingly some especially vulnerable species (e.g., those in temperate latitudes and nearshore, heavily captured as by-catch) are of conservation concern.

sources:  Stehmann (1970); McEachran & Dunn (1998); Musick (1999); Dolgov, Grekov, Shestopal, & Sokolov (2005); Ebert & Bizarro (2007); Ebert & Compagno (2007); Sulak et al. (2009); www.iucnredlist.org, accessed 26 August 2010; FishBase, www.fishbase.org; Mecklenburg, Møller, & Steinke (2011).

Amblyraja hyperborea (Collett, 1879)

Darkbelly Skate, raie boréale

Amblyraja hyperborea

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common names:  A local name is Issittup Tarraleqisaava (Greenlandic). Other common names are Arctic Skate, Blackbelly Skate, Boreal Skate, and raie arctique. taxonomy:  The genus comes from the Greek amblys (blunt), probably in reference to the snout compared to other Skates, and the Latin raia (ray, skate). The species name comes from the Greek hyper (over or beyond) and boreios (northern), in reference to the Arctic distribution. This species was formerly in the genus Raja. Amblyraja (stout skates) is one of 26 genera in the family and subfamily Rajinae. Ten valid species are recognized within the genus, of which three occur in our area. It was originally described from Spitsbergen as Raja hyperborea and referred to as such in older and much of the recent literature. A synonym is R. borea Garman, 1899, described from off the coasts of northern Europe to north of the Faroe Islands and Iceland. This species has been confused with other taxa such as A. radiata in some literature and may be easily confused in the field with A. jenseni; thus, some information may be erroneous if identifications were not confirmed. DNA bar-coding was unable to distinguish this species from A. jenseni. It was noted that A. hyperborea is smaller and appears to mature at a smaller size and that Amblyraja may be a genus for which DNA-based identification is problematical. Recent taxonomic revisions aligned this genus with Rajinae and differentiated three distinct tribes, of which Amblyrajini is one. This work also justifies the elevation of Amblyraja from subgeneric (within Raja) to generic level. description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip; thorns are always present on the tail in a median row, and parallel rows are absent or much smaller; there are 22–32 thorns from the nape to the first dorsal fin; the dorsal fins are separated by a narrow but distinct gap with no intervening thorn (but in some young the gap is wide with 1–2 thorns); the upper-jaw tooth rows number 33–48; the body is thick and flabby, and the tail is very short, 0.7–0.8 times body length; and in adults the ventral disc is darkly blotched with spots or bands. As noted, it may be confused with A. jenseni; however, the leading edges of the pectoral fins are noticeably scalloped in this species, whereas they are more or less straight in A. jenseni. The Darkbelly Skate is a relatively large Skate with a diamond-shaped disc (young) or a spade-shaped disc (adult). The snout is pointed and slightly flexible, with a length that is about 20% of the disc width in adults. The dorsal disc has numerous thorns and many small spinules, but adults have bare areas at the centre of the pectoral fins and along each side of the midline. The shoulder has 2–4 large thorns. No thorns are present on the ventral disc. In adults the body is robust and flabby, thickened in the abdominal area, giving a “hump-backed” profile. The pelvic fins are relatively large with a small anterior lobe. The claspers in males are robust and broad. The upper-surface coloration is bluish grey with bluish-purple overtones and irregular mottling. The lower-surface coloration is white in juveniles and darker in adults with generally symmetrical darkgrey or black patterning. Coloration may be highly variable. The

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Amblyraja hyperborea, dorsal view and egg case (mermaid’s purse) (top), ventral view (bottom)



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lower-surface dark patterning may exceed the remaining whitish surface area. This species attains 106.0 cm in total length.

habitat: The Darkbelly Skate is found in cold (−0.08°C to 3.5°C)

deep waters in northern areas from 167 m to 1,600 m depths. It has been found at 340–412 m in Ungava Bay, at 275–1,600 m and −0.08°C to 1.25°C in Davis Strait and Baffin Bay for 30 collections, to 435 m in the Beaufort Sea, and to 1,067 m in Cumberland Sound, southeast Baffin Island. It is primarily at depths greater than 501 m throughout the Davis Strait and Baffin Bay. In southern Baffin Bay and Davis Strait it is found generally from 517 m to 1,483.5 m and at −0.1°C to 3.8°C. Tagged fish in Cumberland Sound were found between 1.2°C and 2.9°C, and 317 m to 1,355 m, and exhibited resting behaviour and large depth changes of more than 150 m per half hour. This species occurs primarily over silt bottoms. It is most common along upper continental slopes (800–1,500 m) in the east Norwegian Sea, but has not been found deeper (i.e., 2,000+ m). It is one of three common elasmobranchs in NAFO Subareas 0 and 1 (with Amblyraja radiata and Somniosus microcephalus).

association, which places it beyond most present exploitation. However, the by-catch noted above has led to recommendations that it be reassessed.

distribution: The Darkbelly Skate is found in Smith Sound, at Mittimatalik (Pond Inlet) on northern Baffin Island, Lancaster Sound, Baffin Bay and Davis Strait, Cumberland Sound, Ungava Bay, Hudson Strait, off western Banks Island, and the Beaufort Sea. It has been tentatively identified from a video recording on a remotely operated vehicle (ROV) at 1800 m on the Northwind Abyssal Plain (Arctic Ocean northwest of Canada, 74°20' N, 162°19' W), but this record requires verification. It is probably of much wider distribution than implied by the mapped points. It is also found south to the Grand Banks in Atlantic Canada. Globally it is distributed amphitropically in the Atlantic, Pacific, and Indian Oceans. The species is wide spread in the northeast Atlantic from the southeastern Barents Sea and Svalbard south to northern United Kingdom, and northwestwards to the Faroe Islands, Iceland, and northwest, southwest, and southeast Greenland.

biology:  Like many deep-dwelling fishes, its biology is generally poorly known, especially in the north. The species appears to be moderately common where it occurs. Feeding appears to be mostly benthic, with small and large fishes, invertebrates, and crustaceans being prominent items (33% of the diet comprises polychaetes, amphipods, euphausiids, decapods, and other crustaceans; and 67%, fishes). Pelagic amphipods and cephalopod beaks have also been documented from gut contents, as well as Clupea harengus and zoarcids. Trophic studies suggest that members of this genus actively feed at trophic levels of 3.98, approaching those of many sharks. It may be eaten by Narwhals in Mittimatalik (Pond Inlet). Maturity appears to occur late (i.e., a 600 mm male was immature, while a 850 mm male was in breeding condition). Reproduction is oviparous, with females apparently producing two egg cases, each having several eggs. Leathery egg cases are 8.1–12.5 cm long by 5.4– 7.7 cm broad and have two long horns at one end and two shorter ones at the other end. The egg cases are brownish with a layer of golden silk-like hairs covering them when newly laid (these turn brown with age). Sexes are dimorphic; males are smaller and have claspers. Reproduction involves pairing, with males embracing the female. The length at hatching is around 160–180 mm. Distribution of Amblyraja hyperborea

importance:  Directed fisheries do not occur for this species

in Canada; however, it is a significant component of by-catch for other deep-water fishes (e.g., Greenland Halibut) and is usually discarded at sea. It is an important source of income and sustenance for Indigenous and northern communities and is part of Indigenous culture. Small catches may be landed elsewhere and processed into meal. Studies in other parts of the range (e.g., southeastern Barents Sea) indicate that the species comprises 20% of catch in suitable habitats. Although relatively abundant, it is poorly known, and similarly to the other family members, it is likely an important component of the deep-water epibenthic community. It is assessed as of “Least Concern” globally, primarily due to deep-water habitat

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sources:  Jensen (1948); Bigelow & Schroeder (1953); Finley & Gibb (1982b); Anonymous (2000f); Skjæraasen & Bergstad (2001); Jørgensen et al. (2005); Stein, Felley, & Vecchione (2005); Department of Fisheries and Oceans (2007); Ebert & Bizarro (2007); Ebert & Compagno (2007); Sulak et al. (2009); www.marinebiodiversity. ca/skatesandrays/; Coulson et al. (2011); Peklova, Hussey, Hedges, Treble, & Fisk (2014).

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Amblyraja jenseni

(Bigelow and Schroeder, 1950) Shorttail Skate, raie à queue courte

common names:  A local name is Jensens Rokke (Danish). Another common name is Jensen’s Skate.

taxonomy:  The species is named for Adolf Severin Jensen (1866– 1953), a naturalist and ichthyologist of the Copenhagen Zoological Museum who conducted much work on the fishes of Greenland. This species may be confused with A. hyperborea (see distinguishing characters below and for that species). It was originally identified as Raja granulata Gill, 1879 (= Dipturus laevis (Mitchill 1818)) from the Nova Scotia area, a different taxon, and then reassigned as R. jenseni with subsequent realignment to Amblyraja.

description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip; the tail bears thorns in a median row, and parallel rows are absent or much smaller; there are 24–31 large spines from the nape to the first dorsal fin in the midline; the teeth are in 56–66 rows in the upper jaw; and the distance from the rear orbit margin to the pelvic axil is usually shorter than that from the pelvic axil to the first dorsal fin origin. This is a medium-sized Skate with a diamond-shaped disc. The snout is pointed (usually less than 90º angle) and fairly rigid with a stiff cartilage. The anterior margins of the pectoral fins are nearly straight, which differentiates it from A. hyperborea in which they are scalloped. It is firm bodied and relatively thin, lacking any noticeable “hump-backed” appearance. Two small dorsal fins are located far back on the tail with a distinct gap and no intervening thorns in adults, but 1–2 thorns are possible in some juveniles. The dorsal surface has numerous white thorns (24–31 in a mid-dorsal row on the body, and > 10 spines along the midline of the tail behind

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the pelvic fins). There are large patches of spinules dorsally. There is one large thorn on the inner side of each spiracle, 2–3 thorns on the inner side of each eye, and a patch of spines on the snout, which is lost in adults. The underside is smooth. The tail is short (0.9–1.0 times body length). The pelvic fins are relatively large with a small anterior lobe. The upper jaw has 56–66 tooth rows (33–48 in A. hyperborea). The species is sexually dimorphic, with females being larger than males and also exhibiting relatively greater values for some morphometric measurements. The upper surface is grey brown with darker blotches and tiny pale spots. The lower surface has mostly a dark-greyish symmetrical pattern on white, but may be yellowish, and juveniles have more white on the belly. Whitish areas predominate around the mouth. It attains 99.0 cm in total length.

habitat: The Shorttail Skate prefers water temperatures of 2.7°C to 4.0°C and is strictly a deep-water species found from around 365 m on lower slopes to abyssal depths of 3,000 m.

biology: This is virtually unknown but inferred to be similar to that of A. hyperborea. It likely feeds actively on small fishes and crustaceans.

importance: The species is captured as occasional specimens

only in Canadian and other northern waters and so is of incidental importance. It may be captured as by-catch in deep-water fisheries but is likely less vulnerable than other species due to depth preferences. It is globally assessed as “Least Concern” due to presumed lower vulnerability to anthropogenic impacts.

distribution: It is found in Baffin Bay, Davis Strait, Ungava Bay, and Hudson Strait. In Canada indications have it occurring only south of 55° N, which, if this is true, raises doubt regarding the distribution points shown here for the Arctic. Similar to most deepwater taxa it likely occurs in particular habitats associated with preferred water masses; thus, distribution is likely wider than suggested. The species is found on Labrador Shelf, Flemish Cap, Grand Banks, and Scotian Shelf in Canada, and south to New England of the United States. Is is also found in southwest and southeast Greenland waters. It is globally known only from more southerly North Atlantic waters, in the northeast Atlantic to Ireland, the Mid-Atlantic Ridge, and possibly Iceland.

Distribution of Amblyraja jenseni

sources:  Bigelow & Schroeder (1953); Hudon (1990a); Sulak et al. (2009); Orlov, Cotton, & Shevernitsky (2010); Neat & Campbell (2013).

Amblyraja radiata (Donovan, 1808)

Thorny Skate, raie épineuse

Amblyraja radiata

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Amblyraja radiata, with mermaid’s purse (bottom right)

common names:  Local names are Qarlêk (Inuktitut) and Adglernaq (Greenlandic). Other common names are Arctic Thorny Skate, Atlantic Prickly Skate, Maiden Ray, Miller Ray, Starry Ray, and Starry Skate. taxonomy:  The species name comes from the Latin radiatus

(rayed), in reference to the large spinous plates on the body and tail, each with a radiated or stellate base. It was originally described from British waters within Raja. Raia americana DeKay, 1842, described from Staten Island, New York, and R. scabrata Garman, 1913, from Massachusetts Bay are synonyms, and these names have been used in some older literature. In addition, specimens of this species have been misidentified as Raja clavata Linnaeus, 1758, and R. laevis (= Dipturus laevis (Mitchill 1818)).

description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip; thorns are present on the tail in a median row, but parallel rows are absent or much smaller; and large median thorns from the nape to the first dorsal fin number 11–20. In addition, the upper surface is covered with thornlets and is very rough to touch; upper-jaw tooth rows number 27–49; the dorsal fins are often separated by a distinct gap, without an intervening thorn; and the tail is 1.0–1.1 times body length. It is a moderate-sized Skate with a spade- to a heart-shaped disc having rounded corners and generally not being emarginated



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(the leading edge is not scalloped but does bulge slightly behind the eye). The snout is rigid and blunt, forming an angle of about 110º–140º. The dorsal thorns are very large and form a single dominant mid-dorsal row extending down the tail. Thorniness is reduced with age, especially in mature males. Ten or fewer thorns lie behind the pelvic fin axils, and a variable patch of larger thorns is present medially on each wing. The centre of the disc tends to be smooth between the larger thorns there. Thorn counts are inversely related to temperature; conversely tooth-row counts are positively related to temperature. Each shoulder has two to three thorns, each eye has one thorn in front and one behind, and there is one thorn on the inner side of the spiracle. Males have erectile, hooked alar spines in two to five rows near the outer edge of each pectoral fin. Adults have spines on the underside of the snout. Although the dorsal fins are often separated by a distinct gap lacking thorns, the fins may be joined and have up to five intervening thorns. The upper-tooth-row count is 27–49, but 27–36 in Ungava Bay. The pelvic fins are relatively large with small anterior lobes. Sexually mature males have large claspers. Coloration on the upper surface is usually more or less uniformly brown or grey brown in adults, but may have yellow-edged, light pseudo-rosettes on light brown, or may be dark brown. A white spot is typically present near each eye. Lower-surface coloration is generally uniformly white, sometimes with brownish or greyish splotches. The species attains 110.0 cm in total length and 12.5 kg in weight.

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habitat:  The Thorny Skate is primarily found on the continental shelf from about 5 m to 1,478–1,540 m depths in waters from −1.4°C to 14°C, and especially in the mid-range of these depths and temperatures. It has been recorded at 115–860 m in Ungava Bay (36 collections), 0–530 m in Hudson Strait (144 collections), and 146–1,441.5 m and 0°C–5.4°C in Davis Strait and southern Baffin Bay generally. It is primarily found at less than 751 m in Davis Strait in NAFO Division 0A. The species is one of three common elasmobranchs found in NAFO Subareas 0 and 1 (with Amblyraja hyperborea and Somniosus microcephalus). As with other Skates, it is benthic over a variety of substrates ranging from sand to mud, and pebbles to broken shells. biology:  Its biology is considerably better known than that of

most Skates, likely because it occurs in shallower waters and is regularly captured. Its diet is mostly invertebrates (70% polychaetes, mollusks, squids, cephalopods, amphipods, euphausiids, decapods, and other crustaceans) and fishes (30%). Overall it feeds at a trophic level of about 3.82. Food as young consists of amphipods, euphausiids, and mysids and then as adults switches to various worms (e.g., polychaetes) and larger crustaceans (e.g., crabs); fish prey includes Sculpins, Capelin and Sand Lances, and even Greenland Halibut. It consumes by-catch and the discards from commercial fishery vessels. Some seals are known to prey upon them, and egg cases have also been recovered from halibut stomachs and Greenland Sharks. This species lives to at least 20 years (per tag recovery) as generally verified through analyses of aging structures such as vertebrae, with a suggested maximum of 39 years. Both sexes mature at sizes of about 40–50 cm total length (5–6 years of age). Female length at 50% maturity was about 45 cm for fish from Baffin Island and the Labrador Shelf – less than that recorded for other areas in Atlantic Canada (48–70 cm). It is oviparous, and reproduction apparently occurs year round in the south. The egg-capsule size is larger in larger females and ranges 4.2–9.6 cm by 2.5–7.7 cm; size also appears to be geographically correlated (larger in the south). Each female produces 20–80 eggs annually. The total length at hatching is about 10.0 cm. Tagging evidence suggests that they occupy home ranges in a local area. Notwithstanding local demography, genetic evidence suggests long-distance historical dispersal across the North Atlantic and also significant expansion of population sizes in periglacial times.

slower growing and longer lived than was previously thought, which increases vulnerability to perturbations, especially exploitation. Such vulnerabilities are being recognized as conservation concerns. It is listed as “Not Threatened” in Norway; however, commercial harvesting is restricted in U.S. waters due to declining abundances. The Committee on the Status of Endangered Wildlife in Canada has assessed it as “Special Concern.” The global listing is “Vulnerable,” based upon some local declines and potential susceptibility to exploitation. Although it is caught as by-catch and usually discarded at sea in commercial fisheries, it is an important source of income and sustenance for Indigenous peoples and northern communities and is part of Indigenous culture.

distribution:  This species is found in Baffin Bay, Davis Strait, Hudson Strait, Ungava Bay, and northern Hudson Bay. It has also been recorded from the North Water Polynya and in northwest, southwest, and southeast Greenland waters. It is present in the western Atlantic to South Carolina including the Grand Banks and Labrador Shelf. In the northeast Atlantic it occurs to the eastern Barents Sea, and south to the northern British Isles. It is also reported from southern Atlantic waters near South Africa.

importance:  This is a very common Skate whose landings are

converted to fish-meal. It is not presently of economic importance in Canada despite its abundance. Catches in survey tows in NAFO Subareas 0A and 0B varied from 5 to 135 fish. Incidental by-catch in Greenland Halibut and northern shrimp fisheries may be significant but is generally poorly documented in Canadian Arctic waters. The peak catch was in the shrimp fishery in 1994 at 46 t, but it was less than 1 t in 2008 and 2009. It is the most common species in by-catches in Barents Sea fisheries and is marketed as a table fish in Europe. It is especially common in the coastal waters of northern Norway, where biomass increases in northern areas. Similar to other members of the group, this species generally appears to be

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Distribution of Amblyraja radiata

sources:  Jensen (1948); Bigelow & Schroeder (1953); Templeman (1984a, 1987a, 1987b); Hudon (1990a); Anonymous (2000f); Hobson, Fisk, et al. (2002); Dolgov et al. (2005); Jørgensen et al. (2005); Chevolot et al. (2007); Department of Fisheries and Oceans (2007); Ebert & Bizarro (2007); Williams, Helle, & Aschan (2008); McPhie & Campana (2009); Sulak et al. (2009).

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Bathyraja sp.

description:  It is known only from egg cases and young collected in the Northwest Territories. It was originally listed as Raja (Amblyraja) sp. but may represent a new species.

common names: None.

habitat:  It has been recovered from a depth of 335 m.

taxonomy:  The taxonomy is unknown, but if this is a species associated with Bathyraja, it could be one of several species documented for western Alaska or a new species. See the Bathyraja spinicauda species description for more information on the genus.

biology:  Unknown. importance:  Unknown. distribution:  It has been found in Franklin Bay, Cape Bathurst

area of western Amundsen Gulf (CMNFI 1977-0976 and 1977-1692).

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(Greenlandic). Other common names are Spinetail Ray and Spinetail Skate.

taxonomy:  The genus comes from the Greek bathys (deep) and raia (ray, skate). The species name comes from the Latin spina (spine, thorn) and cauda (tail). This species was originally described from Davis Strait (southwest Greenland) in Raja and subsequently placed in a subgenus (Bathyraja) within Raja. Recent phylogenetic taxonomic revision of rajids distinguished two subfamilies, demonstrated that Raja was polyphyletic, and aligned Bathyraja with Arhychobatinae (Softnose Skates). This work also elevated Bathyraja to the generic level; however, derived diagnostic characters for this taxon are lacking so it may be polyphyletic (i.e., composite). The genus includes about 50 species worldwide and achieves its highest diversity at high latitudes.

Distribution of Bathyraja sp.

sources:  See the family sources and the bibliography.

Bathyraja spinicauda (Jensen, 1914)

Spinytail Skate, raie à queue épineuse

description:  This species is distinguished by having the anterior pectoral fin-rays extending almost to the snout tip (which may be seen best by shining a light through the flesh). In addition, the rostral bar is soft and flexible; there is a thorn between the dorsal fins (absent in young), no thorns on the disc, and 21–26 thorns on the tail midline; upper-jaw tooth rows number 30–34; and the tail is short, 0.9 times body length. It is a very large Skate of deeper waters, with a long and pointed soft snout having thin, flexible cartilage. The disc is spade- to diamond-shaped with straight margins on the leading edges. There are no spines on the upper surface of the disc, with the exception of a median row beginning posteriorly and extending along the tail. There is a dense covering of small spinules on the upper surface. The lower surface is smooth, lacking spinules on the tail. Like most other species, mature males have three to four rows of spines on the outer tips of the pectoral fins (alar thorns) and have small claspers. There are two small dorsal fins near the tip of the tail, with the gap between the dorsal fins containing a large thorn. The pelvic fins are bilobed and relatively large. The tail has an obvious lateral fold along the lower margins. The upper jaw has 30–34 tooth rows. The upper surface is light grey to brown in adults. The lower surface is white or light coloured with greyish margins. Smaller juveniles have a few dark bands on the upper surface of the tail. The species attains 170.0 cm in total length. habitat:  This is primarily a cold-water species with temperature

preference appearing to be −1.5°C to 7.5°C, and most often below 3°C. It has been recorded from as shallow as 455 m in Davis Strait, and at 941–1,463 m at 0.6°C–3.5°C in southern Baffin Bay and Davis Strait. There is high abundance near frontal zones mixing southern (warmer, shallower) waters with northern (cooler, deeper) waters on upper slopes, which appears to be a preferred habitat. This species occupies deeper slope and abyssal habitats at 111–2,949 m depths, likely over muddy, silty, and sandy bottoms. Bathyraja spinicauda

common names:  Local names are Taqqalerisaaq for the

adult, and Kanatik for the egg case, presumably of this species

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biology:  Its biology is generally poorly known due to deep-

water habits. It feeds on benthic invertebrates (about 40% of its diet includes polychaetes, amphipods, euphausiids, decapods, and

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Bathyraja spinicauda

other crustaceans) and fishes (60%) including the young of other Skates, which results in a trophic level of 4.02. Its weight at 139 cm equals 18.5 kg. It is oviparous, and egg cases are 13.7–14.2 cm by 9.0– 9.5 cm with a unique set of longitudinal ridges having stiff rod-like structures, which results in a velvety feel to the egg case. Embryonic development is about a year.

importance:  This species is common where it occurs, but over-

all it is rare; however, given the preferences for deeper waters, it is captured infrequently. Like other Skates, it may be at risk from anthropogenic perturbations including fisheries. It is listed as “Data Deficient” in Norwegian conservation literature. Its global conservation assessment is “Near Threatened” based upon presumed life-history constraints (see the Family Rajidae description), regular capture as by-catch in deep-water fisheries, and declines apparent for Canadian waters (it has also been assessed as “Vulnerable” regionally for the northwest Atlantic, and as “Least Concern” for the northeast Atlantic).

distribution:  It is found in Baffin Bay, Davis Strait, Hudson Strait, and Ungava Bay. Its deep-water habit suggests that it is likely more widely distributed than is indicated on the map. It is also found in southwest and southeast Greenland waters. Globally it is known only from the North Atlantic, in the east from the Barents Sea south to the northern North Sea, westwards to the northern United Kingdom, Faroe Islands, and Iceland, and in the west south to the northeastern United States.



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Distribution of Bathyraja spinicauda

sources:  Jensen (1948); Ishiyama & Hubbs (1968); Hudon

(1990a); Skjæraasen & Bergstad (2001); Ebert & Bizarro (2007); Ebert & Compagno (2007); Williams et al. (2008); Sulak et al. (2009).

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Malacoraja spinacidermis (Barnard, 1923)

Soft Skate, raie molle

common names:  A local name is Lodden Rokke (Danish/ Greenlandic). Other common names are Prickled Ray, Roughskin Skate, and raie profonde

taxonomy:  The genus comes from the Greek malakos (soft), in

reference to the dense, velvet-like covering of fine dermal denticles on the upper surface of the type species of the genus, Raja mollis (= Malacoraja spinacidermis), and Latin raia (ray, skate). The species name comes from the Latin spina (spine, thorn) and the Greek akis (point) and derma (skin). It was originally described in Raia, then realigned to Malacoraja. A synonym is Raja mollis Bigelow and Schroeder, 1950, described from south of southern Nova Scotia, 41°53'00" N, 65°35'00" W. It is one of three species in the genus that is a member of the subfamily

Rajinae (tribe Gurgesiellini). Originally collapsed within Raja, this subgenus was elevated to generic level upon recent revision.

description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip, and the disc behind the shoulder, and much of the tail, being without thorns. In addition, fine spinules (minute spines) are present over the whole dorsal surface and the underside of the tail; the dorsal fins are connected, with no thorn between them; the rostral bar is flexible; the upper-jaw tooth rows number 54–63; and the tail is long, 1.3 times body length. The Soft Skate is a medium-sized Skate with a spade-shaped disc that is broader than it is long, with widely rounded outer corners. The snout is short but pointed (the snout angle is 90º or greater) and very soft or pliable (unusual for the subfamily). A few small thorns occur near the eyes and on the shoulders (nuchal area), with no thorns anywhere on the disc behind these. A short row of mid-dorsal small thorns is present on the tail. The dorsal fins are connected, with no intervening thorn or gap. The lower body surface is bare, with fine spinules along the disc. The upper surface is uniform

Malacoraja spinacidermis

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greyish brown in colour. Larger individuals are dark ventrally with medial white remnant patches. In younger individuals the lower disc surface is whitish with scattered grey flecks, and the underside of the tail is uniformly grey. The species is typically around 48.0 cm in total length but has been recorded to 70.0 cm.

sources:  Jensen (1948); Bigelow & Schroeder (1953); McEachran & Dunn (1998); Jørgensen et al. (2005); Ebert & Bizarro (2007); Sulak et al. (2009).

habitat:  It dwells in deeper benthic habitats on continental

slopes to bathyal depths (475–1,570 m), with adults being distributed in deeper waters generally of 3°C–4.5°C. It has been recorded at 1,086 m in the Canadian Davis Strait.

biology:  Its biology is generally poorly known. The species is

oviparous, and a female captured at 52 cm total length was fully mature. A limited sample indicates that the diet consists of decapods (75%) and fish (25%), with a trophic index of 3.7.

importance:  This species is occasionally captured as by-catch. It

is globally assessed as “Least Concern” because it occupies habitats typically beyond the range of general fishing activity.

distribution:  It has been found in Davis Strait as a single record

at 62.6774º N, 60.4071º W from cruise data. It is likely more widely distributed in our area than indicated. It is also known from southwest and southeast Greenland. The Soft Skate is distributed globally in both North and South Atlantic waters. It has been recorded as a rare species south to the Grand Banks in the western Atlantic and from Iceland, Faroe Islands, and likely elsewhere in the North Atlantic. It has also been reported from Namibia and South Africa.

(Holt and Byrne, 1908) Abyssal Skate, raie bathyale

common names:  A local name is Dybhavsrokke (Danish/

Greenlandic). Other common names are Chocolate Skate, Deepwater Ray, and raie chocolat. Chocolate Skate also refers to Rajella bigelowi, thus, the preference for Abyssal Skate as the common name.

taxonomy:  The genus comes from Latin raia (ray) and -ella (the feminine suffix used to indicate diminutive of a noun, hence Rajella, a small ray). The species name comes from the Greek bathy (deep) and phila (loving). It was originally described as Raja bathyphila from the deep waters of the Irish Atlantic slope (about 1,220–1,600 m); however, both previous and subsequent workers have referred specimens and this taxon to Raja lintea (in part). Thus confusion exists in older literature with respect to characters, and some nomenclatural synonymies are inherent. It is considered as a distinct and valid taxon based upon more recent research (see the “Description” section). This genus is phylogenetically associated with Amblyraja, Breviraja, Leucoraja, and Dactylobatus in the Amblyrajini of Rajinae, and thus is related to Raja and Malacoraja species found in our area. As for other presently recognized genera from our area within this family, originally they were collapsed as a subgenus within Raja (which was demonstrated to be polyphyletic) and then raised to generic level upon recent revision. description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip; thorns are present on the tail in a median row, parallel rows being absent or much smaller; and there are 31–44 large spines from the nape to the first dorsal fin in the midline (33 or more in adults). In addition, the lower surface of the disc, pelvic fins, and tail is uniform dark brown in young, darker than the grey-brown upper surface (but lighter in adults); upper-jaw teeth number 34–42 rows; the tail is long, 1.4 times body length; and the dorsal fins are connected, with no intervening thorn. As noted in the “Taxonomy” section, specimens are often referred to Rajella lintea, but the anterior pelvic lobe is smaller and shorter than the posterior lobe in the latter species. The Abyssal Skate has also been confused in part with Rajella bigelowi, which has not yet been recorded from our area but which occurs immediately to the south and to the east (in Greenlandic waters) of southern

Distribution of Malacoraja spinacidermis



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Rajella bathyphila

Davis Strait. This latter species is distinguished by fewer thorns in the mid-dorsal row and a patch of thorns along the rostrum. The Abyssal Skate is an intermediate-sized Skate with a diamond-shaped disc in adults (heart-shaped in juveniles) with slightly emarginate leading edges and rounded outer corners. The snout is rigid, short, and rounded. Thorns are absent on the snout. A few orbital thorns (3–4) and 5–6 thorns in a nuchal triangular-shaped patch are present. The mid-dorsal row consists of 31–44 thorns paralleled by an additional row of 10–30 thorns on each side of the tail posterior to the pectoral insertion. Fine spicules cover the upper surface, and the lower surface is smooth. The anterior and posterior lobes of the pelvic fin are more or less equal in length. The upper surface is brown to dark grey with slight darkening in posterior areas of the disc and pelvic-fin margins. The lower surface is mostly (or entirely) white in adults, with dark marginal markings. The underside of the tail is dark with whitish patches. In juveniles

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the lower surface is dark brown with a white area along the midline. The tail is white in young, darkening to grey in adults. The species reaches 93.0 cm in total length

habitat:  The species is bathydemersal on lower slopes typically

deeper than 1,400 m and possibly also abyssal plains, although it has been captured at 600–2,300 m in western Atlantic areas, likely in waters between 2.5°C and 4°C. It probably occurs over soft substrates, similarly to other Skates in our area.

biology:  It is an uncommon Skate occasionally captured as

by-catch in deep-water fisheries but poorly known. Biological understanding is likely confused due to the taxonomic and identification problems noted earlier. Very limited diet information indicates an overall trophic index of 3.80 (polychaetes, amphipods, euphausiids, decapods, and other crustaceans). The species is

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oviparous, with maturity attained at about 65 cm total length. Its size at birth is about 11 cm in total length. Egg cases are 9 cm by 5 cm flattened structures (excluding corner filaments).

importance:  Presumably it is caught in low numbers as by-catch

in deep-water trawl fisheries, but many are likely misidentified, and most are discarded. It is assessed as “Least Concern” globally due to its occurrence primarily in deeper areas beyond those that are presently fished.

distribution:  The species has been found in Baffin Bay as

a single record from cruise data at 72.01° N, 71.17° W., and also in southwest and southeast Greenland. It is present in Atlantic waters: in the western Atlantic south to the Norfolk Canyon off the United States; and in the eastern Atlantic from Iceland south to the Rockall Trough near Ireland, in the Bay of Biscay, and to the western edge of the Sahara off Africa.

Distribution of Rajella bathyphila

sources:  Bigelow & Schroeder (1953); McEachran & Stehmann (1977); Ebert & Bizarro (2007); Williams et al. (2008); Sulak et al. (2009).

Rajella fyllae (Lütken, 1887)

Round Skate, raie ronde

common names: A local name is Fyllas Rokke (Danish/Greenlandic). Another common name is Deepwater Ray.

taxonomy:  The species name comes from Fylla, the name of the Danish cruiser from which the type-specimen was taken in Davis Strait, Greenland. It was originally described in Raja. Breviraja marklei McEachran and Miyake, 1987, described from off Nova Scotia, Canada, 44°45.1' N, 56°07.1' W, has also been referred to this taxon but is regarded as distinct by authors. description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip, and thorns on the body and the tail are in two to five parallel rows, the median row being much smaller or only in young. In addition, the tail is long (1.3–1.6 times body length); there are 20–30 large thorns on the nuchal-shoulder region in a triangular patch (which differentiates it from the Abyssal Skate); the snout is very short; upper-jaw tooth rows number 30–38; and the lower surface of the disc and tail is bare. A small-sized Skate, it has a distinctly spade-shaped disc with rounded outer corners. The rigid snout is very short and rounded with a small terminal tip. The snout angle is 115º or greater. Thorns continue as three central rows bordered by two dorso-lateral rows along the dorsal surface of the tail (these are claw-like). Younger individuals are often less spinose and have only one dorsal thorn row (however, the tail length can be used to differentiate them from other species). The upper surface is also covered in tiny dense spinules (i.e., it lacks bare areas on the disc or between the thorns). The dorsal fins are connected, with no intervening gap or thorn. The upper surface is light grey to chocolate brown with spots or mottling, a distinct pale rectangular patch may be present between the eyes, and a pale area present on each side of the midline near the tip of the snout. The lower surface is whitish in colour, with darker markings often present marginally and on the pelvic fins. The species reaches 57.0 cm in total length in the western Atlantic but 68.0 cm in Norwegian waters. habitat:  The species is bathydemersal and inhabits deeper

benthic habitats on shelves and slopes in depths centred upon 300–800 m (with upper and lower limits about 170 m and 2,050 m, respectively) and waters of 1°C–7°C (usually 4°C–6°C). In the Canadian Baffin Bay and Davis Strait it has been recorded at 472–1,272 m. It likely prefers soft substrates, as do most other Skates in our area.

biology:  The Round Skate is a moderately common Skate where

it occurs, but overall it is poorly known. Its diet consists of mostly invertebrates (98%, made up of 30% polychaetes, 35% amphipods, 6% euphausiids, 24% decapods, 2% cephalopods, and 1% other



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Rajella fyllae

crustaceans); fishes make up 2%. The overall trophic index is 3.78. It is oviparous, with females maturing at around 50 cm total length, and males at around 44–47 cm. The size at hatching is around 70 mm. Egg cases are black and about 3.6–4.4 cm long and 2.6 cm wide, with long filaments of about 40 mm in length and short filaments of about 22 mm (often damaged). Gastropod predation upon egg capsules is evident.

importance:  It is regularly captured at low levels as by-catch in

bottom-trawl and longline fisheries, but most are likely discarded. It is assessed as “Least Concern” globally due to the potential for moderate-to-high population growth (i.e., it is somewhat resilient to

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fishing), stable populations, and the depth preferences placing them mostly outside heavily fished areas.

distribution:  The species is found in Baffin Bay and Davis Strait, and also southwest and southeast Greenland. Globally it occurs in North Atlantic waters, in the northwest south to Nova Scotia and the Grand Banks, and in the northeast from Iceland, the Faroe Islands, northern British Isles, and Norwegian waters to the southeastern Barents Sea. It occurs south as far as Spanish waters.

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are diagnostic for Dipturus). This was further complicated due to the lack of a mature male specimen from which diagnostic clasper characters could be determined. Accordingly, the recent alignment of this taxon with Dipturus appears to be erroneous, and re-analysis including examination of a mature male aligns this taxon with Rajella as followed here.

Distribution of Rajella fyllae

sources:  Jordan & Evermann (1896–1900); Jensen (1948); Bigelow & Schroeder (1953); McEachran & Dunn (1998); Skjæraasen & Bergstad (2001); Dolgov et al. (2005); Ebert & Bizarro (2007); Williams et al. (2008); Sulak et al. (2009).

Rajella lintea (Fries, 1838)

Linen Skate, raie linon

common names: A local name is Hvidrokke (Danish/Green-

description:  This species is distinguished by the anterior pectoral fin-rays not extending almost to the snout tip; thorns are present on the tail in a median row, and parallel rows are absent or much smaller; and there are 40–51 prominent, large spines from the nape to the first dorsal fin in midline (about 35–36 on the tail). In addition, the lower surface of the disc and the pelvic fins is whitish with a broad grey margin from wing tips posteriorly; the median underside of the tail is grey; a grey blotch is on each side of the vent; the lower surface is paler than the upper surface, which is fawn grey to uniform cream; large thorns are white in contrast with the disc; the upper-jaw teeth number 47–51; the tail is relatively short (0.9– 1.0 times body length); and the dorsal fins are connected or have a very small gap, with no intervening thorn. In our area it may be confused with A. hyperborea or A. jenseni, but the snout is longer in R. lintea and no dark markings are present centrally on the lower surface. Also the number of mid-dorsal thorns is much larger. It may also be confused with R. bathyphila recently confirmed in our area; the short and small anterior pelvic fin lobe and symmetrical markings distinguish R. lintea. Some recent literature does confuse these latter two species. It is a moderately large Skate with a diamond-shaped disc and a long, pointed, and rigid snout (3.2–4.4 times the interorbital distance, compared to 2.0–2.4 times in Amblyraja hyperborea and A. jenseni). The snout angle is usually less than 90º. There are two to four thorns in front of the eyes, one thorn at the inner eye edge, four spines behind the eyes, and three spines on each shoulder. The lower disc is smooth. The anterior lobe of the pelvic fin is small relative to the posterior lobe. The colour is given above. The species attains 125.0 cm in total length and 9.06 kg in weight. habitat:  It is present in benthic areas in deeper waters (270–2,117

landic). Other common names are Pale Ray, Sailray, White Skate, and raie voile.

m in Canada, elsewhere as shallow as 150 m) at colder temperatures (3.6°C–6°C), over sandy, muddy, and rocky bottoms. It is generally found deeper than 250 m.

taxonomy:  The species name comes from the Latin linteum

biology:  Its biology is generally very poorly known. It appears

(linen), in reference to the pale or greyish colour like a linen sail. Originally described from Northern Europe, Bohuslän (Sweden), and the North Sea (eastern North Atlantic) in Raja, it was subsequently realigned to Dipturus and also as Leucoraja in some literature. Raja ingolfiana Lütken, 1898, described from off Holstenborg, Greenland, 66°35' N, 56°38' W, is a synonym. Recent taxonomic revision of rajids distinguished two subfamiles and aligned this genus with Rajinae (tribe Rajini). This work also resurrected Dipturus as a distinct genus rather than a subgenus within Raja, with which this taxon was aligned, in part due to the absence of several distinct characters (none of which



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to be a rare Skate, found at depth and feeds on various benthic organisms including amphipods, decapods, and other invertebrates (65%) and fishes (35%); thus, it is at a moderately high trophic level (3.85). The species is oviparous, and males mature at 97 cm total length. Egg cases are 10.7 cm by 7.7 cm in size, cross-hatched with longitudinal striations, and equipped with four horns ending in fine points.

importance:  This Skate is not commercially important but

is likely taken at low levels as by-catch in deep-water longline and trawl fisheries. It is listed as “Data Deficient” in Norway. It is globally

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Rajella lintea

assessed as “Least Concern,” primarily due to the deep distribution placing it beyond most immediate anthropogenic threats.

distribution:  The species is found in Baffin Bay and Davis Strait. Distribution in our area is likely wider than indicated, given the depth preferences and the lack of sufficient sampling. It is a rare deep-water Skate globally present in North Atlantic waters. In the western North Atlantic it occurs from southwest Greenland south to Flemish Cap and Grand Banks, and in the eastern North Atlantic from Iceland south to the northern British Isles and east to Sweden and southern Norway, where it is relatively common locally. It is occasionally captured in northern Norway and also eastwards into the Barents Sea in areas warmed by Atlantic waters. sources:  Jensen (1948); Bigelow & Schroeder (1953); McEachran & Dunn (1998); Skjæraasen & Bergstad (2001); Dolgov et al. (2005); Ebert & Bizarro (2007); Williams et al. (2008); Sulak et al. (2009); Stehmann (2012).

Distribution of Rajella lintea

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Acipenser fulvescens

Family Acipenseridae

Rafinesque, 1817

Lake Sturgeon, esturgeon jaune

Sturgeons, Esturgeons common names:  Local names are Namew, Nameo, Nemeo, Némèw, and Nimàw (Cree). Other common names include Black Sturgeon, Bony Sturgeon, Common Sturgeon, Dogface Sturgeon, Freshwater Sturgeon, Great Lakes Sturgeon, Red Sturgeon, Ruddy Sturgeon, Stone Sturgeon, and Shellback Sturgeon; Smoothback, Rubber Nose, esturgeon de lac, and camus; and, for the young Rock Sturgeon, escargot maillé and charbonnier.

James D. Reist

The Family Acipenseridae has 26 extant species in four genera, with five species occurring in Canada, of which only one is in the Arctic. Acipenseriform fishes represent an old group (approximately 190–200 million years before present) of bony fishes that have secondarily developed a cartilaginous skeleton. Sturgeons (Acipenseridae) and Paddlefishes (Polyodontidae) represent the extant families, and both are primarily distributed in northern hemisphere temperate waters. Only Sturgeons have a representative in Canadian Arctic waters. Two subfamilies are generally recognized, although the taxa below this level (i.e., tribes and generic composition) in North America are all in dispute. Sturgeons are very large (perhaps 9 m and 1,451 kg) freshwater and anadromous fishes with five rows of bony scutes along the body rather than scales, and a long, pointed snout over-hanging a subterminal mouth. Four barbels occur immediately anterior to the mouth, and teeth are absent. The caudal fin skeleton is upturned, resulting in a heterocercal tail with a large, pointed upper lobe and a smaller lower lobe. Unpaired fins have fulcra, or flat bony plates, distinct from scutes, in front of them. The intestine has a spiral valve that prolongs the passage of food for better digestion. All Sturgeons are long-lived (reportedly over 150 years) and slow-growing fish with low survival to adulthood despite high fecundity. Sturgeon eggs (i.e., caviar) and flesh are sought in fisheries. Benthic migratory habits in fresh water and their biological characteristics make them very susceptible to anthropogenic impacts; thus, most species are considered at high risk. A single record interpreted as Acipenser oxyrinchus Mitchill, 1815, from the George River in Ungava Bay by Scott and Scott (1988) has never been duplicated. The original record (Low, 1896) refers only to Accipenser sp. (sic), and other records in the same paragraph are from James Bay. Note that the La Grande River of James Bay was formerly called the Fort George River.

sources:  Low (1896); Vladykov & Greeley (1963); Bemis & Kynard (1997); Dick, Jarvis, Sawatzky, & Stewart (2006).



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taxonomy:  The genus comes from the Latin acipenser (sturgeon), itself from the Greek akis (point) and pente (five), referring to the five rows of body scutes. The species name comes from the Latin fulvus (tawny) and -escens (becoming), in reference to the olivebrown coloration on the upper half of the body. Acipenser rupertianus Richardson, 1836, described from the Albany River (of James Bay), Prince Rupert’s Land, is a synonym. Moose River (James Bay) tributary stocks were examined using mitochondrial DNA but were not found to be discrete. However, these sturgeons were genetically differentiated from Great Lakes–St Lawrence drainage populations and other Hudson Bay–James Bay populations. description:  The long snout, the subterminal mouth preceded by four barbels on the underside of the snout, the heterocercal tail, and the bony scutes make this fish unmistakable. This species is differentiated from others in North America by having one large scute (plate) on the venter between the anal fin and the caudal flexure (in addition to the fulcrum), and typically 32–35 gill rakers (range is 25–40). It is a very large fish with a torpedo-shaped body that in adults is rounded in cross-section. The scutes are most obvious in young fish, becoming embedded within the flesh as the fish ages. There are 29–43 lateral scutes, 8–17 dorsal scutes, and 6–12 ventral scutes. Scute coverage of the body becomes reduced with age, presumably because such large fish have no predators apart from Lampreys. Dorsal fin-rays number 35–45, and anal fin-rays 25–30. The colour in subadults and adults is olive brown to grey dorsally and on the sides, and whitish on the belly. The flanks may be reddish. The colours of the scutes and fins match those of the adjacent body areas. The body cavity organs are black, but the peritoneum is silvery and only slightly pigmented. Young fish, less than 30 cm, have two black blotches on the upper snout, a black blotch between the dorsal and lateral plates above the pectoral fin base, and another similarly positioned blotch below the dorsal fin, there are smaller spots on much of the rest of the body and on the head, and the lower parts of the body are greenish. This species reaches 274.5 cm in total length (the Canadian record is 241.4 cm in total length and 140.9 kg in weight). habitat:  The Lake Sturgeon is primarily a freshwater species found in large lakes and rivers usually associated with soft-bottomed

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substrates between 5 m and 9 m depths. Anadromy is exhibited in association with larger river discharges in Hudson and James Bays. However, it is likely that marine habitat usage is restricted to brackish areas along coasts near estuaries. A tagged sturgeon moved from the Hayes River into the Nelson River and through Hudson Bay into the Hayes River, Manitoba. A salinity level of 15‰ can be tolerated, but not above 25‰. Brackish-water habitat usage is also exhibited in the St Lawrence River.

biology:  Feeding is primarily upon benthos in fresh waters.

Feeding in brackish marine areas is poorly known but is likely upon estuarine and nearshore benthos. Food is detected by the barbels and taken up by the protruded, tubular mouth. Ageing is difficult, with some estimates ranging to 154 years for the oldest recorded fish. Development is slow (e.g., 10 years to achieve 60 cm length, 25 years to 1 m in Lake Nipigon; 3 cm/year and 100 g/year for the first 7 years, 2.41 cm/year and 388 g/year between 11 and 28 years, and 1.7 cm/year and 147 g/year beyond 28 years in the Moose River system of the Hudson Bay lowland). Maturity is between ages 8 and 20 (males), and 14 and 33 (females), and annual reproduction appears unlikely especially for females, with intervals for the latter at 4–10 years. As with many other fishes, rates for vital parameters are reduced in northern areas. This, and opportunity, may account for anadromy and marine occurrence in our area. Adults migrate to spawn mostly in rivers; however, suitable habitats in lakes may also be used. Spawning occurs in April–June at 10ºC–18ºC in flowing water or on rocky lake margins where wave action oxygenates the eggs. An early June migration upriver from estuaries has been reported for James and Hudson Bay populations. The Nelson River basin fish spawned between 8 and 14 June

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at water temperatures between 16.5°C and 17°C. Each female may be flanked by 2–8 males who lash her flank with their tails to stimulate egg release. Spawning may involve splashing and leaps clear of the water. This leaping or tail-walking behaviour at other times is attributed to attempts to remove parasitic lampreys, or to air gulping by this physostomous fish. Ripe eggs are up to 3.5 mm in diameter, and larger fish are calculated to produce several hundred thousand to millions of eggs. The adhesive eggs are scattered and attach to rocks and logs.

importance:  Due to spawning congregations at the plunge

pools of falls, this fish proved to be a reliable source of food (flesh and roe) for both Indigenous peoples and early explorers of central North America. Extensive and wasteful exploitation in the 1800s included use of the oily fish to fire boilers on steamboats, for animal feed, and for fertiliser. Later use as high-value smoked flesh, caviar, and isinglass (a gelatin made from the large gas bladder and used as a wine and beer clarifier and in jams and jellies) led to continued high exploitation. It has been smoked and eaten by subsistence fishers in James and Hudson Bays. Indigenous harvests in the Hudson and James Bay Lowland, Ontario, in 1989–90 numbered 3,850 fish, with a projected number of 4,768 fish. The average edible raw weight per fish was 5.68 kg. The Cree and Inuit of James Bay and northern Québec harvested an estimated 11,553 sturgeons from 1974 to 1979, mostly in summer and fall (87%), and the rest in winter. Overfishing, combined with sturgeon biology (i.e., slow growth, late maturity, low survivorship to adulthood) and habitat impacts (e.g., dams limiting migrations), has depressed populations, most of which are assessed at high levels of threat (e.g., Western Hudson Bay, Saskatchewan River, and Nelson River populations are all “Endangered,” and Southern Hudson Bay–James Bay populations

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are of “Special Concern” based upon assessment by the Committee on the Status of Endangered Wildlife in Canada). Anglers may pursue this species in more southern fresh waters, including by winter spear fisheries, and it may be hooked or snagged accidentally.

distribution:  Lake Sturgeon are confined to central North American watersheds from the southern Arctic (Hudson and James Bays), south to the Laurentian Great Lakes and drainages to the St Lawrence River, and the central Mississippi River basin of the United States. The known occurrences in Arctic marine waters correspond directly with adjacent freshwater distributions known for Canada, that is, restricted to coasts of southwestern Hudson Bay and James Bay. The distribution map shows estuarine records for the species.

Distribution of Acipenser fulvescens

sources:  Berkes & Mackenzie (1978); R. Morin, Dodson, & Power (1980); Threader (1981); James Bay and Northern Quebec Native Harvest Research Committee (1982c); M. Ferguson et al. (1993); Guenette, Fortin, & Rassart (1993); Berkes, George, Preston, Hughes, Turner, & Cummins (1994); Bemis, Findeis, & Grande (1997); Birstein, Bemis, & Waldman (1997); Ferguson & Duckworth (1997); McDonald, Arragutainaq, & Novalinga (1997b); Wirgin, Stabile, & Waldman (1997); LeBreton & Beamish (1998); Barth & MacDonell (1999); Bernatchez & Saint-Laurent (2004); D.B. Stewart & Lockhart (2005); COSEWIC (2006); Peterson, Vecsei, & Jennings (2007); McDermid, Wozney, Kjartanson, & Wilson (2011).

Family Notacanthidae Deep-sea Spiny Eels, Poissons-tapirs à épines

Brian W. Coad

Deep-sea Spiny Eels or Tapirfishes are found worldwide in the deep sea in both the tropics and the polar seas. The name “Tapirfishes” was coined to avoid confusion with an unrelated family of freshwater fishes called Spiny Eels (Mastacembelidae). The addition of “Deep-sea” to the family renders this unnecessary in English, although “tapir” is retained in French names in the American Fisheries Society list (see the bibliography). There are 10 species, five in Canada with two entering eastern Arctic waters. These fishes attain 1.5 m in total length, although the leptocephalus larva is larger, to 2.0 m. They are characterized by having an elongate body as in eels, but pelvic fins are present (sometimes with three spine-like rays); a relatively small, inferior mouth bearing teeth in a single, comblike row; and up to 40 isolated spines on the back, but no obvious soft rays. A true caudal fin is absent, but the anal fin may regenerate around a broken-off tail tip. The anal fin is long, from the anus to the tail tip, with anterior rays spinous. Scales are small and cycloid and cover both the body and the head. The lateral line is high on the flank anteriorly. Some species have light organs. The overall colour is light grey to dark brown, with the mouth lining and the branchial chamber black. Deep-sea Spiny Eels are found at depths of 125–4,900 m, where they are benthopelagic. Their food is crustaceans, echinoderms, polychaetes, coelenterates, bryozoans, hydrozoans, and other items in and on the bottom. Some feed by cropping and browsing. Food is detected with the aid of well-developed olfactory organs. Some species have many more females than males in a population. Breeding males have darkened anterior nostrils. The eggs are released into the abdominal cavity and laid through two abdominal pores near the anus. The young are leptocephalus larvae, often very large (1.84 m, the largest in the Animal Kingdom). They have no commercial importance.

sources:  McDowell (1973); Smith (1989e).



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Notacanthus chemnitzii Bloch, 1788

Snubnosed Spiny Eel, tapir à grandes écailles

Notacanthus phasgonorus Goode, 1881, described from the Newfoundland Grand Banks is a synonym. Notocanthus nasus Bloch, 1795, described from Iceland and often used in Atlantic Canada waters, is also a synonym.

description:  This fish is identified by a dorsal fin composed only of 5–15 spines, usually 10–11, originating far behind the pectoral fin level. The anal fin has 13–21 spines and 125–145 soft rays; the pectoral fin has 10–17 rays; and the pelvic fin has a fulcral spine, 2–4 fin spines, and 6–11 soft rays. Gill rakers number 12–17, and vertebrae 225–239. The lips and the underside of the snout are covered by dense papillae. The colour is grey to blue grey, becoming dark brown overall with age. The fins are blackish. The peritoneum and pharynx are black in adults, giving a bluish tinge externally when seen through the skin. It attains more than 135.0 cm in total length.

Notacanthus chemnitzii

common names:  A local name is Chemnitz Pigål (Danish/ Greenlandic). Other common names are Cosmopolitan Spinetail, Largescale Tapirfish, Spiny Eel, Swordtail Eel, and poisson-tapir sombre. The French name retains “tapir” as this is used in the AFS list (see the family account). taxonomy:  The genus comes from the Greek noton (back) and

akantha (spine or thorn), referring to the isolated spines on the back. The species is named after Johann Hieronymus Chemnitz (1730–78) a German clergyman and conchologist in Copenhagen who supplied the type-specimen.

habitat:  A benthopelagic species, it is found just above the bottom, usually from 128 m to 1,000 m, down as far as 3,285 m. This species does not appear to school. In Arctic waters it can be found at shallower depths such as 451 m. In Davis Strait it is reported from 500–1,463 m and 0.5°C–4.3°C. biology:  Females are larger than males and have been found with eggs off Iceland in autumn. Food is principally large pink sea anemones thought to be eaten by the fish browsing head down.

importance:  It is not economically important. distribution:  It is found in Baffin Bay and Davis Strait, and worldwide in cooler waters including southwest and southeast Greenland, and south to the Gulf of Mexico in the western North Atlantic Ocean.

Notacanthus chemnitzii

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Polyacanthonotus rissoanus (de Filippi and Vérany, 1857)

Shortspine Tapirfish, tapir à petites épines

common names:  A local name is Savrygget Pigål (Danish/ Greenlandic). Other common names are Shortnose Spiny Eel, Smallmouth Spiny Eel, and Spiny Eel. This species does not appear in the AFS list, and common names using “Tapirfish” and “tapir” are retained. taxonomy:  The genus comes from the Greek poly (a lot of), akantha (spine or thorn), and noton (back), in reference to the numerous dorsal fin spines. The species name comes from the Latin anus (belonging to) and was described from a specimen in the collection of Guiseppe Antonio Risso, called Antoine Risso (1777–1845), author of Ichthyologie de Nice. Notacanthus rostratus Collett, 1889, described from off Newfoundland, and occasionally reported under this name and in the genus Macdonaldia from Davis Strait, is probably a synonym. Distribution of Notacanthus chemnitzii

sources:  Kotthaus & Krefft (1957); Karrer (1973); Jørgensen et al.

(2005).

description:  This species is identified by the dorsal fin comprising solely 26–36 spines that originate anterior to the pectoral fin insertion. The anal fin has 37–42 spines and over 150 soft rays, the pectoral fin has a fulcral spine and 10–16 rays, and the pelvic fin has a fulcral spine and 7–11 rays. There are 22–28 gill rakers on the anterior arch. The mouth ends anterior to the eye. Mature males have black, tubular, anterior nostrils and enlarged posterior nostrils. The colour of the body is light grey to off-white or tan; the anal fin soft rays are dark brown; the opercle margin and posterior part of the mouth are black; and the opercle and the underside of the head and the mouth anteriorly are bluish. The species attains 60.3 cm in total length.

Polyacanthonotus rissoanus



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habitat:  A bathydemersal species, it is found near the bottom at

540–2,875 m, in cooler waters of about 3°C–11°C. In the Davis Strait it is caught at 815–1,470 m and 3.0°C–3.1°C.

biology:  Its food is small bottom crustaceans, particularly amph-

ipods and mysids, and also polychaete worms and coelenterates.

importance:  It is not economically important.

Family Synaphobranchidae Cutthroat Eels, Anguilles égorgées

Brian W. Coad

distribution:  It is found in Davis Strait off southern Baffin Island, and from southwest Greenland south to North Carolina in the western Atlantic, in the Atlantic Ocean generally, and the Mediterranean Sea.

Distribution of Polyacanthonotus rissoanus

sources:  Karrer (1973); Sulak, Crabtree, & Hureau (1984); Crabtree, Sulak, & Musick (1985).

208

This family contains about 29 species, usually in the depths of temperate and warmer oceans. Four species are found in Canada, including two in eastern Arctic waters. Maximum length is about 1 m. All family members have a larva with telescopic or tubular eyes. Adults have one gill opening, or two gill openings close together on the throat region, giving the family its name. Otherwise adults are quite variable. The eyes may be well developed or reduced. Scales and pectoral fins are present or absent. The pelvic fins are always absent. The snout is blunt or pointed. A lateral line is present but may be very short. The mouth may be large or small, and teeth may be needle-like, incisors, or relatively elongate. The dorsal and anal fins are confluent with the caudal fin. Scales are often arranged in a basket-work pattern. The rear nostril is close to the front of the eye, and the anterior nostril is near the snout tip and is tubular. The overall colour is usually dark brown, grey, or black. Cutthroats are bottom-living eels found between 100 m and 4,800 m. Most are predators on fishes or scavengers, even burrowing into carcasses. They have no commercial importance.

sources:  Merrett & Saldanha (1985); Robins & Robins (1989); Sulak & Shcherbachev (1997).

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habitat:  This fish is bathydemersal and has been caught as deep

Simenchelys parasitica

as 3,000 m and as shallow as 100 m. The Arctic specimen was caught at about 963 m. Water temperatures generally are 3.5°C–9.2°C.

Gill, 1879

Snubnose Eel, anguille à nez court

biology:  This species was thought to be parasitic on other fishes

because it has been found burrowing into them, but it is mainly a scavenger, rasping and twisting flesh off moribund or dead fishes. The strongly muscled, blunt, and stout head aids in this process. Invertebrates are also eaten. It produces large quantities of slime or mucus from the skin that literally drips off a captured specimen. Eggs are pelagic and at least 2 mm in diameter.

importance:  It is not economically important. distribution:  This species has been found in northern and southern Davis Strait as two records from cruise data (69.37° N, 65.04° W, and 60.42167° N, 61.24° W), and worldwide including Atlantic Canada.

Simenchelys parasitica

common names:  Other common names are Pugnose Eel, Slime-eel, Snubnosed Eel, and Snubnose Parasitic Eel.

taxonomy:  The genus comes from the Greek simos (flattening or pug-nosed), in reference to the snout shape, and enchelys (eel). The species name comes from the Latin parasitus (parasite) and -ica (belonging to), meaning parasitic, as Gill observed the fish burrowing into the flesh of a halibut. description:  The blunt head and small, circular, and terminal

mouth with strongly plicate lips are distinctive. The tiny mouth is well in advance of the eye level. Elongate scales are deeply embedded in the skin and are arranged in a basket-weave pattern. Juveniles lack scales. There are no teeth on the roof of the mouth, and the jaw teeth are broad, few in number, and in a single cutting row. Pectoral fins are present. The gill openings are small, oblique slits on the throat. Nostril pores are large and evident. The lateral line has 44 pre-anal pores and 2 pre-pectoral pores. The overall colour is grey to grey brown or dark brown, with dark or white fin margins. The lateral line is dark but may be lighter than surrounding tissue in preserved specimens. This species reaches 61.0 cm in total length.

Distribution of Simenchelys parasitica

sources:  See the family sources and the bibliography.

Simenchelys parasitica

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Synaphobranchus kaupii

description:  This species is distinguished by the parallel, slitlike gill openings that are close together on the throat. The head is relatively long compared to that of other eels, more than 10% of the total length. The dorsal and anal fin-rays number about 270. Pectoral fins are present with 14–20 rays. Scales are minute, rod-shaped (3–4 times long as wide), and embedded. The scales are arranged in groups of 4–5 set at right angles to other such groups. There are about 29–35 pre-anal pores and 8 pre-pectoral pores in a distinct lateral line. Small, sharp, recurved teeth occur in bands in the jaws and as a single row on the roof of the mouth, zigzagging anteriorly. The anterior nostril is large and tube-shaped. The posterior nostril is a raised pore near the eye. The anus is at the anterior third of the body, in advance of the level of the dorsal fin origin. The overall colour is black or dark purple grey to brown. The anterior edges of the dorsal and anal fins are lighter than the posterior edges. The mouth cavity is blue black. The species reaches 81.3 cm in total length, perhaps 1.0 m.

common names:  A local common name is Spidssnudet

habitat:  A bathydemersal species, it is found at 131–2,869 m, pos-

Synaphobranchus kaupii Johnson, 1862

Northern Cutthroat Eel, anguille égorgée bécue

Dybhavsål (Danish/Greenlandic). Other common names are Gray’s Cutthroat, Kaup’s Arrowtooth Eel, Longnose Eel, Slatjaw Cutthroat Eel, and anguille égorgée de Gray.

taxonomy:  The genus comes from the Greek synaphes (united) and branchia (gill). The species is named after Johann Jakob Kaup (1803–73), a German researcher who catalogued the apodal fishes in the British Museum.

sibly to 4,800 m. In the Davis Strait, bottom temperatures are 3.12°C to 3.52°C, elsewhere from −1.1°C to 9.8°C. In the southern Davis Strait, 31 collections ranged from 477 m to 1,463 m. In the Davis Strait generally, its depths were 551.5–1,457.5 m, and temperatures 2.1°C–4.6ºC. This eel is a dominant element of the deep-sea bottom fauna and comprises about 40% of the fish by weight and number at 500–1,500 m in the eastern North Atlantic. It is closely associated with Macrourus berglax, Antimora rostrata, and Coryphaenoides

Synaphobranchus kaupii, ventral view showing gill slits

Synaphobranchus kaupii

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rupestris in low latitude–high temperature areas of Davis Strait. From a submersible it has been observed swimming along or above the bottom and hovering head up or down. It can swim backwards as well as forwards.

biology:  Its food is crustaceans, fishes, and large cephalopods,

and it is also known to eat octopus eggs. Roughhead Grenadiers eat this species in the Davis Strait, and Spiny Dogfish are also known predators. Larger squids may also eat this species, as the fish have been found with circular marks on their body. Spawning may peak in spring and summer, but data are contradictory with ripe fish found almost year round. Orange eggs are about 1 mm in diameter. The leptocephalus larva stage may last 18–22 months.

Family Nemichthyidae Snipe Eels, Poissons-avocettes

Brian W. Coad

importance:  This species has been caught on bottom long lines and in baited traps.

distribution:  It is found in Davis Strait, and worldwide including southwest and southeast Greenland and south to the Bahamas in the western North Atlantic Ocean.

Distribution of Synaphobranchus kaupii

sources:  Bruun (1937); Castle (1964); Karrer (1973); Jørgensen (1996); Chambers & Dick (2007).

Snipe Eels or Threadtail Snipe Eels are found in deep waters of the Atlantic, Indian, and Pacific Oceans. There are about nine species, with four species being in Canada including one in Arctic waters. The maximum length is about 1.5 m. These eels are remarkable for their extremely elongate bodies, with over 750 vertebrae in some species, and the unusual jaws. The upper and lower jaws diverge. The two jaws do not meet at the end except in ripe males (and probably, to some extent, females), which lose their teeth and undergo a shortening of the jaw. The name “Snipe” is taken from the shore bird that has a similar beak. The dorsal, anal, and caudal fins are continuous. The end of the body is often lost in captured specimens because it is so thin and the vertebrae so weakly developed. The eyes are large, and pectoral fins are present. The anus is far forward, near the pectoral fins. There is no preopercle bone on the side of the head, and there are no scales. Ripe males and immature fish are so different in appearance that they were often described as different species. In addition to the changes in the male’s jaws, the pectoral fins move posteriorly, the eyes enlarge, tubular anterior nostrils develop, and the fish becomes darker. The overall colour is dark to light brown, pale or even reverse countershaded where the back is lighter than the belly. Snipe Eels are rarely caught inhabitants of deep mid-waters down to 4,300 m. These unusual eels are believed to position themselves vertically, head up or down, in the water column and capture migrating shrimps by entangling their antennae in the diverging jaws. They may also make rapid sideways movements of the head to wedge a prey in the jaws. Some species have an extremely long tail filament. The lateral line extends along this filament, which gives the fish better sensing ability. It may also give a larger echo to predators, which then avoid this apparently large fish. The larva of these eels is a leptocephalus, readily distinguished by its thin, elongate tail. The enlarged, tubular nostrils in males help to locate females by smell at spawning time. Snipe Eels probably spawn once and then die.

source:  Smith & Nielsen (1989).



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Nemichthys scolopaceus Richardson, 1848

Slender Snipe Eel, avocette ruban

taxonomy:  The genus comes from the Greek nema (filament or

thread), referring to the thread-like caudal region, and ichthys (fish). The species name comes from the Latin scolopax (snipe) and –eus (having the quality of), referring to the long bill-like jaws seen in the bird.

description:  This species is distinguished by the snipe-like jaws. It has a caudal filament (a long, thin extension of the tail), five pores in each segment of the lateral line, three pore rows in the lateral line, and no sensory ridges on the head. There are 6–17 postorbital pores, usually more than 10, and 6–13 preopercular pores. The lateral-line pore pattern looks like a five in dominoes, a quincunx. The teeth are fine, sharp, and numerous. The anus lies below the pectoral fins, which have 10–14 rays. Over 750 vertebrae may be present. Both the dorsal and the anal fins have over 320 rays, and the dorsal rays are stiffer than the anal rays. The overall colour is black, dark brown, or grey, with darker brown speckles, the belly often being darker than the back in young, a reversal of the usual pigment pattern. The anal fin and the pectoral fin tips are black. It reaches about 145.5 cm in total length. Nemichthys scolopaceus

common names: A local name is Langhalet Sneppeål (Danish/

Greenlandic). Other common names are Atlantic Snipe Eel, Brittle Watersnake, Glass Snake, Polygon-spotted Snipe Eel, Threadeel, Threadfish, anguille bécasse, long bec, and avocet.

habitat:  This species is bathypelagic and has been taken as shallow as 210 m, south of LaHave and Sable Island banks, and at 30 m off British Columbia. It is occasionally cast on shore by storms but is usually in the depth range of 457–3,656 m. In southern Davis Strait, captures have been made at 820–1,463 m. Leptocephalus larvae are found in shallower water at 90 m but only over the deep ocean.

Nemichthys scolopaceus

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biology:  Its biology is mostly unknown except as described in

the family account. Its food is pelagic shrimps. Spawning in the Sargasso Sea is protracted and may occur in both winter and summer. Goosefish and Pollock eat this species in more southerly waters.

importance:  It is not economically important. distribution:  This species is found in Davis Strait, southeast Greenland, and all temperate to tropical oceans including off both the Atlantic and the Pacific coasts of Canada.

Family Serrivomeridae Sawpalates, Serrivomers

Brian W. Coad

Sawpalates or Sawtooth Eels are usually found in the temperate to tropical Atlantic, Indian, and Pacific Oceans. There are about 10 species, with three in Canada including one from the eastern Arctic. Maximum length is about 78 cm. Sawpalates are characterized by the two or more rows of bladelike or conical teeth in the roof of the mouth that gives them their name. The jaws are elongate, reach beyond the eyes posteriorly, and have minute teeth. The lower jaw projects beyond the upper jaw. The anterior and posterior nostrils are close together, just in front of the eyes. The dorsal fin begins well behind the head, behind the anal fin origin. The dorsal, caudal, and anal fins are continuous. Pectoral fins are rudimentary or absent, and pelvic fins are absent. Gill openings are laterally placed, almost vertical, and confluent on the throat. The anus lies well before the middle of the body length. There are no scales, and lateral-line pores are minute. The tail is thread-like. Sexually mature fish develop the double sawtooth palate, and maxillary and mandibular jaw teeth are reduced to a single and irregular row. The eyes enlarge as do the nostrils, although not as noticeably, presumably to aid in locating a mate. The overall colour is dark brown or black, with an iridescent silvery to bronze layer, which is usually lost when the fish is captured in nets. These are fragile fishes, easily damaged, and are usually caught in good condition only by scientific research vessels using special gear. They are bathypelagic to abyssopelagic, swimming in the ocean depths down to 6,000 m, but they may migrate to surface waters. They can be common in mid-waters. Larvae and juveniles are found near the surface, and hatching is known to occur in the Sargasso Sea for Atlantic species. Larvae are transparent leptocephali, typical of eels, and common in surface waters of the North Atlantic. This family has no commercial importance.

Distribution of Nemichthys scolopaceus

source:  Bossé (1991).

source:  Tighe (1989).



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Serrivomer beanii Gill and Ryder, 1883

Stout Sawpalate, serrivomer trapu

This species has 136–175 dorsal fin-rays, 119–156 anal fin-rays, 5–6 caudal fin-rays, and 7 (rarely 8) branchiostegal rays. Pectoral fin-rays number 6–7. The dorsal and anal fins are delicate. The dorsal fin origin is behind the anal fin origin. The anal fin origin is about a head length behind the gill opening. The overall colour is brown to black, with the head being black or blue black, but the skin is often lost or damaged in captured specimens. The ventral surface is darker than the dorsal surface. There is an overlying silvery iridescent layer in the skin. The species reaches 81.0 cm in total length.

habitat:  It is found in epipelagic and abyssopelagic waters from

0 to 3,000 m, and even as deep as 5,998 m, where they are apparently solitary and fast swimmers. Occasionally they may be found in groups of four to five fish. In the mid–North Atlantic they are found mostly between 450 m and 1,000 m at 8°C–12°C. This species has been caught in Davis Strait at 446–1,472 m and at 739–1,458 m and 3.3°C–3.8ºC. Leptocephali or larvae occur at 0–300 m, mostly between 50 m and 70 m, reaching the surface at night. Serrivomer beanii

common names: A local common name is Beans Næbål (Dan-

ish/Greenlandic). Other common names are Bean’s Sawtooth Eel, Longfin Sawpalate, Stout Sawpalate Eel, and serrivomer à nageoire longue.

taxonomy:  The genus comes from the Latin serra (saw) and vomer (ploughshare or vomer), in reference to characteristic teeth on the mouth roof. The species was named after Tarleton Hoffman Bean (1846–1916), a U.S. ichthyologist and co-author of Oceanic Ichthyology: A Treatise on the Deep-Sea and Pelagic Fishes of the World (1896). Some records were of Serrivomer parabeani Bertin, 1940, now regarded as a synonym.

biology:  Its food is principally large pelagic shrimps, euphausiids, and other crustaceans, along with small fishes. Atlantic Cod are known to eat Sawpalates. Spawning off Bermuda occurs in late spring and summer, and ripe females have been caught in August. Leptocephalus larvae as small as 17 mm long have been captured, but reach 63 mm before metamorphosis, and adolescents in transition to adults are about 96 mm or longer. Metamorphosis can be complete at 90 mm. importance:  It is not economically important. distribution:  It is found in Davis Strait and eastern Hudson Strait, southwest and southeast Greenland, and the Atlantic, Indian and Pacific Oceans.

description:  The characteristic blade- or saw-like teeth on the

roof of the mouth, numbering 50–100 and increasing in number with age, distinguish this species.

Serrivomer beanii

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Family Saccopharyngidae Swallowers, Avaleurs

Brian W. Coad

Distribution of Serrivomer beanii

sources:  Karrer (1973); van Utrecht (1982); van Utrecht & van Utrecht-Cock (1987); Fishelson (1994); Neat & Campbell (2013).

The Family Saccopharyngidae is found in the Atlantic, Indian, and Pacific Oceans. There are about 10 species, including one reported from the Atlantic and Arctic coasts of Canada. The maximum length is about 2 m. Swallowers, Swallower Eels, or Whiptail Gulpers are highly unusual, lacking various skeletal elements such as opercular bones, ribs, and branchiostegal rays, and also scales, pelvic fins, pyloric caeca, and a gas bladder. The caudal fin is absent or rudimentary, but the tail ends in an expanded caudal organ that varies in structure and may be luminous. The eyes are tiny and placed far anterior. The most obvious feature is the very large mouth and distensible pharynx and abdomen that can take in very large fish. The jaws have curved teeth. Mature males develop enlarged eyes and olfactory organs, and the feeding apparatus is reduced. There may be luminous grooves on the body and the tail. The colour is black overall. Swallowers have a leptocephalus larva, which relates them to the eels with which they are sometimes classified. These fishes are rarely seen and are known from less than 100 specimens.

sources:  Bertelsen, Nielsen, & Smith (1989); Smith (1989b).



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Saccopharynx ampullaceus

greatly enlarged olfactory organs, and the abdominal pouch is reduced, probably because feeding does not occur.

Taillight Gulper, avaleur feu-arrière

importance:  It is not economically important.

(Harwood, 1827)

common names: A local name is Almindelig Slugål (Danish/

Greenlandic). Other common names are Gulper, Gulper Eel, and Pelican-fish.

distribution:  It has been found in Davis Strait as two records from cruise data at 63.8717° N, 58.26° W, and 62.629° N, 59.6756° W, and also from southwest and southeast Greenland. It is found in the North Atlantic Ocean between about 10° and 65° N.

taxonomy:  The genus comes from the Greek saccos (sack)

and pharynx (pharynx), in reference to the wide throat. The species name comes from the Latin ampulla (flask, bottle) and -aceus (resembling), in reference to the body being expandable. The holotype was found floating at the surface in Davis Strait at about 62° N, 57° W, in the fall of 1826.

description:  The enormous mouth and the small eyes near the snout tip are characteristic, in combination with the gill openings being closer to the snout tip than to the anus (in contrast to the related Pelican Gulper). The pectoral fins are well developed (minute in Pelican Gulper), the jaw teeth are curved, and the dorsal fin origin is well behind the head. The body and the tail have filaments of varying length sticking out dorso-laterally. The colour is black overall. The caudal organ has spots of luminescent tissue but no well-developed filaments. There is a white line or groove on each side of the dorsal fin. The species attains about 1.61 m in total length, although much of this is the tail. habitat:  It is found bathypelagically down to 3,000 m or more

but may approach the surface. Its food is fishes, some larger than itself. The leptocephalus larva occurs in shallower water. The species has been caught at 1,099–1,429 m in Davis Strait.

biology:  Sexually mature fish lose their teeth, and their jaws

Distribution of Saccopharynx ampullaceus

sources:  See the family sources and the bibliography.

shrink, as bones resorb or disintegrate. Mature males develop

Saccopharynx ampullaceus, underside of body with filaments (bottom left), and open mouth (bottom right)

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Eurypharynx pelecanoides

Family Eurypharyngidae

Vaillant, 1882

Pelican Gulper, grandgousier pélican

Gulpers, Grandgousiers common names:  A local name is Pelikanål (Danish/Greenlandic). Other common names are Big-mouth Gulper, Deep-sea Gulper Eel, Gulper Eel, Pelican Eel, Pelican Fish, and Umbrellamouth Gulper.

Brian W. Coad

taxonomy:  The genus comes from the Greek eurys (broad, wide) and pharynx (pharynx), in reference to the large mouth and throat. The species name comes from the Latin pelicanus (pelican) and -oid (like), again in reference to the mouth and throat. Gastromus bairdii Gill and Ryder, 1883, described from off New England is a synonym. Gulpers or Gulper Eels are found in the Atlantic, Indian, and Pacific Oceans and on the Atlantic and eastern Arctic coasts of Canada. There is only one species in the family. This family is highly unusual, lacking various skeletal elements and also the scales, pelvic fins, pyloric caeca, and a gas bladder. The lateral line has elevated tubules. The caudal fin is absent or rudimentary. The eyes are tiny and placed far anterior. The most obvious feature is the enormous mouth. Gill openings are small, and this is the only teleost fish with five gill arches and six visceral clefts. The Pelican Gulper (Eurypharynx pelecanoides) has a leptocephalus larva that relates it to the eels with which it is sometimes classified. It is not commercially important.

source:  Bertelsen, Nielsen, & Smith (1989).

description:  The enormous mouth and the small eyes near

the snout tip are characteristic, in combination with the gill openings being closer to the anus than to the snout tip (in contrast to the related Taillight Gulper). The body is quite fragile, and specimens brought up in nets are often damaged. Although the mouth is large, the stomach is not markedly distensible as in the Taillight Gulper. The jaw teeth are minute. The pectoral fins are also minute (well developed in the Taillight Gulper). The dorsal fin originates on the head. The tail has a luminescent organ at the tip. Mature males have a large, circular, knob-like, nasal rosette; the jaws degenerate; and teeth are reduced or lost. The colour is black overall with a white groove on each side of the dorsal fin – of unknown function as are the grooves in the Taillight Gulper. The caudal organ is black dorsally and white ventrally. The species rarely exceeds 100.0 cm in total length.

Eurypharynx pelecanoides



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habitat:  This bathypelagic species is found in the deep sea at 500–7,625 m, and perhaps deeper, and is quite common. A Canadian Arctic specimen was taken in Davis Strait at 1,136–1,154 m near the sea bed at 4°C, and others were taken at 758–1,144 m.

biology:  The leptocephalus phase lasts up to two years, and these

larvae are found at 100–200 m. The food is plankton, crustaceans, squid, and fishes that may be attracted to the huge open mouth by the species dangling its luminescent caudal fin tip in front of it or even swimming in a circle. It is unlikely that prey is chased, because most prey are swallowed head first, and the Pelican Gulper is a poor swimmer. This fish probably lunges at its prey with its mouth closed. The mouth is opened very close to the prey, and the huge gape is unfolded like a parachute by the pressure of the water. The prey is engulfed in a pool of water, and this water is gradually removed through the narrow gill openings, enabling the prey to be swallowed. Since the dentition is weak, the mouth structure delicate, and the stomach not very distensible, large food items are probably not taken. The Pelican Gulper is eaten by various other fishes. Males probably use their large nasal organs to detect the pheromones emitted by females.

importance:  It is not economically important. distribution:  It has been found as five records in the Davis Strait off southern Baffin Island as cruise data and ARC 8704123, in southwest and southeast Greenland, and in all oceans.

Distribution of Eurypharynx pelecanoides Eurypharynx pelecanoides

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sources:  See the family sources and the bibliography.

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this species at such localities as Aklavik, Inuvik, Tuktoyaktuk, and Paulatuk in the years 1987 to 1990. The numbers caught range from none up to 6,075 fish.

Family Clupeidae

sources:  Svetovidov (1952); Hildebrand (1963); Messieh (1980); Grande (1985); Whitehead (1985); Grant (1986); Domanico, Phillips, & Schweigert (1996).

Herrings, Harengs

Brian W. Coad

Clupea harengus Linnaeus, 1758

Atlantic Herring, hareng atlantique

Herrings and related species are found worldwide, usually in warmer marine waters, with some species being anadromous or permanent freshwater residents. There are about 180 species, with ten found in Canada, including two in Arctic waters. The maximum length is about 76 cm although many species are smaller, 10–20 cm standard length. These fishes have modified scales on the belly, forming abdominal scutes with a saw-like edge. The lateral line is usually absent or on only a few scales. Silvery cycloid scales are deciduous (easily detached) and are found only on the body. Teeth are small or absent, but the gill rakers are long and numerous for sieving plankton. The fins lack spines, and there is no adipose fin. The pectoral and pelvic fins have a large axillary scale. The caudal fin is deeply forked. Each eye is partly covered by an adipose eyelid. The flesh is particularly oily and is highly nutritious. The overall colour is often dark blue or blue green on the back, with silvery flanks, and often a spot or rows of spots on the anterior flank. Members of this family often form immense schools in surface waters of the ocean where they usually feed on plankton. Herring can feed on smaller plankton at night by filter feeding but during the day can also use particulate feeding, selecting larger plankton by using a specialized part of the retina. Herring are easily caught and are extremely valuable to commercial fisheries. They are the most important fishes economically, as food for both man and many other commercial fish species. An estimated ten billion Atlantic Herring are caught each year, and in one year members of the Herring Family made up 37.3% of all fish caught in the world. Some are used for fish-meal, fertiliser, and bait and as an oil source. Baillie Islands Inuit claimed that Pacific Herring were never caught before the white men came in the 1920s, but the schools came in rarely and then not often close inshore, and the local people did not use long seines. Pacific Herring have been harvested in the western Arctic, but catches are limited; for example, from two to six hunters caught



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Clupea harengus

common names:  Local names are Kavisilak (Inuktitut) and

Ammassassuaa and Kapisilik (Greenlandic). Kavisilak and Kapisilik have also been used in Canada and Greenland for several salmonids. Other common names are Bank Herring, Fall Herring, Labrador Herring, Sardine, Sea Herring, Shore Herring, Spring Herring, and Summer Herring.

taxonomy:  The genus comes from the Latin clupea (herring or sardine), derived from clupeus (shield), presumably in reference to the scales. The species is named from the low Latin harengus (herring) for a fish that swims in armies or schools. description:  Distribution in the eastern Canadian Arctic is the easiest way to distinguish this species as other characters overlap with its western relative. This species has a sharp edge to the belly scales (but not a saw-toothed edge), and teeth on the roof of the mouth. Dorsal fin-rays number 13–22, anal fin-rays 12–23, pectoral fin-rays 14–22, and pelvic fin-rays 8–10, usually 9. Scales before the pelvic fins on the belly number 26–33, behind 11–17, and are more or less well developed in both areas. Gill rakers number 40–49 (fewer in fish smaller than 10 cm). Vertebrae number 51–60, mostly 55–57. Vomerine teeth are well developed. The back and upper flank are green to green blue or dark blue in colour, and the lower flank and belly are iridescent silver or tinged violet. The operculum may be tinted golden, brown, or brassy, and this colour may extend along the flanks. The pelvic and anal fins are translucent, the dorsal and caudal fins are dark grey with some greenish or bluish tinges, and

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Clupea harengus

the pectoral fins have dark bases and upper edges. The peritoneum is dusky. This species reaches 45.0 cm in standard length and 1.05 kg in weight.

habitat:  This herring is benthopelagic and oceanodromous. It is

found in large schools in inshore waters and offshore from the surface down to 364 m. A large school of 17.5 cm long fish was observed in Mittimatalik (Pond Inlet). As well as moving horizontally in schools, herrings move vertically in the water column, approaching the surface at dusk and descending at dawn. However, they may be seen at the water surface on calm days. On the Scotian shelf the preferred depths are 55–90 m and 146–163 m at temperatures of 1°C and 7°C–9°C. The total temperature range there is 0°C–11°C, although this species has a biological anti-freeze that lowers the freezing point of body fluids below that of sea water (−1.9°C) to about −2.2°C. It is an irregular visitor to estuaries in James Bay.

biology:  The size of the diet items consumed increases with the

size of the herring. Larval fish are an important item of the diet, as are various planktonic crustaceans and molluscs. Herrings filter feed at low light intensities and when prey items are small, but they pick out individual food items when light levels are better and such prey is available. Herrings and their eggs are eaten by many other fishes, birds, and marine mammals including Killer Whales. Most herrings mature by age five, although some are mature at three years. The lifespan is 25 years. The spawning time varies between stocks from April to November in Canada. In Davis Strait at the Greenland coast, spawning occurs from August to September. Earlier spawning occurs in northern coastal waters, and later spawning offshore in southern deep water. The timing of herring arrival on spawning grounds

220

depends on temperature, but temperature at spawning ranges from 0°C in spring to 20°C in the autumn in high latitudes. Eggs settle and adhere to the bottom in ribbons, are 1.4 mm in diameter, and number up to 261,000. Males are said to release ribbons of milt 1–2 cm long prior to the release of eggs, which form layers of 1–15 eggs deep and would not be fertilized if laid first. Layers deeper than four eggs have high mortality. Females visually select spawning sites in shallow water, preferring rougher surfaces. Most spawning occurs in areas of mixing where high levels of primary production, and hence food, occur for larval growth. Spring spawners have fewer but larger eggs than do autumn spawners. There are also differences in length and age composition of the stocks, year-class strengths, growth rates, fin-ray and gill raker counts, and ear-stone structure. Although they spawn separately, the stocks mix on feeding and wintering grounds.

importance: Canadian landings have been as high as 528,000 t, in 1968, but were 242,340 t in 1988, second in weight to Atlantic Cod. The Canadian Atlantic catch in 2011 was 134,301 t, worth $38,384,000, second in importance after Greenland Halibut. However, it is not common enough to be commercially important in Arctic waters. distribution:  The Atlantic Herring is found in Davis Strait; the mouth of Frobisher Bay; Hudson Strait; Ungava, Hudson, and James Bays; southwest and southeast Greenland; and south to South Carolina in the western North Atlantic Ocean. It is also found in the eastern North Atlantic Ocean. Records from the Hudson–James Bay region have been termed “adventitious” because they are outside the normal range and may be a relict population. The record by Herlinveaux (1970) from near Mittimatalik

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(Pond Inlet) on northern Baffin Island was from a diver observation of a large “herring” school with fish 17.5 cm long and needs confirmation.

indicate that they are distinct species. Herring entered the Pacific Ocean about 3.0–3.5 million years ago in the mid-Pliocene after the Bering Strait had opened. The gradual cooling of the Arctic Ocean since then has eliminated the continuous distribution from the Pacific across the Arctic to the Atlantic Ocean.

description:  Distribution in the western Canadian Arctic is the easiest way to distinguish this species as other characters overlap with its eastern relative. Dorsal fin-rays number 13–21, anal fin-rays 12–23, and pectoral fin-rays about 17–19. There are 38–54 scale rows along the side, and 10–14 keeled scales between the pelvic and anal fins. Keeled scales are developed only posterior to the pelvic fins. Vertebrae number 46–58, mostly 52–55. Vomerine teeth are less well developed than in its relative. The back and upper flank is blue green to olive in colour, and the flanks and belly are silvery. The peritoneum is black. This species attains 46.0 cm in total length. habitat:  This species is pelagic and neritic. Adults migrate

Distribution of Clupea harengus

sources:  Davidson (1932); Dunbar (1970a); Herlinveaux (1970); Friesen & Nelson (1978); Hudon (1988); Ewart & Fletcher (1990); D.B. Stewart, Bernier, & Dunbar (1991); B. Morin, Hudon, & Whoriskey (1992); D.B. Stewart, Dunbar, & Bernier (1993); Canada, Ministère des Pêches et Océans (1996a, 1996b); Woehrmann (1997); S.H. Ferguson, Loseto, & Mallory (2010).

Clupea pallasii

Valenciennes, 1847 Pacific Herring, hareng du Pacifique

common names: Local names are Iituuq, Kavisilaq Krolleliprark, Pikoaktik, Piqquaqtitaq (Inuktitut); Pirkroartitak, Qaaktaq, Qaantaq, Qaluhaq (Inuvialuktun); and Uqsruqtuuq. Other common names are Blue Back, Blue Herring, North Pacific Herring, and Oriental Herring. taxonomy:  The species was named after Peter Simon Pallas (1741–1811), a German naturalist who worked in Russia, author of Zoographia rosso-asiatica. This herring was regarded as a subspecies of the Atlantic Herring, but biochemical and genetic studies along with life history data



UTP Fishes Book 5pp04.indb 221

to the surface in the evening and to deeper water at dawn. It is reported to be at the surface and possibly down to 475 m, but mostly at depths of less than 150 m. These fish are found in nearshore areas in the Beaufort Sea; and in harbours, inlets, bays, and brackish estuaries when spawning; and may enter fresh water. Eggs in the Fingers area of Liverpool Bay incubate for about 24 days at 10°C–12°C and 6‰–12‰. Adults disperse along the coast to feed after spawning, returning to overwintering sites in late August. The Beaufort Sea coast has temperatures of 2.5°C–14.4°C, depths of 0.7–9.0 m, and salinity of 0.04‰–29.4‰ from June to September. They seek deeper water in the autumn, leaving the Tuktoyaktuk Peninsula. Temperatures as low as −1.4°C are tolerated, as this herring has anti-freeze proteins like many other polar and subpolar fishes do. Migratory runs are influenced by wind-induced tides and the resultant influx of saline water, such as in Tuktoyaktuk Harbour. Large schools have been reported. One between the Booth Islands and Parry Peninsula was 13 m deep and 2.6 km long, and one in eastern Franklin Bay was 12.8 m deep and about 3.2 km long. The Fingers area of Liverpool Bay in the Beaufort Sea is a major spawning and overwintering site, as is the area northeast of Richards Island and Mason Bay. McKinley Bay and the Baillie Islands area may also be spawning areas. Heavy runs of herring are found at the head of Darnley Bay from early September until freeze-up. Herring are reportedly more abundant to the east of the Mackenzie River Delta than to the west, particularly along the Tuktoyaktuk Peninsula and in Liverpool Bay. It is the dominant species in Franklin and Darnley Bays of Amundsen Gulf, and the most abundant fish collected in the Hans Bay region of the Eskimo Lakes. They are particularly abundant in Tuktoyaktuk Harbour, a major overwintering area. In surface gill-net catches near Tuktoyaktuk, this species accounted for 32% of catches. On 3 August 1911 a 200 foot sweep-net run out from shore with a dory at the Cape Bathurst sand-spit caught about 3,000 fish (about 13 barrels). The Arctic warming trend made their appearance at Sachs Harbour more common in the 1990s.

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Clupea pallasii

biology:  Adult Pacific Herring in the Beaufort Sea eat crus-

taceans such as mysids, copepods, ostracods, and amphipods; benthic infauna such as pelecypods, Acarina, polychaete worms, and chironomids; foraminiferans; plant material; and a variety of fishes rather than plankton as elsewhere. In Tuktoyaktuk Harbour copepods predominate, followed by mysids and amphipods. Younger fish eat smaller crustaceans and fish larvae. Herring are of great importance as food for other fishes such as Inconnu in the Beaufort Sea, seabirds, seals, and whales. Glaucous Gulls eat herring in the Beaufort Sea. Herring eggs are eaten by a wide variety of predators, particularly gulls and crabs. Arctic Lampreys have been found attached to this species near Tuktoyaktuk. Herring from Tuktoyaktuk and Eskimo Lakes mature in their sixth to eighth year, about three to five years later than in the North Pacific (Strait of Georgia). In the Alaskan Bering Sea, fish aged seven to nine dominate the spawning population, varying with the year. The lifespan may be 19 years. Beaufort Sea fish grow more rapidly in length, but not in weight, than do Pacific Ocean fish, accumulate surplus energy more slowly as predation pressure is lower, have a lower adult mortality rate, are in poorer condition as adults, and have a smaller size-specific ripe-egg weight to anticipate greater larval mortality because of warmer sea temperatures. Spawning is preceded by an inshore migration to shallow bays in the autumn when feeding stops after a summer spent fattening offshore or in nearshore areas like the Tuktoyaktuk Peninsula. The winter is spent fasting and in ripening sexual products using the energy from stored oil. Strait of Georgia fish have ovary weights that are 2.1 times larger than those of Beaufort Sea fish, they spawn at a smaller size and a younger age, and they have a higher fecundity, these all being effects of latitude and exploitation.

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In Arctic waters of the Mackenzie River estuary, Tuktoyaktuk Harbour, Liverpool Bay, and the Beaufort Sea generally, spawning is variably reported for May–June and June–July under the ice or at spring break-up and is associated with macrophyte growths. Spawning in the harbour is earlier than in the bay. The mean age of these fish is 11.0–12.6 years, and the oldest fish are 16 years. Spawning temperatures are 2.5°C–4.0°C; more generally in the Canadian Arctic the spawning temperature range is 0°C–5°C. The mean lengths of samples in these areas are 265–296 mm, ranging from 210 mm to 320 mm, and mean round weights are 233–306 g. As many as 134,000 eggs are produced, with a diameter of 1.2–1.5 mm, when exposed to sea water. Average fecundity is usually below 50,000 eggs, and eggs per gram range from 116.5 to 151.1, well below the 200 eggs per gram of British Columbia herring. An estimated 8.2 t of herring deposited 568 million eggs in the Fingers area of Liverpool Bay from 12 June to 16 July 1985. The eggs adhere to seaweeds, such as red algae and sea grasses, or rocky substrate but may be arranged in rows on the sea bed as the females extrude the eggs, aided by contact with the substrate. Spawning depth in the Fingers area is 1.0–4.5 m. Egg densities can exceed 4 million per square meter, but egg mortality is high. The herring larvae and juveniles develop and remain in schools along the Tuktoyaktuk Peninsula from mid-July to September. The coastal larval-fish assemblage in the Canadian Beaufort Sea is dominated by this species, an area greatly influenced by the Mackenzie River outflow, defined as a shallow intense plume. High levels of cadmium in livers and relatively high levels of toxaphene and hexachlorocyclohexanes (HCH) in muscle tissues have been found in Pacific Herring at Tuktoyaktuk, measuring 0.86 μg/g, 74 ng/g, and 12.5 ng/g total HCH, respectively. However, toxaphene

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and HCH levels are 10–100 times less than they are on the east coast of Canada and in the Pacific Northwest.

importance:  This species has great commercial importance in

British Columbia, second only to the salmons. The value of flesh in 1985 was $2.23/kg and of roe $30.31/kg. The catch in 2011 was 7,751 t, worth $2,417,000. An experimental shore-based gill-net fishery near Baillie Island (Cape Bathurst) in the Beaufort Sea in 1963 failed because of the high cost of transportation and processing. About 8,000 kg was harvested. As a commercial target in the Beaufort Sea, allowance would have to be made for Beluga feeding. Roe rather than flesh may be a potential product in Arctic waters because of its higher value. Attempts at a roe fishery were made in Tuktoyaktuk Harbour in 1981 and 1982 and at Liverpool Bay from 1981 to 1983, but the spawning stock biomass was too low and too variable year to year to support it. An experimental roe fishery in the Fingers area in 1983 produced 398 kg of marketable roe from 4,580 kg of herring. A commercial fishery quota of about 230,000 kg annually was assigned for Cape Parry near Paulatuk, but no lasting fishery has been established. Catches at Paulatuk in 1986–7 totalled 446 fish, and in June–July 1990, 124 fish. Although sizeable stocks have been reported from Liverpool Bay to Darnley Bay in the central Arctic, they are not considered exploitable. The spawning stock biomass in the Fingers area of Liverpool Bay was 2.6–13.8 t, for example, but this is too low to sustain a commercial fishery. These slow-growing stocks are less productive and more easily overharvested than those in British Columbia waters. Additionally, spawning occurs at ice break-up in some areas such as Tuktoyaktuk Harbour, and fishing then is extremely dangerous. The traditional fishery in Tuktoyaktuk Harbour takes place in autumn after ice has formed in the harbour. An autumn fishery in the harbour with sweep nets in the 1950s occurred at the end of September, but in about one year in five the fish did not enter the harbour, and in another year in five freeze-up occurred. The average annual harvest then was 50 tons over a five-year period. Smoked male carcasses from the roe fishery were potentially marketable, but this still left large numbers of carcasses for disposal. A further problem arose in that the herring are food for Lake Trout in the Eskimo Lakes area, and these Lake Trout support a small sport fishery and a subsistence fishery. Herring have been fished for subsistence in the Cape Bathurst area and by residents of Tuktoyaktuk, Inuvik, and Aklavik at coastal points near the Mackenzie Delta. Some have been used as dog food, and some are pickled or fried for the local market. Catches in the Tuktoyaktuk area peaked at about 2,250 fish in September for the 1988–97 period, for example. About ten residents of the Beaufort Sea region caught 1,880 kg of herring in 1987–8. They are a by-catch of the Mackenzie River Delta coregonine fishery and are occasionally dried for food or frozen as a store of dog food.

Chantrey Inlet (Fenco and Slaney, 1978) is probably the easternmost record of C. pallasii but needs verification by voucher material. It is occasionally found as far upriver as Aklavik in the Mackenzie River system. It is also found from the Norwegian, White, and Laptev Seas east to the Alaskan Beaufort Sea; south to Baja California in the eastern North Pacific Ocean, and Korea in the western North Pacific Ocean.

Distribution of Clupea pallasii

sources:  Anonymous (1963); Dunbar (1970); Percy (1975); Hunter (1975, 1981); Fenco & Slaney (1978); Milne & Smiley (1978); Fraker, Griffiths, & Ward (1979); Byers & Kashino (1980); Bond (1982); LGL Limited & Environmental Sciences Limited (1982); Gillman & Kristofferson (1984b); Lawrence, Lacho, & Davies (1984); Kristofferson & Gillman (1986); Shields (1986); Dickins et al. (1987); Crawford (1989); Raymond (1989); Barry & Barry (1990); Fabijan (1991a, 1991b, 1991c); Lacho (1991); Rass & Wheeler (1991); Kerkvliet & Hamner (1993a, 1993b, 1993c, 1994); Tanasichuk, Kristofferson, & Gillman (1993); Inuuvik Community Corporation, Tuktuuyaqtuuq Community Corporation, & Aklarvik Community Corporation (2006); Muir (2010); Laakkonen, Lajus, Strelkov, & Väinölä (2013); Paulic & Papst (2013).

distribution:  The Pacific Herring is found in the eastern

Queen Maud Gulf, Melville Sound, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and the Beaufort Sea and as a single record from western Banks Island. An identification of C. harengus from



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Argentina silus

Family Argentinidae

(Ascanius, 1775)

Atlantic Argentine, grande argentine

Argentines, Argentines

Brian W. Coad

Argentina silus

common names: A local name is Almindelig Guldlaks (DanArgentines or Herring Smelts are found in all oceans but only on the Atlantic Arctic coast of Canada. There are about 20 species worldwide, with three reported from Canada, including one in Arctic waters. The maximum length is about 70 cm. Argentines are silvery (hence “argentine”) fishes of fusiform body shape and fin arrangement. The Canadian Arctic species has an adipose fin; normal, non-tubular, but large eyes; and a small mouth lacking premaxillary and maxillary teeth and not reaching back to the eyes. The dorsal fin origin is in front of the pelvic fins. The pectoral fins are low on the body. Easily detached cycloid scales are present, and the fin-rays are not spiny. There are no light organs. The gas bladder often has a silvery layer. The overall colour is silvery to light yellowish, often iridescent, and there is often a silvery or brownish flank stripe. These fishes are found on the continental shelf and upper slope and are benthopelagic. Its food items are planktonic invertebrates and small fishes. Argentines have pelagic eggs and larvae. With the exception of the species dealt with here, these fishes are not commercially important.

sources:  Cohen (1958, 1964); Cohen & Atsaides (1969).

ish/Greenlandic). Other common names are Greater Argentine, Great Silver Smelt, Herring Smelt, and saumon doré.

taxonomy:  The genus comes from the Latin argentum (silver). The species name comes from the Norwegian word sild for herring.

description:  This species is distinguished by family characters and a combination of characters outlined here. The scales have minute spines and are easily lost on capture. Lateral-line scales number 64–70 and do not extend onto the caudal fin. The mouth is small. An adipose fin is present, and the caudal fin is deeply forked. There are six branchiostegal rays and 9–17 lower arm gill rakers. The dorsal fin origin is above the tip of the pectoral fins, far forward. The pectoral fins are low on the side of the body. The eyes are large. Dorsal fin-rays number 11–13, anal fin-rays 11–17, pectoral fin-rays 13–18, and pelvic fin-rays 11–14. It is reputed to smell like cucumber. In colour this fish is a pale green-yellow, brown, brassy, or olive with silvery flanks and a white belly. The flanks have golden tints. The adipose fin is yellowish. The base of the caudal fin is darkly pigmented. The gas bladder is silvery. This species is reputed to reach 70.0 cm in standard length, although most are under 50.0 cm. habitat:  This species is bathydemersal and oceanodromous and

found in schools at 55–550 m near the ocean bottom, but it has been caught down to 1,440 m. Canadian Arctic records are from 420–842 m. It is abundant off southwestern Nova Scotia where, on the Scotian Shelf, it is found at a temperature range of 4°C–13°C. Young are pelagic at about 400–500 m depths.

biology:  Its food includes crustaceans, squids, arrow worms,

comb jellies, and small fish. It is eaten by Sebastes sp., White and Silver Hakes, Atlantic Cod, Spiny Dogfish, and other fishes in more southerly waters. Atlantic Argentines may mature as early as four years of age and live for 35 years, although in the Davis Strait maturity is reached at 8–10 years. Males and females mature at similar lengths, but males

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Argentina silus

are almost a year older than females. Growth is rapid up to 7–8 years of age. Females eventually reach larger sizes than males. Spawning occurs in March–April on the Scotian Shelf, as late as May elsewhere, and mature fish are caught year round. The oily eggs float in mid-water. Eggs are 3.0–3.5 mm in diameter when free floating and may number up to 38,599.

importance:  The flesh is white and flaky, and this species

has been marketed in Europe. Catches in Canadian waters are by Japanese and former USSR fleets, but catch limits have decreased the take from 49,040 t in 1966. The sustainable yield is probably about 15,000–20,000 t. The silver pigment from the scales and the gas bladder is used to produce “pearl essence” from which artificial pearls are made. It is not commercially important in the Arctic.

distribution:  It is found in temperate to Arctic waters on both sides of the North Atlantic Ocean, including two records at 63.5483º N, 59.9181º W, and 68.0818º N, 59.3532º W, in the Davis Strait off southern Baffin Island based on cruise data, but it is also known from the central Labrador coast (northern Labrador in FishBase) and southwest and southeast Greenland and is included here on that basis. It is also found south to Georges Bank in Atlantic Canada. Distribution of Argentina silus

source:  Jørgensen et al. (2005).



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Bathylagus euryops

Family Microstomatidae

Goode and Bean, 1896

Goitre Blacksmelt, garcette-goître

Pencilsmelts, Microbecs

Brian W. Coad

common names: A local name is Almindelig Sortsmelt (Danish/Greenlandic). The common name refers to the heavily developed lower part of the head, said to resemble a goitre.

taxonomy:  The genus comes from the Greek bathys (deep) and

lagos (hare), in reference to the big eyes. The species name comes from the Greek eurys (wide) and ops (eye).

This family is found in all oceans from the Arctic to Antarctica. There are about 38 species, with four found in Canada, including one in the Arctic. The sole Arctic member and its relatives were once placed in another family, the Bathylagidae. These are small fishes, usually less than 30 cm in length. Members of this family are also called Deepsea Smelts, Blacksmelts, and Smoothtongues. The small mouth has compressed teeth in single rows, which are absent from the upper-jaw bones (maxilla and premaxilla). The fins lack spines. There is no gas bladder and no light organs. The adipose fin may be present or absent. The eyes are large to medium in size. The scales are large, cycloid, and easily lost. The scales and lateral line extend onto the tail. There are only two to four branchiostegal rays. Pencilsmelts are oceanic, found in midwater and near the bottom. They can be quite abundant. Some species migrate vertically. Their food is zooplankton. The eggs and larvae are pelagic, and the larvae can be distinctive with eyes on stalks in some species. They are not commercially important.

source:  Kobyliansky (1986).

description:  The Goitre Blacksmelt is distinguished by a combination of characters outlined in the family account and here. There are about 38–41 lateral line scales and a small gill opening not extending halfway up the side of the body. There are only 4–6 scales between the dorsal and pelvic fins on the side of the body. The eyes are large, almost half the head length. Dorsal fin-rays number 9–11, usually 10; anal fin-rays 16–21, usually 18; pectoral fin-rays 7–12, usually 9; and pelvic fin-rays 7–9, usually 8. The anal fin base is longer than the caudal peduncle. Gill rakers number 17–18 on the lower arch. The scale pockets are outlined in black and sometimes lined with purplish tissue. The overall body colour is light to dark brown to light purple. The opercle is blue black, and the snout dusky. This species reaches about 21.0 cm in length. habitat:  Juveniles and adults are mesopelagic to bathypelagic

and are found between 500 m and 3,237 m, while post-larvae are found at 20–500 m. This species has been reported at 293–1,457.5 m in Davis Strait and southern Baffin Bay at −0.08°C to 4.5°C, and to 441 m in Ungava Bay. Cruise data from Baffin Bay to Davis Strait indicate a depth range of 216–1,474 m. Adults may be caught in large aggregations. In the pack-ice zone of Davis Strait, larvae of this species dominate the open-water capture stations.

biology:  Its food is small planktonic crustaceans. This species is eaten by Rock Grenadiers and Roughhead Grenadiers in the Davis Strait.

Bathylagus euryops

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importance:  It is not economically important. distribution:  This species is found in Baffin Bay, Davis Strait,

eastern Hudson Strait, Ungava Bay, and northwest, southwest, and southeast Greenland, and southward to Bermuda, and generally in the North Atlantic Ocean.

Family Platytroctidae Tubeshoulders, Circés

Brian W. Coad

Distribution of Bathylagus euryops

sources:  Karrer (1973); MacLaren Marex Inc. (1979a); McKelvie & Haedrich (1985); Jørgensen (1996); Treble, Brodie, Bowering, & Jørgensen (2000); Jørgensen et al. (2005).

Tubeshoulders are small deep-sea fishes found in all temperate to tropical oceans. There are about 37 species, with 13 reported from Canada, including 5 in the Arctic. These fishes are small, usually less than 30 cm in standard length. These fishes are unique in having a special sac under the cleithrum (the large bone supporting the pectoral fin). This shoulder organ is a black-lined pit and produces a blue-green luminous fluid that reaches the outside by means of a tube visible above the pectoral fin. The tube is small but dark in colour. Many species also have light organs, particularly on the belly, where they are not in rows. The dorsal fin is over the anal fin at the rear of the body. The pectoral fin has 13–28 soft rays. The enlarged anterior premaxillary teeth are called “tusks.” There is a canal system under the skin connected to scale pockets by pores. The overall colour is dark brown to black. Many of the more obvious characters like fin-ray and scale counts overlap between species, and these fishes can be difficult to identify. Identifications and distributions of some species are suspect in literature records for Canada. Adults are found mostly at depths of 250–900 m, with some species to over 5,000 m, and are only seen by scientists. They do not seem to migrate vertically. Tubeshoulders produce a few large eggs that are found at similar depths to the adults. Their food is probably small crustaceans. The shoulder organ, with its backward pointing tube, acts as a mechanism to confuse predators by a squirt of luminous fluid, rather like the ink cloud of octopus and cuttlefish. The sudden blinding cloud of blue-green points of light enables the tubeshoulder to escape into the darkness. They are not commercially important.

sources:  Parr (1960); Sazonov (1986); Matsui & Rosenblatt

(1987).



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Holtbyrnia anomala

captures could be shallower than the greatest depths at which the nets fished. An Arctic Canadian record was taken from 1,132 m.

Bighead Searsid, tube-épaule grosse tête

biology:  Unknown.

Krefft, 1980

common names: A local name is Storhovedet Skulderlysfisk

(Danish/Greenlandic). The French common name was coined by Claude B. Renaud. The use of “searsid” in common names here and below refers to the former family name of Searsidae or Searsiidae.

taxonomy:  The genus comes from Ernest William Lyons Holt (1864–1922) and L.W. Byrne, British ichthyologists who worked on deep-sea fishes, among others, off Ireland. The species name comes from the Latin anomalus (anomalous or irregular), in reference to the rudimentary photophores. This species in the Arctic was identified aboard ship, and no voucher specimen was kept. Its presence in neighbouring Greenland waters occasions its inclusion here. description:  This species is distinguished from its Arctic relatives by having weak photophores present or even absent in adults (except for a well-developed PO or pectoral fin photophore), and absent in young – photophores IVO (interventral organ between pelvic fins) and GO2 (posterior gular organ on chin) present but rudimentary in young; pelvic fin-rays number 7–9, usually 9; pectoral fin-rays number 13–17; cleithral symphysis has no spine; and there is white tissue on infra-orbitals 2–3 and, after a gap, on infra-orbitals 4–6. Dorsal fin-rays number 18–20, anal fin-rays 16–17, scales in mid-lateral series 103–113, and total gill rakers 27–28, with 8–9 on the upper arch. Gill filaments are short and united at the base. The colour is dark brown to blackish, with the opercles and inside mouth being black in preserved fish. Photophores found in adults of other species are replaced by colourless spots lacking scales. This species reaches 31.0 cm in total length.

importance:  It is not economically important. distribution:  It has been found in Davis Strait as a single record from cruise data at 63.1019º N, 59.9665º W, off southern Baffin Island, but it is also found in southwest and southeast Greenland, and generally in the temperate to tropical Atlantic Ocean.

Distribution of Holtbyrnia anomala

sources:  Krefft (1980); Neat & Campbell (2013).

habitat:  It is apparently bathypelagic to benthopelagic, perhaps from 70 m to 2,700 m, although, as open gear was used, some

Holtbyrnia anomala

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habitat:  It is a bathypelagic species. Arctic Canadian captures

Maulisia mauli

were at depths of 494–1,360 m.

Parr, 1960

biology:  Unknown.

Maul’s Searsid, tube-épaule de Maul

common names: A local name is Mauls Skulderlysfisk (Danish/

Greenlandic). Another common name is Maul’s Tubeshoulder. The French common name was coined by Claude B. Renaud.

taxonomy:  The genus comes presumably from the same ori-

importance:  It is not economically important. distribution:  It has been found in Davis Strait as four records from cruise data. It is also found in southwest and southeast Greenland and in the Atlantic and Indian Oceans.

gin as the species that is named after the German ichthyologist Günther Edmund Maul (1909–97) of the Museu Muncipal do Funchal, Madeira. This species in the Arctic was identified aboard ship, and no voucher specimen was kept. Its presence in neighbouring Greenland waters occasions its inclusion here.

description:  This species is distinguished by photophores being

weakly developed; pelvic fin-rays numbering usually 6–8; cleithral symphysis produced as a spine; the upper jaw extending far back beyond eye level (to rear of eye in smaller fish); frontal bones being widest over the middle of the eye level, abruptly narrower before and behind; pores in scale pockets being much smaller than scales; the lateral line having papillae or being unmarked; the body being deep (3.5–4.0 times in standard length) and the snout being short (3.8–4.8 times in head length); the shoulder organ opening having several scale rows wide behind the supracleithrum; and premaxilla having 4–8 teeth behind the tusks (enlarged anterior premaxillary teeth). Dorsal fin-rays number 17–22, anal fin-rays 15–19, pectoral finrays 17–20, pelvic fin-rays 6–9, mid-lateral scales 79–95, and total gill rakers 23–26, with 7–8 on the upper arch. The pelvic base and the anus are separated by 14–16 scale rows. The THO (thoracic photophore) is a short bar. There is white tissue on the infra-orbital bones at the posterior border of the eye, extending forward along the posterior third of the eye. The species attains 25.0 cm in length.

Distribution of Maulisia mauli

sources:  See the family sources and the bibliography.

Maulisia mauli



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Maulisia microlepis

Sazonov and Golovan, 1976 Smallscale Searsid, tube-épaule petites écailles

common names:  A local name is Småskællet Skulderlysfisk (Danish/Greenlandic). Another common name is Smallscale Tubeshoulder. The French common name was coined by Claude B. Renaud.

biology:  Unknown. importance:  It is not economically important. distribution:  It has been found in Davis Strait as three records from cruise data, and in southwest and southeast Greenland (and on the Greenland side of Baffin Bay to 71º41' N), and the North and South Atlantic Ocean.

taxonomy:  The species name comes from the Greek mikros (small) and lepis (scale). This species in the Arctic was identified aboard ship, and no voucher specimen was kept. Its presence in neighbouring Greenland waters occasions its inclusion here. description:  This species is distinguished by photophores being absent; pelvic fin-rays numbering usually 6–8; cleithral symphysis being produced as a spine; the upper jaw extending far back beyond eye level (to rear of eye in smaller fish); frontal bones being widest over the middle of the eye level, abruptly narrower before and behind; pores in scale pockets being much smaller than scales; the lateral line having enlarged and modified scales; the body being relatively shallow (4.3–5.2 times in standard length) and the snout long (3.2–3.8 times in head length); no pit being behind the supracleithrum; and premaxilla having 7–15 teeth behind the tusks (enlarged anterior premaxillary teeth). Dorsal fin-rays number 17–21, anal fin-rays 15–18, pectoral finrays 13–18, pelvic fin-rays 6–9, mid-lateral scales 112–125, and total gill rakers 27–33, with 8–9 on the upper arch. The pelvic base and the anus are separated by about 12 scale rows. The body is uniformly blackish brown in colour. The species attains 25.5 cm in standard length.

Distribution of Maulisia microlepis

source:  Sazonov & Golovan (1976).

habitat:  A bathypelagic to benthopelagic species, it has a depth range of 500–2,000 m. Arctic Canadian captures were at 1,231–1,369  m.

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biology:  Unknown.

Normichthys operosus Parr, 1951

importance:  It is not economically important.

Longsnout Manypitshoulder, épaule-criblée long nez

common names: A local name is Grubet Skulderlysfisk (Danish/Greenlandic). Another common name is Multipore Searsiid. taxonomy:  The genus is not explained in the original description but may be named after the British ichthyologist John Roxborough Norman (1898–1944), author of A History of Fishes, who worked on the type material, and the Greek ichthys (fish). The species name comes from the Latin operosus (elaborate). Normichthys operosa islandica Parr, 1960, described from Iceland is a synonym. This species in the Arctic was identified aboard ship, and no voucher specimen was kept. Its presence in neighbouring Greenland waters occasions its inclusion here.

distribution:  It was found in Davis Strait off southern Baffin Island as a single record from cruise data at 66.5119º N, 58.8344º W, but also has been found from southwest and southeast Greenland, off Atlantic Canada, and in the eastern Atlantic Ocean from Iceland to Angola.

description:  This species is distinguished by its lack of photophores, or, if they are present, their development is weak; 6–8, usually 8, pelvic fin-rays; the cleithral symphysis produced as a spine; the upper jaw extending back to about behind the pupil level; the frontal bones being widest over posterior eye level or behind eye level; and about 1–7 scale-sized or twice-scale-sized openings or pores in scale pockets behind the shoulder girdle. Dorsal fin-rays number 17–20, anal fin-rays 16–17, pectoral fin-rays 16–17, total gill rakers 28–30, with 7–8 on upper arch, and –mid-lateral scales 80–99. The gill rakers are broad and elongate, the longest overlapping the sixth adjacent raker. The overall colour is dark. Scale pockets are outlined with dark pigment. This species reaches 16.4 cm in total length.

Distribution of Normichthys operosus

habitat:  It is a bathypelagic species found down to below 1,000

sources:  See the family sources and the bibliography.

m, with shallowest depths generally at 780–785 m; a Canadian Arctic specimen was caught at 721 m in Davis Strait.

Normichthys operosus



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Platytroctes apus Günther, 1878

Legless Searsid, tube-épaule apode

common names: A local name is Højrygget Skulderlysfisk (Danish/Greenlandic). Another common name is Legless Tubeshoulder. The French common name was coined by Claude B. Renaud.

taxonomy:  The genus comes from the Greek platys (flat, broad)

and troktes (gnawer, nibbler), and the species name comes from the Greek a- (without) and pous (foot), in reference to the absence of pelvic fins.

description: This species is distinguished by the absence of pelvic fins; the body is very compressed and deep (3.0 or less times in standard length); about half of the dorsal and ventral body margins have a sharp, non-muscular keel; and the predorsal margin is sharp and one scale wide.

The cleithral symphysis is an elongate and sharp spine, protruding ventrally in preserved fish. Most scales have a single median ridge or keel that is open posteriorly and terminates in a raised peak. The lateral line is straight on the mid-body and has 85–107 scales. The dorsal and ventral margins of the caudal peduncle have black luminous sacs between the scales. Young have mid-dentary teeth, and in adults there is a gap. Dorsal fin-rays number 15–21, anal finrays 15–20, and pectoral fin-rays 19–25. The pectoral fin is very small. Total gill rakers are 29–41. Pyloric caeca number 5, with 5–7 terminal lobes. The body is dark brown in colour with a black band on the upper and lower caudal peduncle edges. The mouth, anus area, and all organs in the abdominal cavity are black. The shoulder region under the opercular flap is silvery. Preserved fish may have a dark head and pale body. The species attains 18.0 cm in standard length.

habitat: It is a benthopelagic species found from 300 m to 5,393 m. The single Arctic Canadian record was found at 1,238 m and 3.8°C. biology:  Its biology is unknown in detail. One specimen was found in the gut of a cod caught at 385 m.

Platytroctes apus

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importance:  It is not economically important. distribution:  It was found in Davis Strait as a single rec-

ord caught in October 2013 (CMNFI 2013-0052) from 62°34.86' N, 59°17.71' W. It is also found in southwest and southeast Greenland, and Atlantic Canada, and worldwide in all oceans except polar ones.

Family Alepocephalidae Slickheads, Alépocéphales

Brian W. Coad

Distribution of Platytroctes apus

sources:  See the family sources and the bibliography.

Slickheads or Smoothheads are deep-sea fishes found in all oceans. There are about 75 species, and 17 are reported from Canada, including 6 in the Arctic. The maximum size exceeds 1 m, although most species are much smaller. These are dark-coloured fishes with anal and dorsal fins far back on the body near the tail; often weak teeth in a small mouth (fangs may be present or teeth may be absent altogether); no tusks on the premaxilla as in many Platytroctidae; no adipose fin; no gas bladder; many long gill rakers; no scales on the head (hence “slick head” or “smooth head”), but large cycloid scales on the body, which are sometimes absent; slippery, loose skin over a flabby body; and no pelvic axillary scale. The body is almost 90% water. If present, the lateral line is composed of a pored tube supported by ring-like scales, or papillae. Light organs are present in some under the skin or on stalks or complexly arranged. The head and eyes are often large. The overall colour is brown to black, although some have bright blue skin on the head and fin bases. Slickheads are found in the deep ocean often below 1,000 m and down to 5,000 m but can be locally abundant. Their food often includes jellyfish as well as crustaceans, echinoderms, polychaetes, decapods, tunicates, and fishes. The eggs are large but do not float to the surface. Some members of this family are only caught by research vessels, and rarely at that, while others, being larger, are occasionally taken by commercial deep-sea trawlers but are not commercially important. The flesh has a poor texture.

sources:  Parr (1952); Sazonov (1999).



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Alepocephalus agassizii Goode and Bean, 1883

Dusky Slickhead, alépocéphale obscur

common names: A local name is Agassiz Glathovedfisk (Danish/Greenlandic). Other common names are Agassiz’ Slickhead and Agassiz’ Smoothhead.

importance:  It is not economically important. distribution:  This species is found in Baffin Bay and Davis

Strait, on both sides of the North Atlantic Ocean including northwest, southwest, and southeast Greenland (to 74º53' N on the Greenland side of Baffin Bay), and south to Honduras on the American side.

taxonomy:  The genus comes from Greek alepos (without scales)

and kephale (head). The species was named after Alexander Emanuel Rodolphe Agassiz (1835–1910), the Swiss and U.S. marine ichthyologist from whose deep-sea researches the type was obtained.

description:  This species is distinguished by fin-ray and scale counts, by the absence of teeth on the maxilla bone of the upper jaw but the presence of teeth on the premaxillae, dentaries, and palatines, and by the dorsal fin origin lying above the anal fin origin. Dorsal fin-rays number 15–18, anal fin-rays 15–18, pectoral finrays 9–12, pelvic fin-rays 6–8, and lateral line scales 80–95. Gill rakers number 23–32. The gill opening extends forward under the eye. The overall colour is black, which may change to purplish brown on capture. It attains 79.0 cm in standard length. habitat:  The Dusky Slickhead is bathydemersal and engybenthic

over sandy, clay, or stony bottoms at depths usually of 600–2,400 m, in schools. It has been caught at 1,145–1,463 m in Davis Strait at 0.3°C–3.0°C and on another cruise in the strait at 668–1,472 m. A record from Baffin Bay was as shallow as 534 m.

biology:  Its food is mainly comb jellies but also includes crustaceans, starfish, and polychaete worms. The eggs are up to 2.8 mm in diameter and number up to 9,150.

Distribution of Alepocephalus agassizii

sources:  Treble, Brodie, Bowering, & Jørgensen et al. (2000); Jørgensen et al. (2005).

Alepocephalus agassizii

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importance:  It is not economically important.

Alepocephalus bairdii Goode and Bean, 1879

distribution:  This species is found in Davis Strait, and in south-

Manyray Smoothhead, alépocéphale multirai

west and southeast Greenland south to the Grand Banks and off the Scotian Shelf. It is found also in the eastern North Atlantic Ocean.

common names: A local name is Bairds Glathovedfisk (Danish/Greenlandic). Other common names are Baird’s Slickhead, Baird’s Smoothhead, and alépocéphale de Baird. taxonomy:  The species is named after the ichthyologist Spencer

Fullerton Baird (1823–87), a head of the U.S. Commission of Fish and Fisheries and first curator and second Secretary at the Smithsonian Institution.

description:  This species is distinguished by fin-ray and scale counts, by the absence of teeth on the maxilla bone of the upper jaw, and by the dorsal fin origin lying above the anal fin origin. Dorsal fin-rays number 18–23, anal fin-rays 20–25, pectoral fin-rays 8–13, pelvic fin-rays 7–9, and lateral-line scales 62–70. Gill rakers number 26–32, and pyloric caeca 13–18. The overall colour is a purplish brown, with the gill and mouth cavities being very dark. Young fish are darker than adults. It reaches 100.0 cm in standard length. habitat:  This species is bathydemersal and engybenthic and is

found in schools over sandy and ooze bottoms at about 365–1,700 m, sometimes shallower. It has been caught at depth ranges of 690– 740 m and 1,100–1,321 m in Davis Strait.

biology:  Its food is comb jellies, sea squirts, fish such as deep-

sea anglerfishes, and crustaceans such as decapods. Demersal eggs are reported to be as large as 4.6 mm in diameter, and spawning may occur in winter. Up to 160,000 whitish to light-orange eggs are produced. Fish are mature only at 55 cm or more for males and 70 cm or more for females, perhaps 1 m. The maximum age is 38 years.

Distribution of Alepocephalus bairdii

sources:  Karrer (1973); Nielsen, Bertelsen, & Nystrøm (1992); Jørgensen et al. (2005).

Alepocephalus bairdii



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Bajacalifornia megalops (Lütken, 1898)

Bigeye Smoothhead, alépocéphale à grands yeux

distribution:  This species has been found in all oceans including three records in Davis Strait off southern Baffin Island (cruise data and ZMB 23712), southwest and southeast Greenland, and off the Scotian Shelf in Atlantic Canada.

common names: A local name is Storøjet Glathovedfisk (Danish/Greenlandic). Another common name is Bigeye Slickhead. taxonomy:  The genus comes from “Baja California” where the type species was taken. The species name comes from the Greek megalo (large) and ops (eye). Bathytroctes drakei Beebe, 1929, described from the western Atlantic Ocean at 39°15' N, is a synonym. description:  This species is distinguished by fin-ray and scale counts, by teeth on the maxilla, and by the protruding lower-jaw tip. Dorsal fin-rays number 16–18, anal fin-rays 13–16, pectoral finrays 12–17, and pelvic fin-rays 7–8. Lateral-series scales number 50–56. The presence of teeth on the maxilla bone of the upper jaw and the location of the dorsal fin origin well in front of the anal fin origin are also characteristic. Gill rakers number 25–29. The overall colour is blackish brown. It attains 39.5 cm in standard length. habitat:  This species is bathypelagic, mesopelagic, and bentho-

pelagic. Adults have been reported from 775 m to 1,425 m, and larvae and young from 250 m to 3,182 m. In Davis Strait this species has been caught at 775–820 m and a temperature of 3.4°C, and one specimen was from 1,247 m.

biology:  Unknown.

Distribution of Bajacalifornia megalops

sources: Karrer (1976); Markle & Krefft (1985); Jørgensen et al. (2005).

importance:  It is not economically important.

Bajacalifornia megalops

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distribution:  Members of this genus are found worldwide in deeper waters. The Canadian Arctic records are from Davis Strait off southern Baffin Island.

Bathytroctes sp. common names: None. taxonomy:  The genus comes from the Greek bathys (deep) and troktes (one who gnaws). This fish was identified only to genus aboard ship, and no material was kept from Canadian Arctic waters. There are several species in this genus that occur in the northern Atlantic Ocean and could reach Arctic waters. The illustration is one of these, the Oarjaw Wingmax or maxailé aviron (Bathytroctes michaelsarsi Koefoed, 1927). description:  The genus Bathytroctes is distinguished by having body scales; 40–80 transverse scale rows; 9–11 anal fin-rays; body flanks that are slightly concave; a deep caudal peduncle; teeth present on the maxilla (upper jaw), lower jaw, and/or vomer bone in the roof of the mouth; teeth near the front of the jaws in a single series, the maxilla ending about below the rear orbit margin level; no prominent knob on the lower-jaw tip; the dorsal fin origin in front of the anal fin origin level; the first pectoral fin-ray not thread-like and elongated; and gill filaments often fused at their bases. habitat:  Its habitat is unknown in detail. Canadian Arctic fish have been caught at 468–1,482 m.

Distribution of Bathytroctes sp.

source:  Sazonov (1999).

biology:  Unknown. importance:  It is not economically important.

Bathytroctes sp.



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Rouleina maderensis Maul, 1948

Madeiran Smoothhead, alépocéphale de Madère

common names: A local name is Nøgen Glathovedfisk (Dan-

ish/Greenlandic). Another common name is Madeiran Slickhead. The French common name came from Claude B. Renaud.

taxonomy:  The genus is presumably named after Louis Roule

biology:  Maturity is attained at about 20–25 cm in standard length. The eggs are up to 3.7 mm in diameter and are pelagic.

importance:  It is not economically important. distribution:  It has been found in Davis Strait as a single record at ca. 62º40' N, 58º56' W (CMNFI 2004-0008), in southwest and southeast Greenland, and in the Central and North Atlantic Ocean, western Indian Ocean, southeast Pacific Ocean, and western tropical Pacific Ocean.

(1861–1942), the chair of reptiles and fishes at the Muséum national d’Histoire naturelle, Paris, who studied alepocephalids; and from the Latin suffix –ina (of, relating to). The species name means “from Madeira.”

description:  This species is distinguished by being scaleless except for the lateral line, which is supported by ring-like scales standing on edge inside the canal. The head is very large, about one-third of the body length, with large eyes. The upper jaw extends a little beyond the eyes. The skin is deciduous and contains small, longitudinal, fluid-filled compartments. Nodular light organs are present and are most numerous below the lateral line. They are less evident in larger fish and are often rubbed off. Papillae are mostly restricted to the fins and fin bases. The teeth are in a single row on the premaxilla, maxilla, and dentary bones of the mouth. The dorsal and anal fins are about equal in size and opposite, far back on the body. Dorsal fin-rays number 19–24, anal fin-rays 19–24, pectoral fin-rays 5–7, usually 6, and pelvic fin-rays 5–6. Lateral-line scales number 50–57, and vertebrae 47–50. Gill rakers number 22–30 in total and are united at their bases (up to 40%–60% of their total length). Pyloric caeca number 10–11. The testes are in separate lobes. The overall colour is black. This species attains 32.0 cm in standard length. habitat:  The species is engybenthic and benthopelagic over 595–

Distribution of Rouleina maderensis

sources:  Markle (1978); Sazonov & Williams (2001).

1,450 m. A Canadian specimen was caught at 1,165 m, 3.9ºC, and 34.91 psu in October 2013.

Rouleina maderensis

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biology:  It feeds on small crustaceans and squids. Spawning

Xenodermichthys copei

occurs from September to November near the bottom. The eggs are large (2.7 mm in diameter), and only a few are laid (about 150).

(Gill, 1884)

Atlantic Gymnast, gymnaste atlantique

importance:  It has no importance, even where it is common outside Arctic waters, because the flesh is watery.

distribution:  This species is found in Davis Strait off southern Baffin Island and in all oceans.

Xenodermichthys copei

common names:  A local name is Kortsnudet Glathovedfisk

(Danish/Greenlandic). Other common names are Black Slickhead and Bluntsnout Smoothhead.

taxonomy:  The genus comes from the Greek xenos (strange),

derma (skin), and ichthys (fish). The species is named after Edward Drinker Cope (1840–97), a U.S. ichthyologist and palaeontologist, after whom Copeia, the journal of the American Society of Ichthyologists and Herpetologists, is named.

description:  This species is unique among Canadian Slickheads by the body being completely scaleless and having numerous light organs as small, pale, and round nodules on the thick skin of the body and on the head. The forehead over the eyes is strongly rounded and leads to a very short and blunt snout. Dorsal fin-rays number 27–31, anal finrays 26–30, and pectoral fin-rays 7–8. The colour is black overall. It reaches 31.0 cm in total length.

Distribution of Xenodermichthys copei

habitat:  This is a bathypelagic species of the continental slope

sources:  Markle & Wenner (1979); Carvalho & Almeida (1988).

found at about 100–2,650 m. It has been caught at 703–1,062 m and 3.8°C off Baffin Island and at 416–1,132 m in Davis Strait.

Xenodermichthys copei



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Mallotus villosus

Family Osmeridae

(Müller, 1776)

Capelin, capelan

Smelts, Éperlans

Brian W. Coad

Mallotus villosus

Smelts are found in coastal sea waters and fresh waters near the coast in the northern hemisphere. Some populations are land locked. There are about 13 species, with 11 reported from Canada, including 3 in Arctic waters. The maximum size is about 40 cm. These small, silvery fishes have a single dorsal fin at mid-body, an adipose fin, no pelvic axillary process (found in Salmonidae – Trouts and Salmons), 8–9 pelvic fin-rays, 19 principal caudal finrays, a forked tail, 5–10 branchiostegal rays, 0–12 pyloric caeca, and a stomach sometimes with a blind sac; the last vertebra at the tail is turned up; and teeth are numerous on the mouth bones and are strong or weak. The scales are cycloid and easily detached. Some Smelts have a strong cucumber smell when fresh, the chemical being trans-2-cis-6-nonadienal. Its function is unknown. Breeding Smelts can develop tubercles, modified scales, and enlarged fins. Reproduction may occur in coastal waters or involve a migration into fresh water. Smelts are all carnivorous fishes. They are very numerous and so are important food fishes for commercially important species and are also used as bait by anglers. They are rich in oil and are excellent smoked for human consumption.

sources:  McAllister (1963b, 1967); Kljukanov (1969, 1972, 1977); Gruchy & McAllister (1973); Kljukanov & McAllister (1973); Nellbring (1989).

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common names: Local names are Amagiak, Angmaggeuck, Axmagiaq, Holili-gah, Ko le le kuk, Kuliligak, Nulilighuk, Panmagrik, Panmaksraq, Qulilirraq, Qoliiligaq (Inuktitut); Angmagiak (Inuvialuktun); Kashkanamesh (Innu-Aimun); and Ammassat (Greenlandic). Other common names are Caplin, Lodde, and capelan atlantique. taxonomy:  The genus comes from the Greek mallotos (villous or tufts of hair). The species name comes from the Latin villosus (hairy). Both the genus and the species allude to the scales in the spawning season. The subspecies Mallotus villosus catervarius (Pennant, 1784), described from Kamchatka, Russia, is not recognized by most North American authors. Russian authors report it from the Arctic coast of Alaska east to Queen Maud Gulf with the type subspecies Mallotus villosus villosus from the islands of the western Arctic and Hudson Bay. Mallotus villosus catervarius natio schulzi Rumyantsev, 1947, has been used for Canadian Capelin in the original description but is an infra-subspecific taxon and is not available for nomenclatural purposes, nor has it been used in North America. DNA bar-coding has shown that fish from the Chukchi Sea, northern Bering Sea, Gulf of Alaska, and Davis Strait form a separate clade from fish from the Maritimes of Canada and Hudson Bay. Salmo groenlandicus Bloch, 1794, described from Iceland, Greenland, and Norway, is a synonym. description:  This species is distinguished by the teeth on the tongue being small and brush-like and by a complete lateral line having about 170–220 scales. Dorsal fin-rays number 10–18, anal fin-rays 16–26, and pectoral fin-rays 16–22. Pelvic fin-rays number 8–9. The last pelvic fin-ray is reduced in size. Pyloric caeca number 3–9, and gill rakers 33–48. Breeding males develop a strong ridge of four rows of overlapping, elongated scales along the lateral line. The ends of the scales give the ridge a hairy appearance. Development starts four to five weeks prior to the start of the spawning season. In addition, the anal fin is enlarged, with the first 10–12 rays thickened, and projects from a ridge on the belly. The pectoral and dorsal fins are also enlarged. Tubercles develop on the head, the lower pectoral, pelvic, and caudal

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Mallotus villosus

fin-rays, and the belly. Overall it is translucent, with an olive-green to yellow-green back, some purplish flank tints, silvery lower flanks, and a white belly. The operculum is spotted. The scales are outlined with pigment. The peritoneum is silvery but has dense dark speckles and appears almost black. The back and head darken at spawning time. The species reaches 25.2 cm in total length.

habitat:  This is a pelagic, schooling species found from the surface down to over 1,000 m. Records in Davis Strait and Ungava Bay generally are from 105 m to 1,086 m. It is usually in coastal areas and on offshore banks. There are daily vertical migrations. It has been caught at 263–356 m in Hudson Strait and may be found stranded in tide pools in Hudson and James Bays. Sometimes it is so numerous as to leap into canoes, as reported in Bathurst Inlet in 1821. It is found in the low-salinity waters of Richmond Gulf and James Bay. Salinities as low as 4‰ are tolerated over winter at −1°C to 6°C in the Eastmain River estuary in James Bay. Eggs on shore can survive temperatures down to −5.2°C for up to six hours, with some eggs surviving longer; and some can survive lower temperatures for short periods. In Rupert Bay and nearby James Bay, larvae are caught mostly at 9–15°C and at salinities above 3‰ from June to August. It is numerous in southern Hudson Bay – at the Belcher Islands and La Grande River, for example – probably as a relict from a warmer period, but its numbers in the cooler Ungava Bay vary from numerous to absent, presumably in response to temperature variations. Several thousand fish were seen in the



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Koksoak River of Ungava Bay on 8 August 1883, said to be the first appearance of this species ever observed. They were rare during 1947–52 surveys but were relatively abundant from 1959 to 1973. At Churchill in western Hudson Bay, schools have been observed at temperatures of 2–16°C in summer. Over a thousand pounds of spawning Capelin were caught in Thetis Bay, western Beaufort Sea, on one occasion, whereas a later study found none. Large numbers (ca. 25,000) were seen spawning on 24 July on a beach at Herschel Island in 1960, when it was noted that their appearance was recent. Capelin spawned in Sachs Harbour for two to three years in such numbers that they could be scooped up in buckets. This variation in numbers is documented for other localities too. They appear to avoid the turbid water associated with the Mackenzie River. Like many other polar and subpolar fishes, Capelin contain anti-freeze proteins that aid in survival in the shallow, icy sea water of nearshore areas. Their eggs too are highly resistant to freezing, which could help their expansion into the Arctic. Recent studies have shown an increase in this species in the diet of Thick-billed Murres that is associated with a general warming of Hudson Bay waters owing to climate change; Arctic Cod, a bigger and fatter fish, is their normal diet, and its replacement by Capelin will mean less food for young murres. Capelin have apparently expanded northward into the mid-Arctic, for example at Coats Island in northern Hudson Bay and at the Minarets on southeast Baffin Island, over the last 30 years as shown by these bird diets.

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biology:  Its food is plankton, particularly euphausiids and

copepods, but also includes polychaete worms and small fishes. In West Greenland waters the dominant prey items by wet weight are euphausiids (61%), amphipods (18%), and copepods (10%). Capelin also eat the eggs of their own species. There are early morning and evening feeding peaks. Feeding is intense before spawning in late and early spring but stops during spawning. This species and Sand Lances replace Arctic Cod as an intermediate carnivore in southern Hudson Bay and James Bay. Capelin are high-energy prey resulting from their lipid content. They are the main food of Ogac when available on the northwest coast of Hudson Bay and in the Belcher Islands of southeastern Hudson Bay. In the estuary of the La Grande River of James Bay they are eaten by Lake Sturgeon, Brook Trout, and Ciscoes. In the Richmond Gulf, Brook Trout feed on this species second only to Pacific (= Stout) Sand Lance. Brook Trout eat Capelin in western James Bay. Arctic Char eat this species in Hudson Bay, including the Belcher Islands, and in Clearwater Fiord, Baffin Island. Ungava Bay cod and Greenland Halibut feed on Capelin at 366 m (200 fathoms) and 549 m (300 fathoms), respectively. Greenland Halibut feed on pelagic Capelin in Cumberland Sound as inferred from stable isotope analysis of halibut tissue. (Most halibut are harvested in winter and early spring and have a high percentage of empty stomachs, such that diet could best be determined by this indirect method.) In Davis Strait Capelin have been found in the stomachs of Atlantic Salmon and Greenland Halibut. In eastern Ungava Bay, when present, they are a principal food for Arctic Char instead of crustaceans, changing the char flesh from pink to white. They are eaten by Saffron Cod, Arctic Char, and Starry Flounder in the Coppermine River Delta. In the decade prior to 2013, Capelin appeared in Cumberland Sound and were newly present in the diet of Arctic Char, a shift from a primarily invertebrate diet. Capelin are extremely important as food for Atlantic Cod, Haddock, Greenland Halibut, Canadian Plaice, Yellowtail Flounder, Roughhead Grenadier, and Atlantic Salmon and are probably the most important prey species in the northern Atlantic Ocean. Capelin may form up to 98% of the Atlantic Cod diet and are the principal food for salmon in Atlantic Canada. In Digges Sound, Hudson Bay, this species is eaten by adult Thick-billed Murres and forms up to 20% of the diet of their chicks at Coats Island but only 1.4% of the chick diet at Akpatok Island, Ungava Bay. Thick-billed Murres have changed their diet from ice-associated Arctic Cod to one involving the Capelin, which is associated with little or no ice cover, as climate change has affected sea ice in Hudson Strait and northern Hudson Bay since 1995. Capelin are eaten by Brünnich’s Murres at Akpatok Island in Ungava Bay. Black Guillemot and Atlantic Puffin chicks are fed Capelin at Coburg Island, Nunavut. Arctic Terns feed on Capelin in the Churchill River estuary. Capelin are probably eaten by Harbour Seals throughout the Arctic wherever they co-occur. Ringed Seals probably eat this pelagic fish in the fall at Sanikiluaq, Nunavut, and it is part of the seal diet in southeastern Hudson Bay. Capelin are the most common food for Beluga in Hudson Bay and are eaten during spawning runs at Churchill, and also at Arviat. Beluga feed on this species in the southern Beaufort

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Sea, Cumberland Sound, and the High Arctic, although consumption is greater in Hudson Bay than in Cumberland Sound (Pangnirtung) and the High Arctic (Aujuittuq, or Grise Fiord) where Arctic Cod are preferred. Minke Whales, Fin Whales, Harp Seals, and Atlantic Cod annually consume 3 million t of Capelin in the northwest Atlantic Ocean. Capelin contribute 89% of the annual gross energy in Harp Seals from Hudson Strait and 98% in a single month. Even Killer Whales may eat this fish at Kangiqliniq (Rankin Inlet). This species is then a very important part of the food chain between crustaceans and Greenland Sharks, Ogac, the larger Sculpins, Eelpouts, and various sea birds; between crustaceans and Arctic Char to other fishes, Belugas, and Harbour Seals; and between crustaceans to the Hooded Seal, Beluga, Harbour Seal, and Harp Seal. The maximum lifespan is ten years, but most fish do not live longer than four to five years. Males grow faster than females until maturity, and then growth is about equal. Populations in the north grow more slowly and mature later than those in the south. For example, Labrador Capelin mature a year later than do Grand Banks Capelin but have accelerated growth at later ages to produce a similar maximum size. Spawning involves a migration onto beaches (the Capelin scull or roll) in Arctic waters. The timing of spawning depends on temperature, latitude, tides, daylight, salinities, turbidity, and winds and can occur from April to September in Canada. Spawning occurred in mid-July at 13°C–15°C in brackish water on beaches near Coppermine River during 24 hours of daylight, rather than in the dark as in southern regions. Spawning has occurred on gravel beaches at Pauline Cove and Simpson Point on the Yukon coast from 24 to 30 July. Ford Lake, a marine bay on the northeastern Keewatin coast, has Capelin preparing to spawn in mid-August. At Fort Churchill, Hudson Bay spawning is indicated for the end of July and the beginning of August, and near Port Nelson it is recorded for June. Spawning on shingle beaches in the Belcher Islands of Hudson Bay occurs from July to September. Large schools have been reported in Bellot Strait at the beginning of August just before the ice leaves, presumably a spawning accumulation. In Rupert and James Bays, the presence of larvae suggests several spawnings during June and July. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae have occurred in July and possibly August, suggesting a July spawning. Temperatures range from 2.5°C to 12.5°C and vary between localities, between years, and between weeks. Numbers and runs are generally low in the Beaufort Sea although large spawning runs occur occasionally at Herschel Island. Mating is reported as most intense at intermediate tides or during ebbing tides at or near a time of greatest tides. Spawning takes place on heavily overcast days and occasionally on sunny days. Large females spawn several weeks earlier than do small females. Reddish eggs are laid on coarse sand, fine gravel, and pebble beaches, and certain areas are known as Capelin-spawning beaches. The grain size of the beaches is 0.5–15.0 mm. The fish swim onto the beach on wave crests, and up to 60,000 reddish eggs, 1.2 mm in diameter, are shed and fertilized. Eggs may also be seen attached to seaweed and rocks. A male presses against the side of a female, and sometimes the female will be flanked by a male on

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each side. The spawning ridges and larger fins of the male help to hold the female. The female and accompanying males rush up the beach as far as possible and are stranded in one spot as the wave recedes. Vigorous movements of the fins and body audibly scoop out a small pool where eggs are shed and buried. Spawning takes less than five seconds, and thrashing movements help the fish regain the water as the next wave rolls up the beach. Wave action buries the eggs, and in this position they are protected from predators and dehydration. The eggs can be buried more than 15 cm in the sand to which they adhere. Spent fish can be stranded in large numbers on a beach, and not all eggs are safely buried, neighbouring rocks being covered by a continuous layer. Those that are buried may exceed 800/sq. cm. Most Capelin die after spawning, but repeat spawning by females may occur. Males gather in schools near the beach prior to spawning while females are in deeper water. More males die after spawning, perhaps because of greater damage in their longer stay in turbid beach water. Larvae emerge from the beach gravel when onshore winds blow warm water into the shallows. This warm water is usually rich in the smaller zooplankton and contains relatively few predators such as jellyfish and arrow worms, ideal conditions for larval growth and survival.

importance:  This species has been an important commercial product in Atlantic Canada. The peak offshore catch in 1976 was 370,000 t by former Soviet mid-water trawlers and Norwegian purse seiners. The Canadian catch in 1988 weighed 85,500 t, making this the third-largest catch by species, and was worth over $21.7 million. In 1985, flesh was valued at $0.17/kg. The Canadian Atlantic catch in 2011 was 32,448 t, worth $5,703,000. Millions of tonnes of Capelin are food for other species of fish each year in the Atlantic and Pacific Oceans, where it is a key forage species. Numbers appear to fluctuate in Arctic waters, however, possibly due to varying temperature regimes, with large numbers appearing at the mouth of the Koksoak River in Ungava Bay in August of some years but not others, for example. It is common in James Bay and southern Hudson Bay (particularly the Belcher Islands) where it has been suggested that it could be developed as food, fish-meal, and fertiliser, but commercial exploitation in the Churchill and Killiniq (Port Burwell) areas failed. A test fishery took place at Wemindji in James Bay in 1987–8 and 1988–9 for export to Japan, but commercial concentrations could not be identified. It has been used for human consumption, as bait, and as dog food in Hudson Bay and Strait. At Kugluktuk (Coppermine) some spawning Capelin have been used as dog food. At Sanikiluaq in the Belcher Islands it is eaten boiled or raw and has been used there, and at Churchill and south coast settlements in Hudson Bay, to supplement local food supplies. Capelin that washed ashore in large numbers in north Baffin Island and the northern Foxe Basin were used as dog food by local people. Its occurrence in the Beaufort Sea is too sporadic for fishery development, although it has been used as dog food locally when it appears. distribution:  This species is found near Aujuittuq (Grise Fiord), Nirjutiqavvik (Coburg Island), and Mittimatalik (Pond Inlet), in Davis Strait, Cumberland Sound, and the entrance to



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Frobisher Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, Foxe Basin, Chantrey Inlet, Melville Sound, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and the Beaufort Sea including western Banks Island. It is circumpolar in the northern Atlantic and Pacific Oceans and is found south to Juan de Fuca Strait on the British Columbia coast and south to Cape Cod in the western Atlantic Ocean. It is also found north to Thule at 76°30' N on the west coast of Greenland and on the east coast.

Distribution of Mallotus villosus

sources:  Richardson (1836a); Vladykov (1933a); Hildebrand (1948); Douglas (1951); Dunbar & Hildebrand (1952); Doan & Douglas (1953); Tuck & Squires (1955); Templeman & Squires (1960); Tuck (1960); Anonymous (1961); Ellis (1962); Bigelow & Schroeder (1963); Abrahamson, Gillespie, McIntosh, Usher, & Williamson (1964); Mansfield (1964); Ryder, Scott, & Crossman (1964); Mansfield (1967); Hunter (1968b); Dunbar (1970a); Robbins (1970); Lear & May (1971); Lalli, Buchanan, Thomson, & Wells (1973); Sergeant (1973b); Galbraith & Fraser (1974); Jangaard (1974); Kendel, Johnston, Lobsiger, & Kozak (1975); Kidd, Greendale, Morin, & Baxter (1975); Percy (1975); Simard & Legendre (1977); Fraker, Sergeant, & Hoek (1978); Fraker et al. (1979); Weir (1979); Dutil & Power (1980); Gaston, Cairns, Noble, & Purdy (1981); Ochman & Dodson (1982); Winters (1982); Dunbar (1983); D.B. Stewart & Bernier (1983); Gaston (1985, 1987, 1989a, 1991); Gaston, Cairns, Elliot, & Noble (1985); Watts & Draper (1986); Gaston, Elliot, & Noble (1987); Crawford (1989); Mikhail & Welch (1989); Stergiou (1989); Hudon (1990a); Baker, Lawrence, & Schneider (1993); Beck, Hammill, & Smith (1993); Gaston & Bradstreet (1993); Ponton, Gagné, & Fortier (1993); D.B. Stewart, Ratynski, Bernier, & Ramsey (1993); Welch (1995); Wein, Freeman, & Markus (1996); Robards, Gilchrist, & Allard (2000); Gaston, Woo, & Hipfner (2003); Dodson, Tremblay,

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Colombani, Carscadden, & Lecomte (2007); Gaston & Woo (2008); Kelley, Loseto, Yurkowski, Stewart, & Ferguson (2008); Dennard, McMeans, & Fisk (2009); Chambellant, in Ferguson, Loseto, & Mallory (2010); M. Ferguson et al. (2010); Kelley, in Ferguson et al. (2010); Hedeholm, Grønkjær, & Rysgaard (2012); Marcoux, McMeans, Fisk, & Ferguson (2012); Provencher, Gaston, O’Hara, & Gilchrist (2012); Carscadden, Gjøsæter, & Vilhjálmsson (2013); Hop & Gjøsæter (2013); Imrie & Tallman (2013).

Osmerus dentex

Steindachner and Kner, 1870 Pacific Rainbow Smelt, éperlan du Pacifique

cut-out of the medial edge of the lateral ethmoid, rarely below this bone through the cartilage). The following characters are repeated below for O. mordax as these two species have not always been separated in literature sources. Dorsal fin-rays number 8–11, anal fin-rays 11–18, pectoral fin-rays 9–14, and pelvic fin-rays 8. Total gill rakers number 26–37, and pyloric caeca 3–9, usually 5 or more. Males have numerous head, body, and fin tubercles, and a lateral ridge may be present. The mouth is large and reaches the rear of the eye. Paired and anal fins are larger in males, and a lateral ridge may develop. The back is olive green, steel blue, or black in colour, and the flank is silvery with purple, blue, or pinkish iridescences (hence “rainbow”). Scales on the back are outlined with pigment, and there may be pigment spots on the anterior flank. There is a silvery stripe along the flank that turns dark in preserved fish. The peritoneum is silvery with dark pigment dots. The species reaches 35.6 cm in total length (based on O. mordax, presumably the current species reaches similar sizes) but usually under 25.0 cm.

habitat:  Pacific Rainbow Smelt are found schooling in coastal

Osmerus dentex

common names: Local names are Eetooknuit (Inuivaluktun); Ilhuagnik, Iqaluarak, and Qaluarak (Inuktitut).

taxonomy:  The cucumber-like odour of these fishes when fresh is distinctive and probably accounts for the name “smelt.” The genus comes from the Greek osmeros (odorous), equivalent to smelt. The species name comes from the Latin dentex (toothed). Populations in the Pacific and western Arctic have been recognized as a subspecies, Osmerus mordax dentex, or as a distinct species, Osmerus dentex, by various authors. They have a general, external, morphological similarity to O. mordax although mitochondrial DNA and allozyme divergence is more marked. DNA bar-coding also indicates distinction. Biochemical characters are not useful in the field although the gap in distribution can be used to aid identification. description:  This species is distinguished by having medium

conical to large canine teeth on the tongue, 1–3 large canines on each side of the vomer bone in the roof of the mouth, and lateral-line scales numbering 51–73 (usually 62 or more), with 13–30 pored. External characters overlap with its relative, O. mordax, and definitive characters are osteological and molecular. Vertebral counts are usually 64–67 (range 63–68), versus usually 61–63 (range 59–65) in O. mordax. The parietal bones of the skull are more or less contiguous, sometimes weakly at the front (versus not in contact with each other), and the orbito-nasal vein exits the nasal capsule usually through the cartilage below the lateral ethmoid, rarely cutting through the lower edge of the bone (versus usually through a

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and pelagic oceanic waters. They overwinter under ice in the Beaufort Sea, for example, where they dominate the fish population numerically. They differ from other anadromous Beaufort Sea fishes by a long residence in the sea as adults. In the Beaufort Sea they are wide spread and abundant in brackish nearshore areas and are often the dominant species in offshore locations. Here they are found at −2.0°C to 13.5°C and at salinities of 0‰–32‰. They are pelagic with a maximum recorded depth of at least 150 m, possibly 425 m in the sea, although this greater depth could be a fish trapped nearer the surface as a deep-set net was retrieved. The Mackenzie River is the major, and perhaps only, spawning habitat in the Canadian Beaufort Sea. The Beaufort smelt enter the Mackenzie for a relatively brief period prior to ice break-up to spawn in the early spring over gravel substrates. Fry leave freshwater spawning areas and drop down to the sea soon after hatching, as early as July. Adults leave rivers immediately after spawning, passing through the outer Mackenzie Delta by July. There are an estimated 6,261,000 smelt in Mackenzie Bay, for example, making this species one of the most abundant in the Beaufort Sea. Fry drift to the Mackenzie Delta on the spring flood and disperse along the coast.

biology:  Its food is a wide variety of crustaceans such as calanoid copepods, hyperiids, euphausiids, mysids, amphipods, isopods, cumaceans, decapod larvae, polychaete worms, gastropods, coleopterans, chironomids, and squid, and small fishes such as Smelts, Sticklebacks, and Arctic Cod in the sea. In Philipps Bay, Yukon, fish occurred in 87% of the Pacific Rainbow Smelt stomachs examined and comprised 78% of the food biomass, with Arctic Cod as the primary food item along with Fourhorn Sculpins, Arctic Cisco, Pacific Rainbow Smelt, and Eelpouts. On the Yukon coast mysids (39.2%), amphipods (30.4%), isopods (22.8%), and fish (7.6%) comprise the average diet. In Tuktoyaktuk Harbour mysids predominate, followed by amphipods and isopods. Fish on the spawning migration in the Mackenzie Delta have empty stomachs.

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Osmerus dentex

Many fishes as well as seals and, in the Mackenzie Delta and southern Beaufort Sea generally, Beluga depend on smelt as a food. The Arctic Lamprey has been seen attached to this species at Pauline Cove, Herschel Island. Females live longer, grow faster, and are larger than males. The lifespan is up to 17 years on the Beaufort Sea coast of the Yukon. Beaufort Sea fish live longer and reach larger sizes than do the related O. mordax on the Atlantic coast. Alaskan and northern Canadian populations mostly mature at ages 4–7, often at 6–7 years and 20–22 cm. The youngest mature females in the Beaufort Sea may be as old as 10 years, the youngest mature males as 7 years. The spawning run in the Alaskan Beaufort Sea takes place at the end of July. The run usually starts when water temperatures reach 4°C or higher, just after the ice breaks up and moves out. The Mackenzie River spawning takes place under ice in April–May. Spawning occurs at night. The eggs are adhesive, up to 1.2 mm in diameter, and number up to 93,000 per female. The outer membrane of the egg ruptures but remains attached to the egg by a stalk. This outer membrane is the adhesive part, and the egg sways in the water at the end of the stalk. They become attached to vegetation or rocks and may be so numerous that some are smothered. As many as 190 eggs/sq cm have been counted. The fry are carried downstream to brackish water. The adults return to the sea after spawning during the openwater season.

western Pacific Ocean and across the Arctic Ocean of Siberia and Russia to the White Sea.

Distribution of Osmerus dentex

importance:  It is not economically important in the Canadian Arctic but has been an important Indigenous food in Alaska.

distribution:  It is found from Bathurst Inlet, Coronation Gulf,

Amundsen Gulf, and the Beaufort Sea coastwise around Alaska, and then south to Vancouver Island, and to Oregon, but is relatively rare in Pacific Canada. It is also found south to North Korea in the



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sources:  Johansen (1926); Kendel et al. (1975); Percy (1975); Slaney (1975, 1976a); Fraker, Sergeant, & Hoek (1978); Bond (1982); Luey, Kreuger, & Schreiner (1982); Craig (1984); Percy, Smiley, & Mullen (1985); Bond & Erickson (1987, 1989); Lacho (1991); D.B. Stewart, Ratynski, et al. (1993); Taylor & Dodson (1994); Mecklenburg, Møller, & Steinke (2011).

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Osmerus mordax (Mitchill, 1814)

Rainbow Smelt, éperlan arc-en-ciel

common names: Other common names are American Smelt,

Atlantic Rainbow Smelt, Atlantic Smelt, Bay Capelin, Boreal Smelt, Freshwater Smelt, Frost Fish, Leefish, Outside Capelin, Saltwater Smelt, Toothed Smelt, éperlan d’Amérique, and éperlan du nord.

taxonomy:  The genus comes from the Greek osmeros (odorous), equivalent to “smelt.” The species name comes from the Latin mordax (biting). description:  This species is distinguished by having medium conical to large canine teeth on the tongue, 1–3 large canines on each side of the vomer bone in the roof of the mouth, and lateral-line scales 51–73, with 13–30 pored. Separation from its relative, O. dentex, is given above, but distribution is the easiest character at present. Dorsal fin-rays number 8–11, anal fin-rays 11–18, pectoral fin-rays 9–14, and pelvic fin-rays 8. Total gill rakers number 26–37, and pyloric caeca 4–9. Males have numerous head, body, and fin tubercles, and a lateral ridge may be present. The mouth is large and reaches the rear of the eye. Paired and anal fins are larger in males, and a lateral ridge may develop. The colour of the back is olive green, steel blue, or black, and the flank is silvery with purple, blue, or pinkish iridescences (hence “rainbow”). The scales on the back are outlined with pigment, and there may be pigment spots on the anterior flank. There is a silvery stripe along the flank that turns dark in preserved fish. The peritoneum is silvery with dark pigment dots. Land-locked smelt are darker than other populations and may have dusky fins rather than the typical clear fins. The species reaches 35.6 cm in total length. habitat:  Habitat and biological information on marine Rainbow

Smelt is based on Atlantic Canada populations. The Hudson Bay and James Bay populations are new and have not been studied in detail. These fish school in coastal waters and enter fresh waters to

spawn. Some smelt spend their lives mostly in estuarine conditions. There are also land-locked populations in numerous lakes. They are found as deep as 150 m, perhaps to 425 m in the sea. Bay of Fundy fish move offshore in summer to deeper and cooler water and enter estuarine water in early winter to avoid cold water. This species has a biological anti-freeze that lowers the freezing point of body fluids below that of sea water (−1.9°C). In the St Lawrence estuary there are seasonal open-water migrations of 80–160 km.

biology:  Its food is a wide variety of crustaceans, squid, and small

fishes in the sea, and this species is voracious. Many fishes, as well as seals, depend on smelt as a food source, including such economically important species as Atlantic Salmon, Atlantic Cod, Inconnu, Burbot, Brook Trout, and Lake Trout. They are also cannibals. Females live longer, grow faster, and are larger than males. The lifespan is five to nine years, with maturity at the age of three years in Atlantic populations, although some fish mature one to two years earlier. Spawning migrations take place from February to June, later in northern seas. The run usually starts when water temperatures reach 4ºC or higher, just after ice break-up. Peak spawning lasts about a week. Males may be on the spawning ground for 14 days, and females for only one to three days. Males are first on the spawning grounds in fresh water, and spawning occurs at night. Two or more males flank each female, keeping station and contact in fast water aided by male breeding tubercles, and eggs and milt are released. The eggs are adhesive, up to 1.2 mm in diameter, and number up to 93,000 per female. The outer membrane of the egg ruptures but remains attached to the egg by a stalk. This outer membrane is the adhesive part, and the egg sways in the current at the end of the stalk. Eggs become attached to vegetation or rocks and may be so numerous that some are smothered. As many as 190 eggs/sq cm have been counted. The fry are carried down to brackish water after hatching. The hatching time varies from 51–63 days at 3ºC–9ºC to 8–10 days at 20ºC. Early spawning adults leave the stream to return to the sea before stream water temperatures reach 10ºC, and late spawners usually leave before temperatures reach 15ºC. The time of spawning is usually specific for each stream.

Osmerus mordax

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Osmerus mordax

importance:  This smelt is important in both sport and commercial fisheries in southern waters. The total Canadian catch in 1988 was 11,000 t, worth about $4.2 million. Smelt are in high demand by consumers in Japan and the United States. The value of flesh in 1985 was $0.28/kg. They are sold fresh, frozen, and precooked. Their impact on salmonids and other fishes in Hudson Bay, where they spread at around the turn of the millennium, has not been assessed but is likely to be deleterious. distribution:  Populations introduced to the Great Lakes have spread (or been spread) to the Hudson Bay drainage of northwestern Ontario and southeastern Manitoba. They have reached Hudson Bay at the Nelson River estuary and are reported to have entered the Churchill River by moving along the coast from the Nelson River against the prevailing currents. A specimen was caught at the mouth of the Moose River, Ontario, in 2011. They are expected to thrive and establish anadromous populations in Hudson Bay. They also enter rivers and are land-locked in lakes throughout eastern Canada and are found from southern Labrador throughout the Maritimes south to New Jersey (and perhaps Virginia) in the sea. sources:  Bigelow & Schroeder (1963); Luey et al. (1982); Ewart & Fletcher (1990); Franzin, Barton, Remnant, Wain, & Pagel (1994); Taylor & Dodson (1994); Remnant, Graveline, & Bretrecher (1997); Woehrmann (1997); Wöhrmann (1997); Zrum (1999); K.W. Stewart, Franzin, McCulloch, & Hanke (2001); K.W. Stewart & Watkinson (2004); D.B. Stewart & Lockhart (2005).



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Distribution of Osmerus mordax

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Family Salmonidae Trouts and Salmons, Truites et Saumons

James D. Reist ( with J. Brian Dempson, Karen Dunmall, Les N. Harris, Michael Power, and Heidi K. Swanson as authors for some species)

Salmonids are fishes found naturally throughout the northern hemisphere, although many species have been transplanted widely including to southern hemisphere locations. There are 11 genera with about 70 species worldwide, although great diversity exists within and among species, which contributes to a high degree of taxonomic uncertainty in some groups. About 42 species are found in Canada, 17 of these occurring in Canadian Arctic marine waters during part of their life history. They are primarily freshwater fishes, but many species exhibit anadromy, that is, reproducing in fresh water (typically in the autumn) but passing at least part of their life cycle in marine waters (typically for feeding and growth). Three different subfamilies are recognized: Coregoninae (ciscoes and whitefishes); Thymallinae (graylings, not in Arctic marine waters, and distinguished by a longbased dorsal fin, which is longer than the head); and Salmoninae (chars, salmons, and trouts); all three of which are elevated to family level by some workers. This family is distributed throughout Arctic Canada, with different species being regionally important. One species, Arctic Char, occupies the northernmost freshwater habitats in the world (about 83o N) and may be anadromous there; hence it is found in nearby marine areas. Currently the subfamily Coregoninae encompasses a widely distributed, relatively speciose group of freshwater and anadromous whitefishes. Coregonines can be distinguished from species in Salmoninae by the presence of larger scales (13 or less from the dorsal fin origin to the lateral line, and 110 or less in the lateral line), and teeth that are weakly developed or absent in the lower jaw, which is short, not extending back to mid-eye (except in Stenodus leucichthys). Within the subfamily Coregoninae, three genera are presently recognized: Prosopium (round whitefishes), Stenodus (a large piscivorous whitefish), and Coregonus (true whitefishes and ciscoes).

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The genus Coregonus has a long history of taxonomic complications and rearrangements attributed to the scarcity of fossil records, exceptional phenotypic plasticity and character displacement associated with co-habiting species, and the evolution of sympatric ecotypes resulting from secondary contact among glacial lineages. Within Coregonus the mid-water plankton feeders with lower protruding jaws (i.e., ciscoes or “lake herring”) were at one time placed in a separate genus (Leucichthys). Subsequent comparative osteological assessments submerged the subgenus Leucichthys within Coregonus. More recent molecular evidence has refuted this classification (i.e., indicated polyphyly of both subgenera), and presently all whitefishes and ciscoes are recognized in the genus Coregonus devoid of subgeneric classification. The subfamily Salmoninae is differentiated from Coregoninae by the small scales (19 or more from the dorsal fin origin to the lateral line, 115 or more in the lateral line); the teeth in the lower jaw being strong and conical; the long lower jaw extending back to or past mid-eye; and the presence of both orbitosphenoid and suprapreopercular bones. Salvelinus are differentiated from other genera in the Salmoninae subfamily by having pale spots or markings (e.g., vermiculations) above the lateral line (there are never dark spots in adults), no teeth along the posterior shaft of the vomer, and more than 175 scales in the second row above the lateral line. Salmonids are typically medium- to large-sized fishes (to 1.5 m in length) characterized by the following: an adipose fin; cycloid scales; an open connection of the gas bladder to the gut (physostomous); deep connection of the eye muscles through the posterior myodome to the trunk musculature; 11–249 pyloric caeca; 7–20 branchiostegal rays; soft-rayed fins without spines; dorsal fin positioned mid-body; abdominal position of pelvic fins; pelvic axillary process present; gill membranes far forward and free from isthmus; 50–75 vertebrae, with the last three turned up; strong lateral line; many species with strong teeth on jaws; various types of sexual dimorphism especially at breeding time that are strongly expressed in some species; parr marks present in young in most species and retained in neotenic adults of some life history types; and tetraploid karyotype. Anadromous fish, particularly salmonids, are ecologically pivotal as keystone species linking marine, freshwater, and terrestrial ecosystems. During their early life history, with freshwater residency of up to eight years in some species, significant amounts of nutrients originating from the freshwater ecosystem are incorporated into the fish. As fish smoltify and begin marine residency, these nutrients become available to nearshore and offshore components of marine ecosystems, for example piscivorous birds, predatory fish, and marine mammals. Considerable growth of anadromous individuals occurs at sea, and marine nutrients incorporated into the fish are transported back into fresh water during subsequent upstream migrations to reproduce or overwinter. These nutrients become available to the freshwater and terrestrial ecosystems as anadromous fishes expel reproductive products, die, or are preyed on by freshwater fishes, terrestrial mammals, and birds. Salmonids exhibit a highly variable life history among taxa as well as between and within species of each group. These life histories

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can be roughly grouped into two major categories: semelparity and iteroparity. Semelparous salmonids are fish such as Pacific salmons that reproduce only once in life, with all individuals dying immediately after such reproduction. Iteroparous salmonids are fish such as Arctic Char, most of which generally do not die after reproduction; rather, they overwinter as adults in fresh water, return to the sea the following year (or less often), and live to reproduce a number of times during their life. Such life-history variation has important consequences for population dynamics, abundance, and fisheries management. Within these general life-history categories, most species exhibit a variety of life-history types. Such types include the following (defined in terms of sexually mature adults): anadromous form (both sexes); co-occurring non-anadromous dwarf individuals (both sexes, but males dominate in some species), that is, freshwater residual or resident individuals of populations containing anadromous fish (residual individuals often exhibit neotenic traits once sexually mature); non-anadromous form (both sexes), which may co-occur with anadromous fishes during freshwater portions of their life history; and allopatric non-anadromous forms (i.e., isolated from anadromous forms by barriers such as waterfalls, distance, or land locking). For some species (e.g., Arctic Char), studies from Eurasia have indicated that switching from a freshwater residency to anadromy occurs frequently in a large portion of the local population. Thus, in some systems the following types co-occur: anadromous throughout life (once having smoltified initially); freshwater resident throughout life; and freshwater resident switching to anadromy. No documentation of a switch back to freshwater residency for the latter form has been made; thus, such changes appear to be unidirectional. Similar studies in Canada are lacking. However, within anadromous Arctic Char, recent evidence suggests a high degree of flexibility in how life history is conducted by individuals. For example, within a population most individuals exhibit anadromy that regularly involves annual migrations to the sea for summer feeding (assuming access is possible every year). However, some individuals of the same population exhibit anadromy for a few years, appear not to go to sea for the next few years despite having access, and then return to anadromous habits later in life. The degree, both within and among populations, and the implications of switching modes or strategies during life are unclear at present for Canadian species. Salmonids that exhibit semelparity tend to do so within fairly narrow life-history limits. As a result most fish found at a particular place or time (e.g., upstream migratory run) represent the same life-history stage and are of similar ages and sizes. In contrast, salmonids that exhibit iteroparity have highly variable life histories. This is exhibited as variable, and sometimes wide, ranges for key life-history transitions such as age of smoltification, age of first sexual maturity, and periodicity of reproduction. Thus, at any particular place or time during life history, fish of many different stages may be present. For example, anadromous char feeding at sea may consist of (a) post-smoltified, large juveniles with ages spanning three to five years and ranging widely in size; (b) adults becoming sexually mature for the first time; (c) adults becoming sexually mature again after some period of non-reproductivity; and (d) adults that



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have reproduced but are presently resting for one or more years and building energy reserves for the next reproduction. Fish in these different stages of life history also tend to exhibit differences in habitat use while at sea as well as in fresh water. For example, recently smoltified juveniles of many species likely remain closely associated with estuarine zones for the first year at sea, whereas older juveniles tend to move further away during summer feeding. This is likely due to the increasing ability of larger fish to tolerate progressively greater salinities, a trait influenced by surface-to-volume relationships, and is especially evident in species that overall are less tolerant of normal marine salinities (e.g., whitefishes). Within a species, adults that have sexually matured and reproduced at least once generally appear to be most tolerant of marine salinities and therefore are the most wide ranging at sea. Diversity in different stages of life history likely results in differential habitat use among stages within a species, theoretically reducing intraspecific competition for food. Diversity in life-history stages may confound species identification especially for younger stages of some species groups; for example, juveniles of whitefish species and cisco species can be easily confused while they are in marine environments. The relevance of the high biodiversity exhibited by salmonids in general and by Arctic salmonids in particular has a number of implications in the Arctic. First, in the Arctic, biodiversity at both taxonomic and functional levels is poorly documented and understood. This complicates taxonomic understanding and in many cases calls into question the definition of a species for some groups. This is especially true for many salmonids such as the chars and some whitefishes. It also means that we cannot predict the effects of anthropogenic impacts, because these are likely dependent upon the taxa present, their proportions in the ecosystem, and their roles in ecosystem structure and function. Second, flexibility in life history, regardless of whether it is manifested as diversity among types, stages, or individuals, likely is highly adaptive in environments such as the Arctic that are subject to extremes of seasonality and variability as well as trends over time. Such diversity enhances the probability of the persistence of populations in the face of environmental extremes by ensuring some individuals survive. Third, from the perspective of the management of salmonid fisheries, diversity of the types outlined above presents significant problems; it is impossible to “manage” for an average fish if the exploited population consists of many types with flexible life histories. Fourth, the apparent truism that biodiversity greatly decreases in polar regions may not be strictly so. In reality, for many Arctic ecosystems, high biodiversity at least within fish species may serve a function similar to that seen between species in more southerly ecosystems. High levels of biodiversity contribute to ecosystem complexity, thus facilitating energy cycling, and are more conducive to ecosystem stability and resilience to perturbations. Such biodiversity is also the basis for future evolutionary options, and present diversity may partially be the result of the Arctic environment. For fish such as anadromous salmonids that occupy marine, estuarine, and freshwater habitats, the Arctic is a highly variable and unpredictable environment at time frames within those of fish generations. The Arctic also has changed and is continuing to change over time. Thus, high levels of

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diversity are advantageous in ensuring the persistence of populations in such areas. As large juveniles and adults in the marine environment, most salmonids are pelagic fish, feeding primarily on food in the water column, which includes being piscivorous on other species as well as members of their own populations. Zooplankton and benthos are also significant components of the diet. The true whitefishes within the coregonines have a subterminal mouth and are primarily benthivorous, at least as adults. Marine feeding is seasonal and typically conducted in a short period in summer because of the following constraints. Pacific and Atlantic salmons have high salinity tolerances and thus, where present in the area, likely feed in both offshore and onshore zones. Chars have somewhat lower tolerances to salinity and thus tend to feed primarily in nearshore zones, but they likely are present in offshore zones in warmer and less saline surface waters. Whitefishes as a group have the lowest salinity tolerances and in the summer feed in close association with heavily freshened nearshore zones and estuaries of suitable Arctic rivers. Arctic marine waters are characterized by the seasonal and perennial presence of ice: land-fast ice onshore and pack ice offshore. Ice formation in northern marine environments results in two changes to the habitat in winter. First, the normal salt content of marine waters prevents freezing until about −1.8°C, and, second, the formation of large amounts of ice results in extrusion of salt into the water column, which in turn increases salinity below the ice and further depresses the freezing point. Land-fast ice also physically affects nearshore habitats by grounding out and scouring the bottom, which makes such areas unsuitable for overwintering fish. Anadromous salmonids are not tolerant of temperatures much below freezing or of hypersalinities. As a major consequence, all Arctic salmonids, unlike southern populations, are obligated to seek winter refuge in freshened areas (e.g., estuarine zones in rivers that continue to flow in winter) or in upstream freshwater areas. This is especially true for species that are less tolerant of high salinities such as whitefishes and chars. The winter ecology in Arctic marine waters for salmonids such as Pacific and Atlantic salmons that are more salinity tolerant is unclear. As a result of these environmental constraints and the iteroparous life history of many Arctic salmonids, Arctic species such as chars and whitefishes are obligated to undergo two seasonal bouts of osmoregulatory transition between fresh and salt water each year (spring and late summer or autumn) for many years. This presumably represents a major annual energy expenditure that may limit growth and reproduction. The consequences of such life-history constraints have been little studied in most Arctic anadromous salmonids in Canada. Many species are much sought after in sport, domestic food (subsistence), and commercial fisheries. Salmonids are the most significant group of fishes exploited by Arctic peoples globally, as well as in Canada. In the Arctic most exploitation occurs on anadromous life-history types in estuarine areas or in freshwater environments immediately upstream of estuaries during the autumn migrations. In Arctic Canada, exploitation of salmonids is important, but great regional differences exist depending upon the species present in the area. In the western portion of the Northwest Territories,

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coregonines of the lower Mackenzie River (five species) are the most heavily exploited group in subsistence fisheries and have occasionally been commercially exploited. In the eastern portions of the Northwest Territories and Nunavut, Arctic Char is the most exploited taxon and supports most subsistence fisheries and several commercial fisheries, as well as being a much sought-after sport fish. The Atlantic Salmon supports substantial fisheries of all types in northern Québec and Labrador. Chars (Arctic Char and Brook Trout) and whitefishes are locally heavily exploited, primarily in subsistence fisheries, where these species are abundant in Hudson and James Bays. These fisheries confer cultural, social, and economic benefits on northerners as well as Canada as a whole. Subsistence fisheries based on anadromous fishes provide a calculated economic benefit to the north of about $6,000,000 (about 40% of the total estimated benefit of all subsistence fisheries; estimates are for 1988 dollars). Such figures do not include the cultural or social benefits that contribute to the maintenance of traditional lifestyles, self-sufficiency, and quality of life of northern peoples. Commercial fisheries for Arctic Char, Broad Whitefish (Coregonus nasus), and Atlantic Salmon are conducted either for local trade or as export fisheries in which the product is sold in the south. In 1987–8, together these accounted for an additional direct economic benefit of about $160,000, which, though small, contributes significantly to local economies. A further gross benefit (i.e., value added to the economy) of about $659,000 resulted from commercial export fisheries. Sport fisheries primarily exploit Arctic Char and Atlantic Salmon, with the former being the most wide spread and significant. Sport fishing for Arctic Char in 1985 was conducted from 23 lodges, accounting for 172,692 angler days (7% of total days for all of the then Northwest Territories including present-day Nunavut). Arctic Char was the most preferred species, with about 16,500 fish caught. The direct economic benefit from this sport fishing is estimated at $300,000. Additionally, angling for anadromous fishes is estimated to have had a gross benefit of about $1,600,000. Thus, the total estimated benefit to the Canadian economy of Arctic anadromous fisheries for the late 1980s was about $8,719,000. More recent summaries are not available, but this value has almost certainly increased since that time. There is an extensive literature on these fishes, in both research papers and popular outlets. This literature cannot all be cited here, but the former is given in the bibliography.

sources:  Dymond (1933, 1963); Bigelow (1963a, 1963b, 1963c,

1963d); Lindsey & Woods (1970); Marshall & Woods (1971); Salonius (1971, 1973); Hunter (1974); Balon (1980); Lindsey (1981b); Dodson, Lambert, & Bernatchez (1985); Bodaly (1986); Reist & Bond (1988); Kawanabe, Yamazaki, & Noakes (1989); Kemp, Bernatchez, & Dodson (1989); Kemp & Dodson (1990); Groot & Margolis (1991); Chiperzak (1992); Clarke (1993); Stearley & Smith (1993); Willson & Halupka (1995); Osinov (1999); Watson (1999); Quinn (2005); Reist & Sawatzky (2010); Nielsen, Ruggerone, & Zimmerman (2013); Reist, Power, & Dempson (2013).

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Coregonus artedi Lesueur, 1818

Cisco, cisco de lac JAMES D. REIST

common names: Local names are Arnaqsleq, Kapisilik, Kaviselik, and Kavisilik (Inuktitut); Anmagiak, Armagiak, and Qaluhaq (Inuvialuktun) – although several of these names likely refer to ciscoes collectively – and Nùtimĭwàsù, Nùtimĭweshish, Nùtimĭwesù, Uchùlipĭsh, Utùlipĭ, and Utùlipĭsh (Cree). Other common names are Bear Lake Herring, Blueback, Blueback Tullibee, Freshwater Herring, Grayback Tullibee, Herring Salmon, Lake Cisco, Lake Herring, Sand Herring, Shallowwater Cisco, and Tullibee. It is probably also grouped with riverine ciscoes in the western Arctic collectively as “herring,” with Pacific Herring being distinguished as Blueback Herring. taxonomy:  The genus comes from the Greek core (pupil of the eye) and gonia (angle). The species is named after the Swedish naturalist Peter Artedi (1705–35), called the Father of Ichthyology because Linnaeus used his notes for fish descriptions. First described as Coregonus Artedi from Lake Erie and Lewiston, Niagara River, this New World cisco exhibits considerable taxonomic variation within and among locations, and thus it is often referred to as a complex. This situation is exemplified by the listing of up to 22 subspecific forms being present within the Laurentian Great Lakes. As for other North American ciscoes, this taxon has

been variously associated with Argyrosomus Agassiz, 1850, and Leucichthys Dybowski, 1874, at the generic level; including Coregonus, the specific epithet for these three was for many years artedii, recently corrected to artedi. A variety of synonymous taxa involving all these names and additional specific epithets thus results. Only synonymies potentially referring to northern taxa are listed here: Salmo tullibee Richardson, 1836, described from Cumberland House, Pine Island Lake, Hudson Bay; S. harengus Richardson, 1836, described from Penetanguishene, Ontario; Leucichthys nueltinensis Fowler, 1948, described from the northwestern extremity of Nueltin Lake at Windy River, southwestern Keewatin, Northwest Territories; and L. churchillensis Fowler, 1948, described from the Churchill River at Churchill, Manitoba. Some authors also include three taxa described by Harper and Nichols (1919): Coregonus athabascae from Lake Athabasca at the mouth of Charlot River, northern Saskatchewan; C. entomophagus from the Tazin River at the foot of Kolethe Rapids, Mackenzie District, Northwest Territories; and C. macrognathus from Great Slave Lake near Fort Resolution, Mackenzie District, Northwest Territories. However, these are more properly synonyms for a different taxon – Shortjaw Cisco, C. zenithicus (Jordan and Evermann, 1909) – found in inland lakes only. Some authors also consider other ciscoes that are typically recognized as distinct species as part of this taxon (e.g., some genetic evidence suggests that inland North American ciscoes are all part of one lineage). Within-system (lake or river) variation in Cisco in the narrow sense also exists (e.g., ecomorphs in lakes, life-history variants in rivers); similarly, among-system variation of the same putative taxon is present, leading to difficulties in aligning individuals from various locations with a particular taxon. Accordingly, as a reflection of these difficulties, the Cisco, C. artedi

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complex, can be viewed as pertinent at two levels, that referring only to Cisco and close derivatives (sensu stricto) and that referring to Cisco and its wider taxonomic relatives (sensu lato). Superimposed upon this there appears also to be life-history differentiation (e.g., anadromous, lacustrine, and riverine or fluvial types). In some areas of the Canadian marine Arctic it appears that these forms possibly co-occur (e.g., Hudson and James Bays) and also occur with other anadromous ciscoes in other areas (e.g., Beaufort Sea area). Thus, careful identifications are required when examining marine occurrences that are suspected to be this species. Pleistocene glaciations undoubtedly resulted in significant diversification, rearrangements of taxa (e.g., through extirpation and hybridization), and episodic and uneven recolonizations of deglaciated areas. These processes continue presently, contributing to ongoing taxonomic problems. Resolution of the taxonomic complexities appears to defy present approaches; despite much recent work over parts of the range, neither a complete picture regarding the number of distinct taxa nor their recognition at specific or various subspecific levels has yet emerged. The common name of Lake Cisco has been dropped, and this species is now known only as Cisco.

description: This species is distinguished from its congeners in Arctic Canada by having the profile of the upper lip sloping backwards in line with the forehead; the gape of the mouth is not vertical; the tip of the lower jaw is equal to or projects slightly beyond the upper jaw; scales between the dorsal fin origin and the lateral line number 7–10; and the chin and the pelvic fin tips are speckled or dark. The Cisco is an elongate coregonine, round in cross-section, with the greatest depth being in front of the dorsal fin. The head is around 20%–24% of total length. The eyes are relatively large, about 21%–26% of the head length. The maxilla extends to the anterior edge of the pupil. The pelvic fins tend to be far back on the body (the snout–pelvic origin distance falls on caudal fin-rays, differentiating this species from Least Cisco). Dorsal fin-rays number 9–15, anal fin-rays 10–15, pectoral fin-rays 14–18, and pelvic fin-rays 8–12. Lateral-line scales number 63–94. Gill-raker numbers range widely (36–64), likely a reflection of wide geographic range and ecology; usually there are 40–50 relatively long and slender rakers. Males have one nuptial tubercle in the centre or near the edge of each scale. The coloration overall is silvery with bluish to greenish hues above, whitish or silver on the sides with pink to purplish iridescences, and white on the belly. The anal and pelvic fins tend to be light in colour and typically with black sprinkling or tips (differentiating this species from Arctic Cisco where they co-occur). Other fins have dark pigment especially along the outer halves or margins. As with other ciscoes, dark fin pigmentation usually is present in older fish, and species identification of fish less than about 10.0 cm is very difficult. The characters above are derived primarily from lacustrine forms in the centre of the range; no detailed description of the anadromous form is available, and thus some differences may be present in this life-history type. The species attains about 57.2 cm in total length in some lakes, while northern anadromous populations are somewhat shorter, attaining lengths of 35.0–37.5 cm. Its weight reaches 3.63 kg.

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habitat:  The Cisco is primarily lacustrine, occupying lakes and

ponds of all sizes suitable for fishes, although it may also be found in larger rivers. River habitat requirements are not known. In lakes, Cisco are mid-water pelagic and benthopelagic fishes. Young often inhabit the shallow waters of protected bays over rocky substrates and are usually associated with vegetation. They school with whitefish in these areas until they are approximately one month old. By summer they have moved to deeper water and become pelagic. They are typically found at depths of 10–60 m, moving from shallow waters in spring to deeper waters in summer. Diel migrations are also made away from shore at sunrise and towards shore at sunset. Adfluvial populations are known, which utilize tributary rivers for spawning and downstream lakes for feeding and growth. Details of their migratory patterns and nearshore marine habitat use in much of the Arctic are generally unknown scientifically, although local peoples likely understand them quite well. The exception is the rivers flowing into James Bay where out-migration of Cisco larvae occurs in the spring into marine waters, followed by autumn in-migration to overwinter in freshened river mouths. Thereafter, seasonal migrations to feed in the marine system and overwinter in fresh water occur to maturity. Details of adult usage of marine waters are unclear; adults appear to remain mostly associated with freshwater habitats; however, some information suggests alongshore movements during summer between the Nelson and Churchill estuaries of Manitoba. This pattern is similar to that observed for other coregonines in the Mackenzie River area of the Beaufort Sea; however, Cisco appear to be less migratory there.

biology:  Its diet generally consists of plankton, but both benthic

and aerial prey are consumed in fresh water. Its marine diets are generally unknown but likely parallel those of other ciscoes. Strongly swimming prey is taken by a dart-and-suck action. Gulping is also used, in which the fish opens and closes its mouth two to three times per second, taking more than one prey item at a gulp. Ciscoes are preyed upon by a wide range of piscivorous fishes in fresh and marine waters; again, information on marine predators is limited. The latter probably include marine mammals and birds as well as co-occurring fishes. Anadromous Cisco in the James Bay area live to around 13 years, growing to about 37.0 cm in length. Growth is latitudinally dependent, declining in more northerly locations along the southeast Hudson Bay coast. Females mature at five years, and males at six years; this parameter also increases with latitude such that more northerly females mature at age seven (although limited samples may bias these conclusions). The maximum age for the species is 31 years. Several life-history forms exist including anadromous, lacustrine, and adfluvial. Whether wholly riverine (fluvial) forms also exist is unknown. Anadromous forms appear to follow a general life history parallel to that of other anadromous coregonines by spawning and overwintering in freshwater areas, followed by a seasonal pattern of summer marine feeding and return migrations to overwinter in fresh or freshened estuarine waters. Unlike most other coregonines, except for Arctic Cisco, larvae appear to be very tolerant of saline waters and therefore use marine areas earlier in life

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history, at least in James Bay. Further north in Hudson Bay this does not appear to be the case, and the possibility of spatially disjunct, juvenile overwintering areas exists. Annual marine sojourns continue to maturity, whereupon adult fish remain associated with fresh water or highly freshened estuaries. Cisco are iteroparous fall spawners for the most part; however, some indications exist for spring-spawning populations in fresh water. Spawning occurs in lakes in the fall, most often over sand and gravel substrates and occasionally over boulders, rubble, clay, mud, and vegetation. Cisco are generally shallow-water spawners (1–5 m) but have also been known to spawn at depths between 120 m and 140 m. Males arrive on the spawning grounds first, two to five days before females. As many as 12 males will follow each female, but at spawning time the female is accompanied by two males. The female descends to 15–20 cm above the bottom, leading the males whose heads are level with her anal region. The eggs are shed, mostly at night, fall to the bottom, and are abandoned. They are slightly adhesive and become attached to the bottom. Ripe females produce up to 29,000 eggs of 2.1 mm diameter. The eggs hatch in spring before the ice break-up. Evidence of skipped spawning (i.e., mature individuals resting for a spawning period) exists and also appears to be latitudinally dependent (i.e., more frequent in more northern locations). Fecundity is not latitudinally correlated; it and growth appear to be similar to those of Cisco occupying cold oligotrophic lakes at similar latitudes, a reflection of colder conditions and shorter growing seasons in northern marine and estuarine environments.

west in the Mackenzie River Delta area. The Cisco is mostly confined to the mainland, and a record from the Thomsen River on northern Banks Island (Wilkinson et al., 1977) is an error for C. sardinella (Stephenson, 2010). The Cisco complex (both sensu stricto and sensu lato) is strictly a New World taxon that appears to have originated in the centre of the continent during Pleistocene times; recolonizations from the upper Mississippian refugium south of the present-day Laurentian Great Lakes appear to have occurred through proglacial lakes to both the northwest and the northeast. Presently Cisco are distributed from the Great Lakes basin through the northern portions of the three Prairie provinces and Ontario and Québec. Distribution in the Northwest Territories extends primarily to the lower Mackenzie River basin and episodically to the coastal areas of the southern Beaufort Sea, where it may co-occur with other anadromous ciscoes. It occurs throughout much of Manitoba, Ontario, and western Québec north through Nunavik to Hudson Strait drainages, and eastwards to the western margin of Ungava Bay. Thus, it occurs along the entire coasts of Hudson and James Bays to western Ungava Bay. In this area there are no co-occurring ciscoes as found in the western Arctic; therefore, identifications of marine specimens should be reasonably straight forward. Throughout many of these coastal areas Cisco are likely restricted to freshwater locations; however, in estuarine and freshened nearshore areas close to large rivers they may be locally common.

importance:  Freshwater Cisco populations are the basis of

significant fisheries throughout their range. The anadromous populations are also fished by local peoples, particularly upstream migrations (i.e., runs) of spawners. Some spring fisheries occur through land-fast sea ice prior to break-up; similarly some coastal locations are fished during summer. As with other Arctic coregonines, fisheries may also occur at inland locations, particularly where the fish congregate to spawn or at obstructions such as falls. Statistics are difficult to accumulate because either all ciscoes occurring in an area are locally grouped together (e.g., the Beaufort Sea and Mackenzie River region) or ciscoes and whitefishes are grouped (e.g., James and Hudson Bays). For southwestern Hudson Bay and western James Bay, in 1990 around 45,000 whitefishes (which included Cisco) were reported to have been harvested by fishers from seven communities (projected estimates indicate that up to 58,000 fish may have been harvested). Regardless, similar to other salmonids, the Cisco figures prominently in local subsistence harvests and socio-cultural aspects of Arctic Indigenous peoples. No commercial harvests targeting anadromous populations of Cisco are known, although those Cisco may occur as by-catch in harvests for other species.

distribution:  It is found in Ungava, Hudson, and James Bays; southern Foxe Basin; Bernier Bay on Baffin Island (Ellis, 1962); Pelly Bay; Rasmussen Basin; Coronation Gulf; and the Beaufort Sea. It also occurs along the northern continental coast from the Boothia Peninsula westwards to about Darnley Bay, then disjunctly further



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Distribution of Coregonus artedi

sources:  Harper & Nichols (1919); Ellis (1962); Wilkinson et al. (1977); R. Morin, Dodson, & Power (1981, 1982); Berkes (1982a); A. Kemp, Bernatchez, & Dodson (1989); Bernatchez & Dodson (1990a); Berkes et al. (1994); Stephenson (2010).

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Coregonus autumnalis (Pallas, 1776)

Arctic Cisco, cisco arctique JAMES D. REIST

common names:  Local names are Kaktak, Kapisilik, and

Kraaktak (Inuktitut); Amagiak, Anma, Anmagiak Piqquaqtitaq, Kapihillik, Qaaktaq, Qaluhaq, and Qanktaq Siirgarq (Inuvialuktun); Treeluk (Gwich’in); and Qaaktaq and Qaatag in northern coastal Alaska. Local names in the western Arctic refer collectively to all riverine cisco species found in the area. Other common names in Arctic Canada are Herring, Blue Herring, and Salmon-Herring, confusingly as these may also refer to Pacific Herring.

taxonomy:  The species comes from the Latin autumnus (autumn) and –alis (pertaining to), referring to the fall congregations into migratory runs. It was first described from western Siberian waters and associated with Salmo and is often confused with its closely related congener the Bering Cisco, C. laurettae Bean, 1881; however, taxonomically these are distinct (see notes below). Several taxa aligned with this species have been described from Eurasia, but uncertainty exists as to the actual composition and relationships of the various subspecific taxa; for example, C. autumnalis migratorius (Georgi, 1775), the Baikal Omul, is now recognized as a distinct species and is genetically a form of the C. lavaretus complex despite showing convergent morphological characters with ciscoes. The Penzhina Cisco, C. subautumnalis Kaganowsky, 1932, from northeastern Asia appears to be either a variant form or a very closely related species of Arctic Cisco. The situation with the Irish Pollan, C. pollan Thompson, 1835, disjunctly isolated in Irish loughs (lakes), is similarly uncertain. Within the coregonines, ciscoes generally exhibit a terminal mouth associated with feeding in the water column; however, convergence with whitefishes (which generally have subterminal mouths) occurs in many locations. Terminal mouths and other shared (but likely non-homologous) characters led to the association of all ciscoes in Leucichthys, considered variously a subgenus of Coregonus or a genus. This taxon is now demonstrated to be of mixed composition, thus invalid. Northwestern North American ciscoes (so-called Old World ciscoes – C. autumnalis, C. laurettae, and C. sardinella) genetically represent a distinct group from inland “New World” ciscoes (i.e., C. artedi and related nominate forms). Within the former group, some question the validity of the three species, but taxonomic, genetic, biological, and life-history differences are all apparent. Unlike other ciscoes, there appears to be only one form of Arctic Cisco in North America exhibiting coastal anadromy. description:  This species is distinguished from other congen-

eric coregonines in Arctic Canada by the gape of mouth not being vertical; the tip of the lower jaw does not project beyond the upper jaw; scales between the dorsal fin origin and the lateral line are 11–12;

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the chin and pelvic fin tips are pale; the body lacks black spots, and the fins lack white spots; and gill rakers on the lower limb of arch number 26–31 (41–48 total). The Arctic Cisco is a pelagic coregonine with an elongate but slightly compressed body and a moderately sized head (about 24% of total length). Arctic Cisco are also more terete (elongate) than Bering Cisco. The moderate mouth is terminal, with upper and lower jaws of equal size (the lower jaw projects slightly in Cisco and Least Cisco). The maxilla extends to below the middle of the eye (differentiating this species from Least Cisco). Dorsal fin-rays number 9–12, anal fin-rays 9–14, pectoral fin-rays 14–17, and pelvic fin-rays 11–12. Lateral-line scales number 82–111. Gill rakers number 26 or more on the lower limb (distinguishing this species from Bering Cisco with < 25). Pyloric caeca number 113–183. Males develop nuptial tubercles on the flank scales. The coloration overall is silvery with greenish or brownish hues on the back, and silver or white on the sides and belly. Paired and anal fins are immaculate or with very light dusting of pigment (cf. the dark fin tips on Least Cisco and Cisco). There is no dark spotting on the head, the back, or the fins (as found in Least Cisco). Individuals of all cisco species of less than 15.0 cm in length each generally exhibit similar characteristics, which are less well defined than in adults, resulting in difficulties of identifying young fish. The species attains about 45.0–50.0 cm in total length, up to perhaps 64.0 cm, and 7.0 kg in weight.

habitat:  Spawning occurs upstream in the freshwater reaches of

tributaries to the Mackenzie River. Young emigrate to the delta and estuarine areas and enter nearshore habitats early in life (ages 0–3 years), although some may be flushed into delta lakes and entrapped until the following spring freshet. Emigration, aided by wind-driven coastal currents, occurs along shores westwards to the Colville River of Alaska, and perhaps eastwards as well, but to a lesser extent. Summer feeding areas for all life stages include these coastal areas; those used early in life history presumably are highly freshened. As fish age, they likely occupy more saline areas and perhaps offshore freshened surface waters for summer feeding. Hypersalinities combined with cold temperatures require migrations to, and overwintering in, freshwater habitats. These appear to include the lower reaches of rivers that maintain winter flows, and adjacent estuarine zones that remain oxygenated and highly freshened. There are few such locations on the continental north coast (e.g., Colville, Mackenzie, and Anderson Rivers), and all appear to be used in this fashion by some segments of these and other coregonine populations.

biology:  Its diet consists of marine crustaceans and other

invertebrates, as well as small fishes. Recent stable isotope studies suggest that feeding occurs at relatively high trophic levels for a coregonine. Arctic Cisco exhibit a single life history in Arctic North America, that of anadromy. Siberian lacustrine populations that have been attributed to this species have recently been shown to be allied to different species. Despite a wide overall range in North America, and numerous studies, only one complex of indistinct breeding populations is known from the Mackenzie River basin. Perhaps

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Coregonus autumnalis

others exist in nearby suitable rivers (e.g., Anderson River and its tributaries); however, definitive studies are lacking. The maximum lifespan is about 19–21 years, with sexual maturity at 5–10 years. Arctic Cisco exhibit iteroparity. Reproduction occurs in the Arctic Red, Peel, Great Bear, Carcajou, and Mountain Rivers of the Mackenzie basin. Eggs are shed over gravel in the fall in fast current and may number up to 90,000 per female, with a diameter of 1.1 mm. Age-0 fish emerge the following spring under the river ice and are flushed downstream into the Mackenzie Delta. Wind-driven coastal currents aid westbound movements by young fish to the Colville River (i.e., 80% of variation in recruitment to fisheries in a particular year depends upon the strength and duration of easterly summer winds in the nearshore Beaufort Sea lagged the appropriate number of years). Fish rear in the Colville system to around age seven or eight years and make annual feeding forays to the lower reaches of the river and nearby coastal waters. At the onset of sexual maturity seven- to nine-yearold fish migrate eastwards to the Mackenzie River and join mature adults to participate in late summer upstream migrations to spawning areas. Once mature, individuals remain associated with the Mackenzie River basin and nearby shelf areas of the southern Beaufort Sea, annually migrating between marine feeding areas and freshwater spawning areas. Upon spawning, adults likely move downstream to areas in the lower river and the delta to overwinter. It is unknown whether adults, especially females, consistently skip spawning, as seen in other anadromous fishes. The migratory scenario developed above is best understood for the Mackenzie basin and northern Alaska. Wind-aided recruitment (i.e., persistent summer easterly winds) to western areas such as the Colville River is highly correlated with a fishery yield of large subadults lagged seven or eight years. In years when winds are persistently from the west, recruitment to Alaskan fisheries is low. The converse expectation – that is, recruitment to easterly locations in years with high westerly



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winds – appears likely but has not been adequately tested. Large rivers and embayments (e.g., Wood Bay, Anderson River to the east of the Mackenzie Delta) may contain populations as well, but their distinctiveness from the Mackenzie River complex and their life histories are unknown.

importance:  Arctic Cisco are culturally important to western Arctic Indigenous peoples in both Canada and Alaska, where they support substantive subsistence fisheries. This is especially so along the coasts, but harvests also occur inland of migrating adults of this species and Least Cisco. Collectively in Canada these fishes are known as “herring” by local peoples. Harvests traditionally have been quite large for consumption by both humans and dogs. They are tasty when smoked because they are a fatty fish on the migration. Commercial fisheries also occur in Alaskan coastal locations (especially the Colville River) primarily during autumn and early winter once freshwater ice has formed and when the fish return to freshwater areas to overwinter. This fishery targets large subadults (6–8 year olds) from the Mackenzie system (see the “Biology” section). Catch data, corrected with the coastal wind patterns experienced by 0–2 year-old fish, support the Mackenzie-origin hypothesis of life history; thus, this species is truly a trans-boundary taxon. Catches and catch-per-unit effort in the Alaskan fisheries are highly variable, related in part to coastal wind patterns and in part to local environmental conditions in the river mouths (especially salinities). Estimates of total numbers of fish captured annually range from a high of about 80,000 (1993) to about 5,000 (2001). Climate-driven shifts may significantly affect this species and harvests in the future. Arctic Cisco are a keystone component linking freshwater, estuarine, and marine ecosystems through horizontal transfers (i.e., migrations) of material and energy. These processes (and thus the fish themselves) may be particularly important in Arctic ecosystems with low productivities.

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Coregonus autumnalis, juvenile

distribution:  It is found in the Rasmussen Basin, Chantrey Inlet, around the Adelaide Peninsula and Prince William Island, Queen Maud Gulf, Melville Sound, Bathurst Inlet, Coronation Gulf, Dolphin and Union Strait, eastern and western Victoria Island, Amundsen Gulf, and the Beaufort Sea. This species is associated with larger north-flowing Arctic rivers from Russian waters (e.g., Pechora River) eastwards through Siberia and Chukotka, and from northern Alaska and Canada to the Chantrey Inlet area (Kitikmeot region of Nunavut, west of the Boothia Peninsula). They are most common in central and downstream areas of large rivers, but, as seen from life history, their occurrence is seasonally related. They are less common at geographic-range peripheries, including occurrences on the southern edges of the western Arctic Archipelago (i.e., Banks Island) and Victoria Island. Inland distributions are uncertain but generally are common to several hundred kilometers from river mouths. Occurrences reported east of the Boothia Peninsula are likely misidentified Cisco (C. artedi); those on the northern edge of Victoria Island require confirmation and may be misidentified Least Cisco. As noted in the “Habitat” section, summer occurrences in freshened marine areas are mostly coastal, but they are also found on nearby shelves. Bering Cisco, Coregonus laurettae Bean, 1882, present in western Alaska and on the Alaskan north coast eastwards to the Colville River, has been suggested to occur in Canadian Beaufort Sea waters. Coastal research on the Yukon North Slope and Northwest Territory waters west of the Mackenzie Delta has failed to capture any specimens despite explicit searching. This taxon, accordingly, has been excluded from the Canadian Arctic marine fish fauna.

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Distribution of Coregonus autumnalis

sources:  McPhail (1966); Alt (1973a); Craig & Mann (1974); Fechhelm & Fissel (1988); Reist & Bond (1988); Bond & Reist (1989); Dillinger (1989); Fechhelm & Griffiths (1990); Edge (1991); Chereshnev & Skopets (1992); Dillinger, Birt, & Green (1992); Turgeon & Bernatchez (2003); Sukhanova, Smirnov, SmirnovaZalumi, Kirilchik, & Shimizu (2004); Yakhnenko, Mamontov, & Luczynski (2008); C. George, Moulton, & Johnson (2009); Moulton, Seavey, & Pausanna (2010); Zimmerman et al. (2013).

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Coregonus clupeaformis (Mitchill, 1818)

Lake Whitefish, grand corégone JAMES D. REIST

common names: Local names are Anadleq, Anâdlerk, Anahik, Jikuktok, Kakiviaktok, Kakiviartût, Kakkiviartoq, Kapihilik, Kapisilik, Kaviselik, Kavisilik, Keki-yuak-tuk, Pi-kok-tok, Pikuktuuq, and Qelaluqaq (Inuktitut); Anakheek, Kapihilik, Pikuktung, Pikuktuq, Pikuktuuq, and Qalupiaq (Inuvialuktun); Dalts’an and Dalts’in (Gwich’in); and Atihkamàkw and Atihkamekw (Cree). Other common names are Buffalo Back, Crookedback, Common Whitefish, Eastern Whitefish, Gizzard Fish, Humpback, Inland Whitefish, Labrador Whitefish, Lake Herring, Sead, Whitebait, and Whitefish. taxonomy:  The species name comes from the Latin clupea (her-

ring), forma (shape), and –is (having). Coregonus atikameg Bajkov, 1933, described from Churchill, Hudson Bay, and C. atikameg manitobensis Bajkov, 1933, described from Northern Manitoba, are synonyms. Lake Whitefish are one taxon of the humpback whitefishes that form a species complex, often termed the Coregonous clupeaformis complex, which in northern North America includes the Lake Whitefish, the Alaskan Whitefish (C. nelsonii Bean, 1884), and the Humpback Whitefish (C. pidschian (Gmelin, 1789)). Gill-raker

counts are considered the best meristic characteristic for separating species in the complex, with C. clupeaformis usually having 26 or more, C. nelsonii having 24 or 25, and C. pidschian having 21–23. Variation caused by environmental modification, however, may confound simple classification on the basis of gill-raker counts alone. Range separation may also help to distinguish among members of the complex, with C. clupeaformis being found throughout much of Canada and parts of the United States, C. nelsonii being found largely in the Yukon River basin, and C. pidschian being found in most Alaskan rivers that drain to the Bering, Beaufort, and Chukchi Seas. Recent morphological and genetic studies have further subdivided the complex, recognizing five groups within the previously described forms: Mississippian Lake Whitefish, widely distributed across Canada; Alaskan Whitefish, Beringian Lake Whitefish, and Nahannian Lake Whitefish, found primarily in central Alaska; and Humpback Whitefish, distributed along the north and west coasts of Alaska. Genetic comparisons to the closely related Broad Whitefish (C. nasus) have indicated that none of the five forms was sufficiently different to be considered a separate species, but, given the pattern of distributional separation, they should be treated as subspecies within a complex. In eastern and central Canadian Arctic waters the Mississippian form appears to dominate and thus be present in adjacent marine areas. In western Canadian Arctic waters, particularly those of the Peel River draining into the Mackenzie basin, a mixture of forms is likely present (Mississippian, Alaskan, Nahannian) in freshwater areas, with the Beringian form penetrating eastwards along the coasts (and perhaps upstream in the lower Mackenzie). Thus, marine migratory Lake Whitefish in this area likely consist

Coregonus clupeaformis



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of at least two forms. External morphology among forms is similar, and so discriminating these in marine areas is difficult, if not impossible.

description:  This species is distinguished from its congeners in Arctic Canada by the profile of the upper lip being vertical or overhanging; the forehead is concave in profile; the adult usually has a hump behind the head; the maxilla length is more than twice its breadth; total gill rakers usually number 24–33; and the length of the longest gill raker is more than one-fifth of interorbital width (1.32%– 1.97% of standard length). Lake Whitefish have an elongate, laterally compressed body that is greatest in depth immediately anterior to the dorsal fin and more terete than that of Broad Whitefish where these species co-occur. Large fish develop a hump on the back, particularly in northwestern Canada. The head is short in comparison to the body length, and the eye diameter is small relative to the head. The snout comes to a rounded point and overhangs the maxillary, which extends posteriorly to the anterior edge of the eye. The snout of Lake Whitefish is longer and more pointed than that of Broad Whitefish. The mouth is slightly inferior. Teeth are not readily apparent in either the mouth or the jaws. Scales are large and cycloid and vary in number from 70 to 97 along the lateral line. Meristic characteristics, particularly the gill-raker number (the total-count range is 19–37), vary among populations. Gill rakers number 19–33, usually 24–33, dorsal fin-rays number 10–13, anal fin-rays 10–14, pectoral fin-rays 14–17, pelvic finrays 10–12, pyloric caeca 140–222, and vertebrae 55–64. At spawning time sexual dimorphism is evident, with males developing hard, white, conical nuptial tubercles on the lateral scales, particularly those immediately above (three or more rows) and below (six rows) the lateral line. Each scale has one central tubercle with smaller tubercles on each side or on one side on some scales. Tubercles develop to a lesser extent on the head. Tubercles are also less well developed or absent on females. Whitefish vary in colour from dark brown to greenish in dorsal areas, and silver to yellow in ventral areas. The overall appearance may be silvery in some waters, and the flanks may have a bluish tinge. Scales are outlined with pigment in northwestern Canada. Fins can be clear in the south, to black-tipped in the north. The species reaches 80.0 cm in total length, possibly 125.0 cm, and 19.05 kg in weight. habitat:  The Lake Whitefish is a cold-water stenothermic species.

Over most of the southern part of its range Lake Whitefish descend to cooler hypolimnetic waters during the summer. In the north where waters are cooler, Lake Whitefish can be found in both littoral and pelagic habitats. In lakes juveniles are usually found in shallow surface waters (< 1 m) in areas with emergent vegetation and woody debris over cobble and boulder substrates that provide protection from predators. As temperatures warm seasonally, juveniles migrate to cooler, deeper waters. As adults, lacustrine Lake Whitefish are primarily bottom feeders and generally associated with habitat areas where benthic prey may be obtained. Planktonic feeding in adult Lake Whitefish does occur. In the north, anadromous Lake Whitefish inhabit the estuarine areas of large rivers – e.g., Lake Whitefish

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are a common component (6%–16% of the catch) of estuarine net sets in the estuaries of rivers that are sampled along the eastern coasts of James and Hudson Bays – and are caught in waters with salinity of up to 20.5‰. Other studies have noted downstream spring migrations of larvae to river mouths; eventual occupancy of the saltwater zone as juveniles; and repeated spring-fall, downstream-upstream migration patterns as fish mature. Juveniles remain longer in the marine environment, often being found in concentrations at river mouths in the autumn. Returning larger and more mature, adults migrate directly upstream to spawning grounds for the autumn spawning period. As mature adults, Lake Whitefish move further from natal rivers, and longer-range (16–43 km) movements have been documented along coastlines. It has been speculated that Lake Whitefish may move further offshore than recorded catches indicate; however, similarly to most other coregonines, this is likely not extensive and may be associated primarily with river plumes. In the western Arctic, Lake Whitefish have been found in coastal areas throughout the summer but have not been captured in abundance; similar to Broad Whitefish, they apparently do not undertake extensive coastal migrations. More recent investigations of anadromy in Hudson Bay, using strontium stable isotopes, have indicated that Lake Whitefish originating from the Nastapoka River estuary remain largely within the influence of the freshwater plume, with only a minority of fish venturing into higher-salinity waters.

biology:  Lake Whitefish have been extensively studied throughout their range but primarily in freshwater habitats. They may occur as adfluvial, lacustrine, or anadromous life-history types, but the correspondence of these with the various postglacial lineages is unknown. Lacustrine and adfluvial Lake Whitefish are bottom feeders and feed on snails, clams, chironomid larvae, and small fishes. Lake Whitefish are known to feed along the coastal margin of the Beaufort Sea during the summer months, although a high incidence of empty stomachs is typically reported. In contrast, anadromous Lake Whitefish sampled along the Hudson Bay coast are typically well fed. As anadromous individuals, Lake Whitefish feed on a variety of organisms including gastropods, pelecypods, amphipods, chironomids, notostracans, cladocerans, ostracods, and various insects. There is evidence from scarring that this species may be attacked by Arctic Lampreys, the marine parasitic copepod Coregonicola, and bears, as well as birds and piscivorous fishes. The lifespan is at least 30 years, perhaps over 50 years. Northern populations mature later than southern ones, as late as age eight years. Spawning typically occurs in November and December in southern populations and in late September to early October in northern populations when temperatures fall below 8ºC. The eggs hatch in the spring, usually April or May. Post-spawning adults move to overwintering areas such as the Mackenzie River Delta. Northern populations may only spawn at two- to three-year intervals. Lake Whitefish are broadcast spawners and spawn in both rivers and lakes. In lakes, spawning occurs in shallow waters and typically over hard substrates of stone and gravel, where eggs will

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lodge in crevices for overwinter incubation, although spawning over sand and silt substrates has been reported. In rivers, spawning occurs in shallow, running waters or rapids. Spawning often occurs at night with much splashing and jumping. Pairs of fish, or one female and two males, rise to the surface and shed eggs and milt. Fecundity varies by population and declines towards the northern edge of the range. For example, Lake Whitefish populations of the James-Hudson Bay area exhibit reduced fecundity northwards independent of variations in growth. Fecundity may exceed 415,000 eggs per female, or 27,400 eggs per kg of body weight, and eggs are up to 2.6 mm in diameter. Development and hatch are temperature dependent. In lakes, larvae move to deeper water from shoreline spawning areas by early summer. In rivers, larvae are carried downstream by the spring freshet to nursery estuarine areas or move into nearby lakes to develop. In the Mackenzie River, nursery areas are found in lakes in the delta that are flooded during the spring freshet, in the many channels and back eddy areas of the inner and outer estuary, or in nearshore coastal areas. In the coastal rivers of Hudson Bay, large aggregations of juveniles occur in the estuaries. Lake Whitefish overwinter in freshwater environments, with anadromous fishes returning to rivers for spawning, beginning in August. Juveniles will migrate upstream later, remaining in marine-influenced environments until late August or early September to continue growing. Data collected to date have not conclusively demonstrated natal homing on the part of migratory Lake Whitefish, although, it being a salmonid, homing is assumed to predominate.

southern Victoria Island, but there is no other distribution on any of the other islands of the Canadian Arctic Archipelago. Lake Whitefish are absent from the Ungava Peninsula north of about 60° N and from the Labrador coast north of about 58° N but are widely distributed in inland Canadian and Alaskan waters, with the exception of insular Newfoundland, and of Nova Scotia where it is replaced by the Atlantic Whitefish (Coregonus huntsmani Scott, 1987). Along the Labrador coast, Lake Whitefish appear confined to lake habitats.

importance:  Lake Whitefish are and have been commercially

fished in many of the large inland lakes in Canada. For example, the Lake Whitefish fishery in the Laurentian Great Lakes in the year 2000 was worth over US$18 million and accounted for roughly onethird of the total value of the entire Great Lakes fishery at the time. In the north, anadromous Lake Whitefish are also captured in subsistence fisheries, particularly in northern Canada along the coasts of the Beaufort Sea and in James, Hudson, and Ungava Bays. Fisheries occur at river mouths and in rivers, coincident with expected upstream autumn migrations starting in August. Throughout the Mackenzie River valley Lake Whitefish comprise a large portion of subsistence fishery catches. Fisheries for Lake Whitefish also exist in the larger inland lakes of the north (e.g., Great Slave Lake), although recent trends have shown a decline in catch associated with the declining economics of the fishery. They are marketed fresh or frozen in such categories as jumbo, large, and medium and have been canned and the eggs made into “golden” caviar.

Distribution of Coregonus clupeaformis

sources:  Backus (1957); R. Morin, Dodson, & Power (1980, 1981, 1982); Bond (1982); D.B. Stewart & Bernier (1983, 1984); Bond & Erickson (1985, 1992b); Reist, Bodaly, Fudge, Cash, & Stevens (1987); Bernatchez, Dodson, & Boivin (1989); E.S. Richardson, Reist, & Minns (2001); McDermid, Reist, & Bodaly (2005); Rennie, Sprules, & Johnson (2009).

distribution:  Lake Whitefish are ubiquitous across northern Canada. They are found in Ungava, Hudson, and James Bays, but general reports for Hudson Strait are not confirmed by specimens, and along the western Hudson Bay coast; the northern limit of distribution is at Chesterfield Inlet, on each side of the Boothia Peninsula (Perry Bay and Spence Bay), Queen Maud Gulf, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and Beaufort Sea, to a northern limit on Banks Island and a limited distribution on



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Coregonus nasus (Pallas, 1776)

Broad Whitefish, corégone tschir LES N. HARRIS

common names:  Local names are Aanaaksiiq, Aanaaliq,

Anaakliq, Anah’lih’, Anaklek, Anaklik, Ananaaklik, Anaqkik, Anaqklik, An-ark-hlirk, Angnaklin, Kausriluk, Kavasilik, and Kavisilik (Inuktitut); Aanaarliq and Anaakłiiq (Inuvialuktun); and Łuk dagaii, Łuk digaii, and Łuk zheii (Gwich’in). Other common names are Large Bottom Whitefish, Round-nosed Whitefish, Sheep-nosed Whitefish, and Whitefish.

taxonomy:  The species name comes from the Latin nasus (nose).

Although described centuries ago by Pallas (1776), the distinctiveness of Broad Whitefish (Coregonus nasus) was questioned in North America until the 1960s. This situation was exacerbated by fish from European waters that included a C. nasus form within the European whitefish complex; however, the distinctiveness of these taxa in Siberian waters was well established. Older European literature that refers to C. nasus, particularly in Scandinavian waters, represents misidentification of an ecotype of European Whitefish (C. lavaretus complex), and care is required when using literature from this area. In North America several authors considered C. nasus to be synonymous with Lake Whitefish (C. clupeaformis), and it was not until an extensive examination had been conducted that the true distinctiveness of this species was authenticated.

description:  This species is distinguished from its congeners in Arctic Canada by the profile of the upper lip being vertical or overhanging; the forehead is rounded in profile; the adult is without a distinct hump behind the head; the maxilla length is less than twice its breadth; total gill rakers number 18–25, and the length of the longest gill raker is shorter than one-fifth of the interorbital width (0.85%–1.35% of standard length). The majority of Canadian populations of C. nasus are anadromous, although both lacustrine and riverine forms exist. Few external characters are presently known to efficiently distinguish anadromous from lacustrine or riverine individuals; that is, there is great similarity and overlap of character states among these forms. There is some evidence of differences in growth rates, reproductive investment (fecundity), and various aspects of morphology; however, most of these are subtle and generally not reliable for field discrimination of forms. Where anadromous and lacustrine populations do occur in sympatry, however, they are genetically differentiated, which suggests, at least, partial reproductive isolation among these forms. Similar to all whitefishes, anadromous C. nasus are characterized by large cycloid scales (84–102 in the lateral line, 9–11 in the suprapelvic row, and 25–29 scale rows around the caudal peduncle). The scales develop tubercles during spawning. The body is elongate, and the sides are flattened or compressed

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laterally (often called “slab-sided”). The snout is blunt and projects marginally beyond the tip of the lower jaw. Pyloric caeca number approximately 148, and vertebrae 60–65. C. nasus has a deeply forked caudal fin, and dorsal fin-rays number 10–13, anal fin-rays 11–14, pectoral fin-rays 16–17, and pelvic fin-rays 11–12. In colour it is olive brown to almost black dorsally, with silver flanks, sometimes grey or brown, and a white to yellowish belly. The sides of the head are brown with light spotting. The scales are outlined in black. The fins are grey to dusky or black. The pectoral and anal fins may have blue to purple iridescence. Females develop a white first ray in the pectoral fins during the spawning season. The species attains approximately 71.0 cm in total length and 16.0 kg in weight but is usually smaller in Canada.

habitat:  Most of what is known regarding marine habitat use

by the anadromous form of this species is based on work conducted primarily in the lower Mackenzie River in Canada and the Yukon River in Alaska. Generally information on marine habitat use for this species is lacking. Typically, in larger river systems and their tributaries, newly hatched Broad Whitefish are flushed to marine habitats during the spring freshet, and these fish inhabit nearshore coastal areas for approximately 30–60 days. Young of the year then migrate into freshwater systems in July, where they overwinter and presumably rear until maturity. There is, however, evidence that juvenile fish may migrate out of these freshwater systems and inhabit marine environments prior to maturation, as evidenced by the capture of juvenile Broad Whitefish of all ages in coastal habitats. It is believed predominantly that mature Broad Whitefish feed in marine habitats and spawn and overwinter in the fresh waters of large rivers or delta areas. Some overwintering by both large juveniles and adults may take place in brackish waters or coastal habitats where there is a continuous outflow of fresh water. Recent synthetic aperture radar imagery, however, suggests that Broad Whitefish prefer large freshwater pools in larger river systems for overwintering. Little is known regarding the extent and distribution in marine waters, but otolith microchemistry results suggest that Broad Whitefish use varying degrees of marine habitat from brackish coastal habitats to full-strength sea water. Regardless, similarly to most coregonines, the marine habitat preferences of this species appear to be primarily coastal and do not involve extensive offshore movements. Field-collected fish have been shown to withstand salinities of up to 27‰, and preferences appear to be somewhat lower.

biology:  With the exception of a few populations, North American Broad Whitefish are primarily anadromous, and only recently have lacustrine and riverine forms been confirmed. In Canada very few freshwater populations are known, and the majority of biological data available for this species stems from studies and observations of anadromous populations of the Mackenzie River system. Anadromous Broad Whitefish feed on a variety of marine species, mainly preying on mysids, amphipods, and copepods. They are eaten by various other fishes and, in the Mackenzie Delta, by Brown Bears. There is evidence from scarring that this species may

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Coregonus nasus

be attacked by Arctic Lampreys and the marine parasitic copepod Coregonicola, as well as birds and piscivorous fishes. Compared to freshwater populations, anadromous Broad Whitefish have faster growth rates earlier in life, have slower growth rates later in life, grow to a larger overall size, have a higher age at maturity, and are more fecund. Most of these differences reflect the trade-offs incurred as a result of being non-migratory. Anadromous Broad Whitefish mature between five and eight years of age, which translates to a length of approximately 450 mm. In Canadian waters they can reach lengths of 600 mm and maximum ages of 35 years. Much larger fish, however, have been recorded in Europe and Eurasia (e.g., > 16 kg). The typical life cycle of the anadromous form is as follows. Spawning takes place in larger rivers and the tributaries to these systems in late October and early November, often under the ice, when the water temperature is approximately 0ºC. Upstream migrations from coastal habitats can commence as early as mid to late July. During spawning, sexes are dimorphic, with males developing larger and significantly more nuptial tubercles on their scales. Fecundity is 15,000–90,000 eggs, and the diameter of the pale yellow egg is 2.3– 2.6 mm. Broadcast spawning occurs over gravel substrates, where the eggs remain over the winter until hatching in the early spring. Adults overwinter in larger rivers or outer delta areas. Fry actively migrate to coastal habitats or passively drift to these locations during the spring freshet, where they remain until freshwater rearing systems become ice free. Rearing until sexual maturity takes place in these freshwater systems, although some juvenile Broad Whitefish may migrate back to marine habitats if the lakes chosen for rearing are not suitable. These juveniles overwinter in outer delta areas and likely migrate back into freshwater systems the following



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spring. Preceding maturity, they migrate to coastal habitats, where feeding takes place until July–August, at which time they commence upstream migrations to spawning locations. Anadromous Broad Whitefish likely spawn on a biennial (or less frequent) basis.

importance:  Broad Whitefish have been the focus of intense

subsistence fisheries and have long provided an important local resource through its range. Typically, this species is harvested by gill-net and is used for consumption by both humans and dogs. Additionally it has been the target of commercial fisheries; however, these have been intermittent within the Canadian portion of the range. The flesh of Broad Whitefish is highly sought and has been marketed as fresh, frozen, smoked, and dried. The eggs are considered a delicacy in many cultures. There is an aquaculture industry for this species in Europe; however, most attempts at stocking Broad Whitefish have been unsuccessful. Sport fishing for Broad Whitefish is negligible.

distribution:  The species is found in Queen Maud Gulf, Mel-

ville Sound, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, Beaufort Sea, and western Victoria Island. It is distributed from the Perry River, Nunavut, westwards in numerous river systems along the mainland coast (such as the Coppermine, Mackenzie, and Anderson Rivers) to the Kuskokwim River, Bering Sea drainage, in Alaska. The inland distributions in Nunavut, Northwest Territories, and Yukon Territory are restricted to northward-flowing rivers with direct drainage to Arctic marine waters. The furthest upstream regular occurrence in the Mackenzie River system appears to be Rampart Rapids, upstream of Fort Good Hope. They are distributed throughout many of the larger river systems on the North Slope of

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Alaska such as the Colville and Sagavanirktok Rivers and have been reported throughout the Yukon River and in the headwaters as far upstream as Teslin Lake (i.e., southern Yukon Territory). Anadromous Broad Whitefish are distributed throughout Arctic and Bering Sea drainages in Eurasia and North America. In Eurasia they occur as far west as the Pechora River in Russia and are distributed eastward to the Penzhina River (Sea of Okhotsk).

Coregonus sardinella Valenciennes, 1848

Least Cisco, cisco sardinelle JAMES D. REIST

common names:  Local names are Kapahilik and Kraaktak (Inuktitut); Amagiak, Anma, Anmaglak, Armagiak, Iriqpaligaurat, and Qaluhaq (Inuvialuktun) for all riverine ciscoes of the western Arctic; Treeluk (Gwich’in); Kalushak Qaqtak, Iqalusaaq, Qaaktak, and Qanktaq in northern coastal Alaska, the last three also being used in Canada. Other common names are Sardine, Big-eyed Cisco, Blue-eyed Cisco, Black-eyed Cisco, Lake Cisco, and Mackenzie Cisco. They are also referred to as herrings collectively with other ciscoes. taxonomy:  The species comes from the Latin sardina (a sardine)

Distribution of Coregonus nasus

sources:  Berg (1962); Lindsey (1962, 1981b); Ferguson, Himberg, & Svardson (1978); Reist, Bodaly, Fudge, et al. (1987); Baker & Reist (1988); de March (1989); Bernatchez, Colombani, & Dodson (1991); Bodaly, Vuorinen, Ward, Luczynski, & Reist (1991); Chang-Kue & Jessop (1992, 1997); Tallman & Reist (1997a); Treble & Reist (1997); Treble & Tallman (1997); Tallman, Abrahams, & Chudobiak (2002); Brown, Bickford, & Severin (2007); Kottelat & Freyhof (2007); McPhail (2007); Van Gerwen-Toyne, Walker-Larsen, & Tallman (2008); Barker & Derocher (2009); Brown, Duguay, et al. (2010); Harris & Taylor (2010b, 2010c); Harris, Taylor, Tallman, & Reist (2012).

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and –ella (small). It was described initially from specimens from eastern Siberian rivers, where it is clearly differentiated from the Arctic Cisco. Northern American populations identified as C. pusillus Bean, 1889, described from the Kuwuk (Kobuk) River, Alaska, represent this species, and thus this taxon is a synonym. As with other coregonines, it has also been aligned with Leucichthys under both specific epithets (see Arctic Cisco), and with Argyrosomus in keeping with the generic association of inland North American ciscoes with that old genus. As with the Arctic Cisco, this taxon is an “Old World” cisco. At least two life-history types, anadromous and lacustrine, are known, leading some authors to question the composition and validity of the taxon and also refer to it as a species complex. Additional freshwater forms found well inland or associated with particular areas exhibit variation that differentiates them from anadromous forms; however, these appear to be different from other co-occurring cisco taxa and allied to Least Cisco (as classically defined, based upon the anadromous type). Fluvial forms migrating within rivers also appear to exist in Siberia. It is not known if a similar situation occurs in North America. The following information refers to the anadromous form unless otherwise noted.

description:  This species is distinguished from other congeneric coregonines in Arctic Canada by having the profile of the upper lip sloping backwards in line with the forehead; the gape of the mouth is vertical or nearly so; the mouth is superior; and pelvic fins are dusky or black in adults. The Least Cisco is a pelagic coregonine with an elongate but slightly compressed body and a moderately sized head (about 24% of total length). Least Cisco are terete (elongate), being both dorso-ventrally and laterally more compressed than either the Arctic or the Bering Cisco. Each eye is relatively large (about 26%– 32% of head length). The moderate mouth is terminal and slightly supra-terminal, that is, the lower jaw projects beyond the upper jaw (cf. Arctic Cisco). The maxilla is at a distinct angle downwards and

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Coregonus sardinella

extends to below the anterior half of the eye (differentiating this species from the Arctic Cisco). Dorsal fin-rays number 12–14, anal fin-rays 11–13, pectoral fin-rays 14–17, and pelvic fin-rays 8–12. There are 78–98 lateral-line scales. Gill rakers are 48–53 in the anadromous form and are relatively long and slender, but are 41–47 in freshwater forms. Pyloric caeca number 74–111. The coloration overall is silvery with brown blackish to dark green hues on the back, and silver or white on the sides and belly. Unlike the Arctic Cisco, the pelvic fins in adults are heavily dusky, grading to black at the tips; in young fish (< 15.0 cm) they are pale or yellowish. The pectoral and anal fins are generally immaculate or with a very light dusting of pigment, and occasionally with dark tips. Dark spotting on the head, back, or fins is present in anadromous forms but not in non-anadromous types. Heavy, dark pigmentation on the ventral fins differentiates the freshwater form from co-occurring other ciscoes. The species attains about 47.0 cm in total length.

habitat:  Migratory populations exhibit anadromy that is typical of other Arctic coregonines; however, Least Cisco appear to not venture far from natal rivers. They use coastal marine areas for summer feeding. Typically during summer they are found in freshened areas (usually with salinities of less than 15 psu) associated with river deltas and nearby coastal areas. Although they are migratory within systems, it appears that long-distance movements such as those described for Arctic Cisco do not occur; thus, local systems likely have somewhat distinct populations. Upstream migrations of adults occur in late summer and early fall in rivers, and spawning occurs over sand or gravel areas near river margins and perhaps also lakes. Nursery areas for young fish and feeding areas for adults include river deltas, estuaries, and freshened nearshore locations.



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Upon attaining maturity, adults likely remain associated with larger water bodies within the systems. Extensive use of the lakes of the Mackenzie Delta and those of the Tuktoyaktuk Peninsula occurs. Overwintering occurs in freshened estuarine areas of rivers or in the river channels themselves, similarly to other coregonines.

biology:  Its food in fresh water includes aquatic insects, terrestrial insects taken at the water surface, worms, molluscs, crustaceans, and fishes. They are eaten by Lake Trout, Inconnu, and Burbot. The age of first maturity appears to be around year four, and individuals as old as 16 years have been captured. Individuals are iteroparous, and adults may not spawn in consecutive years. As noted, given their propensity for only short-distance migrations, Least Cisco from different systems likely form distinct populations, completing their life history in association with their natal river and nearby marine areas. The upstream runs of mature individuals to spawning areas occur late in the open-water season, fall spawning is typical (i.e., September–October), and spent adults likely move downstream to overwintering areas in lower river reaches and delta lake systems. Eggs are laid over gravel or sand bottoms at 1.0ºC–1.5ºC. A female can have up to 23,600 eggs. Young emerge in spring and are likely flushed downstream on the river freshets. Unlike Arctic Cisco, this species makes no concerted coastal westward migrations from any single river; rather, young fish appear to concentrate and remain in the deltas. Those from the Mackenzie River move generally eastwards along the Tuktoyaktuk Peninsula, occupying river mouths and embayments in this area. Young Least Cisco make some use of the various river systems present in this

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area, but this is not as extensive as that by other coregonines (e.g., Broad Whitefish). Migratory anadromous forms and non-migratory lacustrine forms likely co-occur in many coastal systems (the latter can usually be differentiated in that they lack the black spots found on anadromous fish), thus complicating the understanding of life history and biology.

importance:  Least Cisco, similar to other harvested coregonines, are culturally important to western Arctic Indigenous peoples in both Canada and Alaska, where they support substantive subsistence fisheries. This is especially so along the coasts, but harvests also occur inland on migrating adults of this species and Arctic Cisco. In Canada these fishes are known collectively, as “herring” by local peoples. Harvests traditionally have been quite large, primarily for use as dog food. Although tasty with firm white flesh, Least Cisco are less popular as human food. Least Cisco is generally not commercially fished, and it likely represents a by-catch in such fisheries in North America. Extensive commercial fisheries occur in Siberian waters. Similar to other anadromous Arctic fishes, Least Cisco are a keystone component linking freshwater, estuarine, and marine ecosystems through material and energy transfers. These processes (and thus the fish themselves) may be particularly important in Arctic ecosystems with low productivities. Given the apparent large numbers of Least Cisco present (one estimate for the Colville River indicates that over one million fish between 11–30 cm in length were present), this ecological role may be extremely important. distribution:  It is found in the Rasmussen Basin and Chantrey Inlet, northern and western King William Island, Queen Maud Gulf, Melville Sound, Coronation Gulf, Dolphin and Union Strait, Victoria and Banks Islands, and the Beaufort Sea. This species is associated with virtually all freshwater and nearshore coastal habitats present in Arctic areas and is most common near to the mouths of large river systems. Geographically it extends from the White Sea (northern Russia). Records east of the Boothia Peninsula (not mapped) require confirmation and are likely to be Cisco, C. artedi. The species occurs on the southern margins of Banks and Victoria Islands and in river systems (and presumably coastal embayments) on the northern margin of the former. No occurrences are reported from more northerly islands. It extends to inland areas, likely mostly as different variants (e.g., to lakes near Great Slave Lake in Canada); however, the taxonomic relationships of these are unclear. Unlike Arctic Cisco, it occurs in Alaskan and Chukotkan drainages to the Bering Sea.

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Distribution of Coregonus sardinella

sources:  Corbin (1986); Reist & Bond (1988); Turgeon & Bernatchez (2003); C. George et al. (2009); Moulton et al. (2010); Stephenson (2010).

Oncorhynchus gorbuscha (Walbaum, 1792)

Pink Salmon, saumon rose KAREN DUNMALL

common names:  Local names are Amaatuuq (Inupiatun, Alaska); and Pukkellaks (Danish/Greenlandic). Other common names are Autumn Salmon, Dog Salmon, Humpback Salmon, and Humpy.

taxonomy:  The genus comes from the Greek onkos (hook) and rhynchos (snout). The species name comes from the Russian name for “humpback” in Alaska. The genus Oncorhynchus in the subfamily Salmoninae contains the Pacific salmons and Pacific trouts. While some believe that the Family Salmonidae has a marine origin, the more widely accepted theory is that the Pacific salmon species diverged from the trout genus Salmo and have a freshwater origin.

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Oncorhynchus gorbuscha, male silver

description:  This species is the smallest of the congeneric salmon species and is distinguished by the presence of distinct oblong black spots on the back and the caudal fin, with the largest being as large as the eye. Dorsal fin-rays number 10–16, anal fin-rays 12–19, pectoral fin-rays 13–19, and pelvic fin-rays 9–11. Lateral-line scales number 147–205, and in the first row above and paralleling the lateral-line scales 169–229. Gill rakers are of moderate length and number 24–35, pyloric caeca number 95–224, and vertebrae 63–72. Spawning males also have extended, hooked snouts, gaping mouth, enlarged teeth, a large hump before the dorsal fin, and embedded scales. The adipose fin is always larger in males than in females. At sea the colour of the fusiform, streamlined body is steel blue to blue green on the back, silver on the sides, and white on the ventral surface. Large black spots are present on the back and extreme upper sides and on the adipose and caudal fins. Breeding males become darker on the head and back, and the sides become pale red with brown to olivegreen blotches. Females change colour less dramatically than do males. Pink Salmon parr have blue-green backs and silvery sides, with no black spots on the fins and no parr marks. Adult Pink Salmon at sea are approximately 61.0 cm in length. This species attains 76.0 cm in total length and 6.4 kg in weight. habitat:  Pink Salmon exhibit an anadromous life-history type.

Pink Salmon enter rivers and travel upstream to spawn, which usually takes place between August and September at 10ºC–16ºC, with later runs at lower temperatures. Adult Pink Salmon die after spawning (i.e., are semelparous). The eggs hatch between December and February, and the alevins remain in the gravel until April or May. Emergent fry promptly migrate downstream to estuarine waters, where they adapt to higher salinity and feed. Young Pink Salmon may stay in inshore waters for several months before moving to the ocean. Pink Salmon spend 18 months at sea before returning to spawn, and their ocean migration is influenced by temperature and ocean currents.



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biology:  Fry in the estuary are active during the day and form

schools, feeding primarily on invertebrates, such as copepods, euphausiids, amphipods, ostracods, larvae of decapods, cirripedes, tunicates, and dipterous insects. Pink Salmon may spend several weeks feeding in the estuary before heading to sea. In the marine environment the most abundant prey items include euphausiids, amphipods, fishes, squid, copepods, and pteropods. Adults do not eat while in fresh water and die a week or two after spawning. They are cannibals and are eaten by various other fishes and by birds and mammals. Lampreys are frequently parasitic. Pink Salmon have a fixed age at maturity of two years, although individuals at three years and at one year have been reported. Due to this two-year life cycle, distinct odd-numbered and even-numbered runs have developed. These runs exhibit little gene flow between them and may vary in the timing of return, the size of fish, and the local adaptations to natal streams. In western Alaska, Pink Salmon are more abundant in even-numbered years than in odd-numbered years, although a similar pattern is not as obvious in the Canadian Arctic. While Pink Salmon home to their natal streams to spawn, vagrants are relatively common. Pink Salmon usually move only a short distance (e.g., ~65 km) upstream before spawning, although some move as much as about 480 km upstream whereas others spawn in the lower tidal area. The female builds a redd in the gravel usually in a riffle area where there is little silt or sand and the water depth is less than 1.0 m and frequently between 0.2 m and 0.25 m. Redds, excavated by lashings of the tail, can be up to 1 m long and 0.5 m deep. Females may build and spawn in more than one redd, and males may spawn with more than one female. A spawning pair, sometimes accompanied by 10 or more other males, positions itself in the redd, gapes, and releases the eggs and sperm, usually at night. The spawning pair consists of the female and the longest, most hump-backed male, with the other males arranged by size behind them. The other males fight and jockey for position and have deep

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Oncorhynchus gorbuscha, male spawner

Oncorhynchus gorbuscha, female spawner

cuts from the enlarged teeth. There may be an outlier male, small and coloured like a female, which maintains a position to one side of the nest. The other males and the outlier all release sperm when the main pair spawns. After the orange-red eggs are fertilized and deposited, the female covers the redd and its eggs by dislodging upstream gravel. The egg number varies with the size of the female, the area, and the year but usually ranges from 1,500 to 1,900 per female, and the egg size is large, approximately 6.0 mm in diameter. Incubation usually takes four to six months, with hatching occurring in late December to late February, at least in more southerly distributed populations. The fry then emerge from the gravel and move at night, hiding in the gravel during the day. If the migration distance to estuary areas is longer, the fry also move during the day and form schools.

importance:  The lack of an Inuvialuit name for Pink Salmon

suggests that it is not frequently captured and there is no history of established populations in the western Canadian Arctic. Established breeding populations exist between Point Hope and Point Barrow, Alaska, and early records show that Pink Salmon were captured and sold in 1875 at the mouth of the Colville River, Alaska. No established

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populations are known east of Point Barrow, although local knowledge suggests some might occur. Pink Salmon have been incidentally captured in subsistence fisheries in the Mackenzie River Delta region of the western Canadian Arctic. Occurrences of vagrants appear to be increasing, which may increase competition between Pink Salmon and established populations of other salmonids in the Arctic. All species of Pacific salmon are exploited to significant degrees wherever they occur in reasonable abundances. This group has formed the mainstay of subsistence fisheries by Indigenous peoples on both sides of the Pacific Ocean and similarly has a long history of exploitation in commercial fisheries. Significant recreational fisheries also exist throughout the ranges of the various species. This species has been less regarded in Indigenous and commercial fisheries than other salmons on the Pacific coast of Canada. However, this has changed, and catches have exceeded ten million fish annually in Canada. Catches are made with purse seines and gill-nets and by trolling. The 1988 catch was 31,355 t and was worth $42 million. The flesh is pink (hence the common name) and tasty though reputedly not as good as Sockeye, Coho, or Chinook. Most of the catch is canned. Anglers catch this species by trolling bait in the ocean or large lakes or by flies and artificial lures on the spawning run. After Coho and Chinook it is

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the third most important Pacific salmon for anglers. Like most migratory fishes, it is very susceptible to environmental changes.

distribution:  Pink Salmon range from the Lena River, Russia, to the Mackenzie River, Canada, in the Arctic Ocean and from Peter the Great Bay to the Sacramento River, California, in the Pacific Ocean. In the western Arctic, spawning populations of Pink Salmon exist between Point Hope and Point Barrow, Alaska, and perhaps in the Colville River. Stray Pink Salmon have been reported infrequently in the western Canadian Arctic, most often in or near the Mackenzie River, since 1890. The proximity of the Colville River to the Mackenzie River may help to explain the increase in vagrants noticed in the western Canadian Arctic in some years. In almost all cases of incidental catch, only single specimens have been captured, and all captures have been made in August or early September. Only one Pink Salmon has been captured further east of the Mackenzie River, at Sachs Harbour on Banks Island. The Peel River represents the furthest inland that Pink Salmon have been captured in the Canadian Arctic; half of all captures have been made in coastal areas, and the rest were caught in the lower reaches of rivers. This situation, however, appears to be changing in recent years as indicated by increased captures. Spawning has not been recorded in the western Canadian Arctic, so it is unlikely that colonization has occurred. A small freshwater resident population exists in Lake Superior, introduced in 1956 from Skeena River, British Columbia. In 1958, Pink Salmon were introduced in Newfoundland, and by 1967 strays from this population had spread to Nova Scotia, Labrador, and the Gulf of St Lawrence to the Great Watchichou River, Québec, and possibly to southwest Greenland. Similarly, introductions were previously made in Hudson Bay, but these appear not to have survived.

sources:  Takagi, Aro, Hartt, & Dell (1979); Craig & Haldorson (1986); Heard, in Groot & Margolis (1991); Babaluk, Reist, Johnson, & Johnson (2000); Quinn (2005); Stephenson (2005, 2006); C. George et al. (2009).

Oncorhynchus keta (Walbaum, 1792)

Chum Salmon, saumon kéta KAREN DUNMALL

common names: Local names are Iqalugruaq and Iqaluruaq (Inupiatun, Alaska); Paiirluq (Inuvialuktun); Shii (Gwich’in); and Geo Sahba (North Slavey). Other common names are Autumn Salmon, Calico Salmon, Dog Salmon, Keta Salmon, and Silverbrite Salmon. “Chum” may be from the Chinook word for “spots or writing.” Chum Salmon are referred to by many different names throughout their range. For instance, there are at least nine names for Chum Salmon used in Russia, and more than ten names used in Japan. taxonomy:  The species name comes from the language of Nanai,

from northeast Russia and southeast China, and means “fish.”

description:  This species is distinguished from congeneric species by the presence of short, stout, widely spaced gill rakers that number 16–28 on the first gill arch, and the absence of distinct black spots on the tail and the back. Dorsal fin-rays number 10–14, anal fin-rays 13–17, pectoral finrays 14–16, pelvic fin-rays 10–11, lateral-line scales 124–153, pyloric caeca 140–249, and vertebrae 59–71. Spawning adults have extended hooked snouts, a gaping mouth, bared teeth, a slight hump before the dorsal fin, and embedded scales. Females change less than males do, and the colour change is more distinct on males than on females. At sea the fusiform, streamlined body is steel blue on the back and upper sides, silver on the sides, and silvery to white on the ventral surface. There may be fine black specks on the upper sides and the back of the fish, but there are no distinct black spots on the back, sides, or fins. The tip of the anal fin is white, although males have tinges of black on the tips of the caudal, anal, and pectoral fins. When Chum Salmon return to fresh water to spawn, they become dark olive to almost black on top, and grey red on the sides with dirty-green vertical bars. Parr are marked with 6–14 narrow, short parr marks, most of which do not cross the lateral line. Mackenzie River Chum Salmon are approximately 66.7 cm in length and weigh 3.82 kg. The species attains 109.0 cm in total length and 20.8 kg in weight.

Distribution of Oncorhynchus gorbuscha



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Oncorhynchus keta, male silver

habitat:  Chum Salmon enter rivers and travel upstream to

spawn. They are not strong jumpers and are halted by the first barrier. However, the Yukon River has a run that travels 3,200 km upstream. A migratory speed of 115 km/day has been recorded. Eggs hatch in the spring, and alevins spend several weeks in the substrate before emerging. Once emerged, fry promptly migrate downstream to estuarine waters, arriving in June or July of the year following spawning, where they adapt to higher salinity and may spend several weeks feeding. In the Mackenzie River system, juveniles may use lakes as nursery and feeding areas during their migration. Juvenile Chum Salmon correlate their ocean entry with the warming of nearshore waters and the shift in prey availability from inshore to offshore. Juveniles also move offshore when they are large enough to avoid predators, and feed on larger neritic prey. It is unclear where Chum Salmon (or indeed any of the Arctic populations) spend their winters while at sea to avoid freezing temperatures. They may follow currents and a plume of warmer Mackenzie River water west to the Bering Sea, they may overwinter in warmer waters of Atlantic origin at depths greater than 200 m and then return to the surface to feed and grow in the Beaufort Sea, or they may overwinter in brackish or fresh water that is near zero in temperature and spend their summers in the Beaufort Sea.

biology:  It is not clear to what extent Chum Salmon fry feed as

they move downstream, although migrations are extensive in the Mackenzie River system and young likely feed along the way. Once in estuaries, the juveniles feed primarily on invertebrates, but the preferred food varies by location. Chum Salmon are selective for larger prey in both epibenthic and neritic feeding. Copepods, tunicates, and euphausiids are the most abundant prey items of Chum Salmon in the marine environment, and other prey items include fishes, arrow worms, pteropods, and squid. Adults do not eat while in fresh water. Chum are eaten by various fishes, birds, and mammals. They are parasitized by Lampreys.

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Four-year-old Chum Salmon predominate in spawning populations in the Mackenzie River, with some three- and five-year-old fish being captured. The lifespan in British Columbia is about seven years, perhaps nine years, the fish spending as much as seven years at sea before returning to spawn. The time at which mature fish migrate to spawning waters differs with location. Although Chum Salmon often return to spawn in two modes each year in many locations, in Arctic Canada spawning Chum Salmon return to the Mackenzie River in the fall, arriving at the mouth in late summer and travelling upstream until after the river ices over in November. Chum Salmon frequently have already undergone physiological changes and are in a reproductive state prior to entering the fresh water and can therefore spawn almost immediately or, as in the Mackenzie River spawning populations, travel approximately 2,000 km before spawning. The female builds a nest that is up to 3.2 m long and 2.1 m wide in the gravel, usually in an area of upwelling, where there is little silt or sand and the water depth is usually less than 1 m. In some cases no redd is excavated, and the eggs are shed over and between boulders. Females can build more than one nest and may be attended by more than one male. The orange-coloured eggs usually number from 2,400 to 3,100, and up to 7,779, per female, and the egg size ranges from 5 mm to 6 mm in diameter in the ovary. The females guard the nests. Chum Salmon are semelparous, like all Pacific salmon species, which means that adults die after spawning. The timing of hatching depends on the water temperature. Once hatched, the alevins wait in the gravel for conditions that favour their seaward migration. The fry then emerge from the gravel and move at night, hiding in the gravel during the day. If the migration distance is longer, the fry move during the day as well and form schools.

importance:  The existence of an Inuvialuktun name for Chum

Salmon provides historical evidence of the long-term presence of a spawning population in the Mackenzie River basin. Chum Salmon

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Oncorhynchus keta, male spawner

Oncorhynchus keta, female spawner

have been captured in subsistence and commercial fisheries in the Canadian western Arctic since 1914; however, they are uncommon outside the Mackenzie River. Capture frequency has increased in the past decade, perhaps as a result of climate change. More frequent straying may increase the likelihood of competition with indigenous salmonids. This fish is caught commercially in British Columbia during September and October with the use of gill-nets and purse seines in sheltered marine waters. A catch of 29,998 t, worth $72 million, was made in 1988. Some are sold fresh, but most are canned. The flesh is white and is favoured for smoking. It ranks third in commercial fisheries after Sockeye Salmon (Oncorhynchus nerka) and Pink Salmon. They are not fished extensively by anglers because they may spend only two to three weeks in fresh water. Its use as sled-dog food is indicated by its common name of Dog Salmon.

distribution:  One spawning population of Chum Salmon is known on the Mackenzie River in the Slave River near Fort Smith,



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Northwest Territories. Another spawning population may also exist in the upper Liard River, northern British Columbia (drainage to the Mackenzie River). Chum Salmon are also captured in the Peel River, a tributary of the lower Mackenzie River. Vagrants are caught irregularly in subsistence fisheries at Aklavik, Northwest Territories, along the length of the Mackenzie River, and in Great Bear and Great Slave Lakes. The number of Chum Salmon estimated in the western Canadian Arctic ranges from 1,000 to 5,000. Chum Salmon have been infrequently reported east of the Mackenzie River, although a single specimen was taken near Kugluktuk (Nunavut), and they have been reported at Paulatuk. Records of individuals from Hudson Bay indicate introduced fingerlings that failed to survive. Chum Salmon have the largest natural distribution of all Pacific salmon species, ranging from the Lena River (central Siberia) to the Mackenzie River (Canada), in the Arctic Ocean, and from Korea to Monterey (California) in the Pacific Ocean.

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Distribution of Oncorhynchus keta

sources:  Bakkala (1970); McLeod & O’Neil (1983); Babaluk, Reist, Johnson, & Johnson (2000); E.S. Richardson et al. (2001); Quinn (2005); Stephenson (2005, 2006); C. George et al. (2009); Irvine et al. (2009); Dunmall, Reist, Carmack, Babaluk, HeideJorgensen, & Docker (2013).

Oncorhynchus kisutch (Walbaum, 1792)

Coho Salmon, saumon coho KAREN DUNMALL

common names: Other common names are Blueback, Hooknose, Hoopid Salmon, Sea Trout, Silver Salmon, Silversides, and White Salmon. taxonomy:  The species name comes from the Russian kizhuch, which is the local name in Alaska and Kamchatka. Extensive interest, with a long history of taxonomic resolution, and in most cases clear distinguishing characteristics, combined with lower degrees of phenotypic variation (ecological, life history) than those of many other northern fishes, have resulted in resolution of most of the taxonomic issues associated with Pacific salmons.

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description:  This species is distinguished from congenerics by the presence of distinct small black spots (pupil-sized or smaller) on the back and the top lobe only of the caudal fin, and the absence of black pigment along the lower gums. Dorsal fin-rays number 9–13, anal fin-rays 12–17, pectoral finrays 12–16, pelvic fin-rays 9–11, and lateral-line scales 112–148. The gill rakers are rough, widely spaced, and total 18–25, pyloric caeca number 45–114, and vertebrae 61–69. The snout on males becomes thicker and hooked, but less so than in other species of Pacific salmon, there is a slight hump, and the teeth are enlarged. At sea the fusiform, streamlined body is steel blue to slightly green in colour on the back, silver on the sides, and white on the ventral surface. Small black spots are on the back, the upper sides, and the upper lobe of the caudal fin. The adipose fin is dusky. Breeding males become darker blue green and have dull sides with a brilliant red stripe. During reproduction, females change less dramatically than do males. Coho Salmon parr are blue green on the back and have 8–12 narrow, vertical parr marks along the sides. The lateral line extends through the middle of the parr marks, and the pale area between the parr marks is greater than the width of the parr mark. At sea, adult Coho Salmon range in length from 457 mm to 610 mm and in weight from 3.6 kg to 5.4 kg. It attains 108.0 cm in total length and 17.7 kg in weight. habitat:  Coho Salmon exhibit an anadromous life-history type,

although some fish, known as residuals, remain in fresh water and do not spawn. Maturing adults enter rivers in late summer and autumn and travel upstream to spawn. The alevins remain in the gravel for two to three weeks. Emergent fry initially form small schools in backwater areas, shallow pools, and eddies but then become territorial and move into pools and areas having little current. Juvenile Coho are found in areas of cover, including back eddies of rivers, and near vegetation or river banks and move to deeper, slower water that has an abundance of cover in which to overwinter. The length of time that juveniles spend in the fresh water before migrating to the ocean depends on the geographic location of the population. In Alaska juvenile Coho remain in fresh water for one to two years, and rarely three or four years, and in the Yukon they remain for two years. Coho usually remain inshore for a few months and then enter the open ocean, spending a year in the sea before returning to their natal stream to spawn. About 15% of fish are strays from their natal stream.

biology:  Juvenile Coho Salmon in fresh water in Alaska feed

mainly on insects including dipteran larvae, Trichoptera, Plecoptera, and Coleoptera but also eat large numbers of juvenile Sockeye Salmon, Chum Salmon, and Pink Salmon. Juveniles in the nearshore environment feed on the fry of Pink Salmon and Chum Salmon, other fishes, squid, and crustaceans. The diet of adult Coho Salmon in the marine environment is more pelagic and varied than that of other Pacific salmon species and consists of fish (80%), such as Pacific Herrings and Sand Lances, and invertebrates (20%). Adults do not eat while in fresh water. They are eaten by other salmon, Sculpins, birds, and such mammals as seals and Killer Whales. Lampreys regularly attack Coho Salmon.

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Oncorhynchus kisutch, adult silver

Oncorhynchus kisutch, male spawner

Coho Salmon characteristically migrate from the sea to rivers to spawn, preferring to travel during the day, and the fry remain in the stream for a year or more before migrating to the ocean. Northern populations typically have a larger proportion of Coho Salmon that spend two years in fresh water before migrating to the ocean, and the proportion of four- and five-year-old individuals returning to spawn increases northward. Coho Salmon typically school at the mouth of the river and move upstream when rainfall increases the water flow. The timing of spawning varies with geographic location, although most spawn later than do other Pacific salmon species. Coho Salmon spawn in the tributaries of larger rivers and select riffle areas below pools. The female builds a redd in the gravel where there is little silt or sand and the water depth is at least 0.15 m. More than one redd may be built and be attended by more than one male. The eggs are deposited and fertilized, and the female covers the redd. Sneaker males may rush in and try to fertilize the orange-red



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coloured eggs. Adult Coho Salmon die after spawning. The eggs hatch in six to eight weeks, depending on the water temperature. The number of eggs, up to 5,700, varies with the size of the female, geographic location, and year, but in British Columbia egg numbers range from 2,100 to 2,789, and egg size is approximately 4.5–6.0 mm in diameter. Emergent fry initially school but then become territorial, feed voraciously, and grow quickly. Overwintering juveniles move into deep pools that have abundant cover. When they are ready, juvenile Coho Salmon migrate to the ocean in spring in small schools, mainly at night or on freshets.

importance:  Extensive fisheries occur for this species throughout its range except in most populations present in Arctic locations that typically have low abundances. Coho Salmon are the rarest of all Pacific salmon species present in the Canadian Arctic.

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The 1988 catch in Pacific Canada weighed 6,122 t and was worth over $32 million. The fish are caught by gill-nets, purse seines, and trollers from July to September and sold fresh, frozen, cured, or smoked. They are also canned as medium red salmon. The flesh is pink to red but turns white during spawning. Anglers take this fish from June to October in its first or last year of ocean life. Coho Salmon fight strongly and leap dramatically.

distribution:  Coho Salmon range from the Anadyr River (Russia) to Chongjin (North Korea) in western Pacific drainages and from the Kukpuk River near Point Hope (Alaska) to Monterey Bay (California) in eastern Pacific drainages. Coho Salmon have been extensively introduced, for example to the Laurentian Great Lakes, but those populations are usually maintained by fish culture rather than by natural reproduction. This species also occurs in Pacific drainages in southwestern Yukon, and in Bering Sea drainages (i.e., Yukon River) upstream to the southern portion of the Yukon Territory. The northernmost spawning population of Coho Salmon exists near Point Hope, Alaska, and strays have been captured further east to Prudhoe Bay, Alaska. Coho Salmon are the rarest of all Pacific salmon species in the Canadian Arctic. Only two captures have been reported; one was in the subsistence fishery on Great Bear Lake in 1987, and the other was made through the ice on the Mackenzie River Delta near Inuvik in 1998. Spawning has not been recorded in the western Canadian Arctic, so it is unlikely that colonization has occurred.

Distribution of Oncorhynchus kisutch

sources:  Craig & Haldorson (1986); Babaluk, Reist, Johnson, &

Johnson (2000); Stephenson (2005, 2006).

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Oncorhynchus nerka (Walbaum, 1792)

Sockeye Salmon, saumon rouge KAREN DUNMALL

common names: Other common names are Kokanee or Kokani in particular, Kennedy’s Salmon, Kickinee, Koke, Little Redfish, Silver Trout, and Yank (for the freshwater form); and Blueback Salmon and Red Salmon (for the sea-run form). taxonomy:  The species name comes from the Russian name for the anadromous form. “Sockeye” is an anglicization of its name in Halkomelem, the language of Indigenous people along the lower Fraser River. “Kokanee” comes from the Okanagan word for this type of fish. This species exhibits two forms, the permanent freshwater form, called the Kokanee, and the anadromous form, called Sockeye Salmon. Although Sockeye Salmon were originally described as Salmo nerka Walbaum, 1792, and Kokanee were originally described as a separate species, Salmo kennerlyi (Suckley, 1862), they are now considered subspecies O. n. nerka and O. n. kennerlyi. The two forms are very similar except the Kokanee is smaller than the Sockeye that goes to the ocean, and when the two forms occur together, they exhibit differences in gill-raker number, male spawning characteristics, and coloration. description:  This species is distinguished from its congeneric species by the presence of long, slender, serrated, closely spaced gill rakers that number 28–44 on the first gill arch (31–44 in Kokanee), and the absence of distinct black spots on the tail and back. Dorsal fin-rays number 11–16, anal fin-rays 13–18, pectoral finrays 11–21, pelvic fin-rays 9–11, lateral-line scales 120–150 (121–140 in Kokanee), pyloric caeca 45–115 (50–87 in Kokanee), and vertebrae 56–67 (62–67 in Kokanee). Breeding males have a prolonged, hooked snout, a gaping mouth, and a small hump before the dorsal fin, but these are not as marked as in some other Pacific salmons. The head and mouth of spawning Sockeye are less deformed than in other forms of Pacific salmon. At sea the fusiform, streamlined body is steel blue to blue green in colour on the back with no distinct spots, silver on the sides, and white on the ventral surface. The fins are clear to dusky. Breeding males become bright red to red grey on the back and sides, and dirty red to grey on the ventral surface. The head to lower jaw becomes bright green to olive, with black on the snout and white on the lower jaw. The dorsal, adipose, and anal fins become red, and the pectoral, pelvic, and caudal fins turn green to black. Females change colour similarly, but the body is darker grey red in colour. In some freshwater resident populations, spawning salmon are dull green to yellow without prominent red colouring. Sockeye Salmon parr have blue-green backs, silvery sides, and 8–14 short, dark, oval parr marks that do not extend below the lateral line. Light areas are greater than dark areas along the lateral line.

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Oncorhynchus nerka, male silver

Oncorhynchus nerka, male spawner

Juvenile Sockeye become silvery when preparing to start their seaward migration in spring. The species attains 84.0 cm in total length and 7.0 kg in weight as Sockeye, and to 53.3 cm in total length as Kokanee.

habitat:  Anadromous Sockeye Salmon enter rivers and travel

upstream to spawn, which usually takes place in the fall and varies by area. Alternate years have dominant and subdominant runs, the latter being less numerous. The eggs hatch during the winter or early spring, and the alevins remain in the gravel until April–June. Emergent fry remain in the nursery lake, feeding and exhibiting schooling tendencies. Sockeye Salmon can spend one to two years, and rarely three to four years, in the lake before migrating to the sea and can spend two to three years, and rarely one or four years, at sea before returning to spawn. Freshwater resident or residual populations have similar habitat requirements to those of anadromous populations in the freshwater environment.



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biology:  Juvenile salmon stay in the lake environment, feeding

primarily on plankton crustaceans and chironomid pupae. Sockeye Salmon in the ocean feed on zooplankton and squid, and infrequently small fishes. Young Kokanee and Sockeye are eaten by a wide variety of other fishes, including other salmonids and Sculpins, and some adults are taken by seals and bears. The biology of freshwater resident populations is similar to that of anadromous fish except, instead of migrating to sea, juveniles remain in the lake to maturity. Kokanee mature mainly in their fourth year, and occasionally in their third or fifth year. Some fish may be as old as eight years. The life history of Kokanee in the Peace River system is assumed to be similar to that of Kokanee in other areas. Juvenile Sockeye exhibit active migration, swimming faster than the current, and remain in schools to avoid predators. Sockeye Salmon are primarily anadromous, although landlocked Kokanee populations exist. Sockeye characteristically make use of a lacustrine habitat for juvenile rearing, and this adaptation to

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lake environments requires this species to be more precise regarding timing and location when homing to spawning areas in order to take advantage of optimal rearing conditions. Sockeye Salmon enter coastal streams from May to October, the early entrants having further to travel to the spawning site, being earlier spawners, or spending up to three to four months in fresh water before spawning. Sockeye usually spawn in streams that have lakes in their drainage. Spawning occurs from July to December at 3ºC–7ºC. The female excavates a redd in the gravel where there is little silt or sand in inlet streams to the lake where they rear, or in upwelling areas in gravel beds along the shores of the lake. The redd is about 10.2 cm deep and longer than the fish. The spawning process is similar to that of other Pacific salmons. Orange-red eggs are deposited and fertilized, and the female covers the redd. Females can build more than one nest and may be attended by more than one male. Adult Sockeye Salmon die after spawning. The egg number varies with the size of the female but usually ranges from 368 to 1,764 eggs per female (up to 4,300), and the egg size is 4.5–5.0 mm in diameter. The timing of hatching is dependent on the water temperature but can occur up to five months after spawning or after several weeks. Alevins remain in the substrate for two to three weeks, emerging in April–June, when they typically move to a nearby lake. Smolts and fry tend to move at night to avoid predators.

Bering Sea, and eastern Sea of Okhotsk north of 40° N. Sockeye Salmon are extremely rare in the Arctic; however, they have the greatest geographic distribution of the Pacific salmon species in the Canadian western Arctic. Spawning populations of Kokanee exist in drainages to the Canadian Arctic at Arctic and Thutade Lakes, headwater lakes in the Peace River system. Stray Sockeye Salmon have been recorded east of Point Barrow in Alaska to the Mackenzie River. Vagrant Sockeye Salmon have also been reported at Bathurst Inlet (Nunavut), Uluhaktok (Holman), Victoria Island, Prince Albert Sound, Victoria Island, and Sachs Harbour, Banks Island. This species also occurs in Pacific drainages in southwestern Yukon. The southernmost capture site for Sockeye Salmon in the Canadian Arctic is the Slave River, near Fort Smith, and it may have in fact been a Kokanee. One Kokanee has been captured in the commercial fishery in Great Slave Lake and was likely from the native populations in the Peace River system. It has also been widely introduced, for example, to the Laurentian Great Lakes.

importance:  Significant fisheries for this species are present

throughout much of its range, decreasing in more northerly locations due to lower population sizes. Sockeye Salmon are infrequently found in the Canadian Arctic and are only incidentally captured in subsistence fisheries. Kokanee present in the Peace River system of northern British Columbia may be the source population for strays to Slave River, a tributary of Great Slave Lake, via the W.A.C. Bennett and Peace Canyon Dams. One of the factors limiting distribution is low water temperature, so climate change may influence the marine distribution of Sockeye Salmon by increasing water temperatures, which may result in more frequent straying and straying longer distances from natal streams. Increased straying or the establishment of a spawning population of Sockeye or Kokanee as a result of climate change would have an impact on native populations of fishes. This is the most important of the Pacific salmons, caught by gillnets and purse seines and sometimes by trolling. In 1990, 22 million Sockeye returned to the Fraser River system of British Columbia, for example. About eight out of ten of the fish returning to the Fraser are caught by commercial, Indigenous, and recreational fisheries. The 1988 Canadian catch weighed 11,808 t and was worth almost $82 million. The run is also a tourist attraction. Dams, chemical and thermal pollution, destruction of spawning sites, and probably overfishing have caused declines in runs.

Distribution of Oncorhynchus nerka

sources:  Foerster (1968); Hunter (1974); Craig & Haldorson (1986); D.W. Welch, Ishida, & Nagasawa (1998); Babaluk, Reist, Johnson, & Johnson (2000); Babaluk, Reist, & Low (2000); Stephenson (2005, 2006).

distribution:  Spawning populations of Sockeye Salmon extend from the Anadyr River to the southern tip of the Kamchatka Peninsula (Russia), and along the north shore of the Sea of Okhotsk in the western Pacific area, and range from the Sacramento River (California) to the Pilgrim River (western Alaska) in the eastern Pacific. In the ocean, Sockeye Salmon are found in the North Pacific Ocean,

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habitat:  Chinook Salmon exhibit an anadromous life-history

Oncorhynchus tshawytscha

type. Chinook Salmon enter rivers and travel upstream to spawn, the timing of which varies throughout their geographic range and can also vary between populations within a river. The travel distance to spawning areas also varies depending on the river and can be as long as 4,827 river km. Some fish go 1,600 km or more out to sea. Freshwater temperatures of 12°C–14°C are preferred. Emergent fry in some rivers promptly migrate to sea, but, more commonly in northern populations, the fry spend one or two years in the freshwater environment before migrating. The smolts stay in inshore waters for some time before moving to the ocean. Chinook Salmon usually spend two or three years, and rarely up to five years, at sea before returning to spawn. Northern populations usually remain in the ocean for longer periods.

(Walbaum, 1792)

Chinook Salmon, saumon chinook KAREN DUNMALL

biology:  Chinook Salmon exhibit large variations in their life

Oncorhynchus tshawytscha, juvenile

common names:  Local names are Sahti Eda (Dogrib) and

Tarjaxfaq (Inuktitut). Other common names are Black-jaw, Blackmouth, Chub Salmon, Hookbill, King Salmon, Quinnat, Spring Salmon, Tyee, Winter Salmon, and saumon quinnat.

taxonomy:  The species name comes from the Russian, which in turn is a derivation of a common name used among Indigenous peoples in Alaska and Kamchatka. The species may be spelled incorrectly as tschawytscha. description:  This species is distinguished from its congeners by the presence of distinct small (pupil-sized or smaller) black spots on the back and on both lobes of the caudal fin, and of black pigment along the base of the teeth in the lower jaw. Meristic counts vary greatly over the geographic range of distribution. Dorsal fin-rays number 10–14, anal fin-rays 13–19, pectoral fin-rays 14–17, pelvic fin-rays 10–11, and lateral-line scales 130–165. The gill rakers are rough, widely spaced, and total 16–30, pyloric caeca number 90–240, and vertebrae 67–75. Spawning males have a prolonged kype with enlarged teeth and an enlarged adipose fin. At sea the colour of the fusiform, streamlined body is iridescent green to blue green on the back, the top of the head, and the upper sides, silver on the sides below the lateral line, and white on the ventral surface. The lower gums are black, and the tongue is dark. Small black spots are present on the back, the upper sides, the top of the head, and all fins, including both lobes of the caudal fin. The adipose fin has a dark margin and a clear central area. Breeding males become olive brown to purple overall, or even dull yellow with a purple sheen, but are much darker than on fish in the sea. Females change less dramatically than do males. Chinook Salmon parr have 6–12 large, wide parr marks that extend well below the lateral line, and the dark area of the parr mark is equal to or greater than the light area between each parr mark. The length of adult Chinook Salmon at sea is variable but is usually between 839 mm and 915 mm. It attains 160.0 cm in total length and 61.2 kg in weight, but is rarely over 23.0 kg. The Chinook is the largest of the Pacific salmons (Tyee is the Chinook word for “chief ” or “large”).



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history and have developed into two races with little gene flow: the stream type and the ocean type. Ocean-type Chinook Salmon are found in North America south of 56° N, and the fry migrate to the ocean during their first year. Stream-type Chinook Salmon are found in Asia and northern North America, and the fry spend one or more years in fresh water before migrating to the ocean. Streamtype fish are found in areas distant from the ocean or are those with low growth opportunity. They have a longer time to grow and be strong enough for the migration and to be large enough to avoid some predators on a short migration. As Alaska populations are almost exclusively stream type, the vagrants found in the Arctic are likely stream-type Chinook Salmon and therefore are the focus of this description. Juvenile Chinook Salmon in fresh water do not eat fish but feed on terrestrial insects, crustaceans, chironomids, corixids, caddisflies, mites, spiders, aphids, Corethra larvae, and ants. Fingerlings move downstream in summer and spend time in the nearshore environment before moving to the ocean. Juveniles in the nearshore environment feed on chironomids. Adults in the marine environment feed primarily on other fishes including Pacific Herring, Sand Lances, and Smelts. Adults do not eat while in fresh water. Various other fishes including salmonids feed on Chinook, and adults at sea are taken by marine mammals, and by birds and bears on the spawning run. The maximum lifespan is nine years, and spawners are three to nine years old. Small three-year-olds are called jacks and are usually males. Chinook Salmon home to their natal stream to spawn; however, vagrants occur. Although the timing of migration and spawning varies throughout the geographic range of the species, in northern populations Chinook Salmon typically have a single peak of migratory activity during summer. In the Yukon River the Chinook Salmon spawn from July to September in gravel and cobble. Chinook usually spawn in deeper water and use larger gravel than do other Pacific salmon species. The female builds a large redd, up to 3.7 m long, in the gravel in an area of upwelling where there is little silt or sand and the water depth is between 0.05 m and 7.2 m. Females can build more than one nest and may be attended by

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Oncorhynchus tshawytscha, male silver

Oncorhynchus tshawytscha, male spawner

more than one male. The eggs are deposited and fertilized, and the female covers the redd. Sneaker males often rush in to fertilize some eggs. Adult Chinook die a few days to several weeks after spawning although some males may survive for five months. The eggs hatch in the spring, and the alevins remain in the gravel for two to three weeks. The egg number varies with the size of the female and stock, but in Alaska the average egg number is 8,517, and the size of the orange-red egg is large, approximately 6–7 mm in diameter. Emergent fry are found in areas of cover including back eddies of rivers, and near vegetation or river banks, and move to areas of current once they grow. Overwintering juveniles move from the tributaries into deep pools in the main stem.

importance:  Similarly to other species of Pacific salmon, significant fisheries occur for this species in southerly areas of its

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distribution. Chinook Salmon are extremely rare in the Canadian Arctic, likely due to their intolerance for low water temperatures. This is the most important ocean sport fish in British Columbia taken on trolled plugs, on spoons, by mooching herring, or on stationary live or dead bait and are fished deeper than other salmons. It does not leap as much as the Coho Salmon, but it is very strong and makes long sounding runs. As many as 93,000 are taken each year, mostly in the Strait of Georgia. Yukon River fish are important in the subsistence fishery taken by fish wheels and gill-nets. Commercial fisheries use trollers, purse seines, gill-nets, and long lines. The 1988 catch was 5,108 t, worth over $37 million. Some are sold fresh or frozen, but many are canned. The flesh is red or white, the former receiving a higher price.

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distribution:  Chinook Salmon range from the Anadyr River (Russia) to northern Hokkaido (Japan) in the western Pacific and from Point Hope (Alaska) to the Ventura River (California) in the eastern Pacific. The northern limit for spawning populations of Chinook Salmon in North America is the Pilgrim River in western Alaska and perhaps Kotzebue Sound. This species also occurs in drainages to the Pacific Ocean in southwestern Yukon. Chinook Salmon have been extensively introduced worldwide with limited success; the only self-supporting anadromous populations that resulted have been in New Zealand, although there has been some occasional spawning in Lake Superior. Continual stocking operations take place in the Laurentian Great Lakes for sport fishing. Single strays have been captured between Point Hope and Point Barrow, and also on the Colville River and in Prudhoe Bay, Alaska. In the western Canadian Arctic, Chinook Salmon have been captured on the Liard River, the Slave River, the Coppermine River, in subsistence fisheries near Aklavik, and in marine waters on the Yukon North Slope. Chinook Salmon may also have been captured in the Peel River and Arctic Red River in 1914. Map points represent river mouths that the fish must have passed on their upriver migrations. Spawning has not been recorded in the western Canadian Arctic, so it is unlikely that colonization has occurred.

Distribution of Oncorhynchus tshawytscha

sources:  Craig & Haldorson (1986); Dogrib Divisional Board of Education (1996); Quinn (2005); Stephenson (2005, 2006).

Prosopium cylindraceum (Pennant, 1784)

Round Whitefish, ménomini rond JAMES D. REIST

common names: Local names are Okeugnak (Great Bear Lake) and Osungnak (Back River) (Inuktitut); Aanaaliq, Anahluk, Kapisilik (Inuvialuktun); Savigunnaq in northern Alaska; and likely many other local names throughout the range. Other common names are Cross Whitefish, Frost Fish, Lake Minnow, Menominee Whitefish, and Pilot Fish. taxonomy:  The genus comes from the Greek prosopon (face or mask) in reference to the large bones in front of the eyes. The species comes from the Latin cylindraceus (like a cylinder) in reference to its body shape. The following are synonyms: Coregonus quadrilateralis Richardson, 1823, described from Fort Enterprise, Hudson Bay, and the Arctic Sea; Coregonus preblei Harper and Nichols, 1919, described from the Tazin River, about one mile above its confluence with Talston River, Mackenzie River basin; and Prosopium hearnei Fowler, 1948, described from the northwestern extremity of Nueltin Lake, Windy River, one-quarter mile above its mouth, southwestern Keewatin, Northwest Territories. Additional synonyms exist for Asian populations. Although the species was first described from eastern Siberian rivers as Salmo Cylindraceus, in North America it was first described as Coregonus quadrilateralis. Delineation of a distinct genus, Prosopium, was made due to the presence of basibranchial bones in the floor of the mouth, which are absent in Coregonus. Prosopium also has a notch below the rear margin of the pupil in the transparent membrane surrounding each eye. A significant range disjunction occurs in North American populations, with an eastern group (Ontario, Québec, and northeastern United States) and a northwestern group (northern Manitoba, Saskatchewan, Northwest Territories, and the Kivalliq and Kitikmeot areas of Nunavut). The latter has high mean counts of gill rakers (about 17–19) and pyloric caeca (about 94–106) compared to low counts for the former group; no clinal variation exists within forms. This likely reflects origins from two isolated refugia during glacial times (i.e., Beringian and Mississippian). Variation in habitat associations (lacustrine in southern areas; riverine in northern locations) and their tolerance of brackish waters add an additional layer of potential taxonomic diversity (i.e., lacustrine, adfluvial, fluvial, anadromous). The consequences of this, if any, have not been adequately investigated. description:  This species is distinguished by a single flap

between the nostrils, the body is rounded in cross-section, and young have parr marks (other coregonines lack this combination of the three characters).



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Prosopium cylindraceum, parr (top right)

This species is an elongate, medium-sized coregonine with a strongly forked tail. The snout is laterally pinched, the upper jaw extends beyond the lower, and the small mouth is downturned and lacks teeth. The head is short, about 20% of total length. Each eye is moderately sized, less than snout length, and maxillae extend almost to the front margin of the eye. Dorsal fin-rays number 11–15, anal fin-rays 9–13, pectoral fin-rays 14–17, and pelvic fin-rays 9–11. Lateral-line scales are 74–108, and usually there are 22–24 around the caudal peduncle. Short gill rakers number 14–21, and pyloric caeca number 50–130. Males have a prominent nuptial tubercle on each scale in the five scale rows above and below the lateral line in the spawning season. Generally this species is more heavily coloured than other coregonines, with a brown, bronze, or greenish back underlain by silvery white; the sides are silvery, and the venter is silvery white. The scales, especially those on the back, are well delineated and conspicuously lined by dark pigment. The pectoral, pelvic, and anal fins all tend to be amber in colour, darkening to reddish orange during spawning. Blackish dusting may be present on the dorsal and caudal fins and possibly also on the margins of other fins. Eastern specimens tend to have brown spotting on the adipose fin. Young exhibit distinct round or lengthwise oval black spotting in two to three rows of 7–13 spots along the sides, the dorsal one coalescing across the back. The lowest row is at the lateral line. This spotting is similar to the parr marks of salmonines. The species attains 56.1 cm in total length.

habitat:  Northern populations tend to be associated more closely with rivers than with lakes but also occur in the latter. They tend to occupy river reaches that are further inland; however, individuals

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are quite common in deltas, estuaries, and nearby freshened coastal areas associated with larger rivers (e.g., Mackenzie, Coppermine, Little Whale, and La Grande Rivers) draining to the sea. Detailed understanding of their habitat use in rivers in the north is generally poorly known. Anadromous forms use freshwater habitats for most life-history functions (spawning, early rearing); as larger juveniles and adults they probably undertake annual feeding migrations to brackish waters, but details of this remain unclear. Fishery gear that is set to catch other coregonines is inefficient at capturing Round Whitefish; thus, their coastal and estuarine occurrences are likely underestimated in the North.

biology:  Its food is primarily benthos in fresh water, including

insects, gastropods, and the eggs of other fishes; its marine diet is unknown but likely is opportunistic upon small estuarine benthos. Predators include larger fishes (e.g., Lake Trout) and perhaps aquatic mammals. Round Whitefish exhibit lacustrine, adfluvial, fluvial, and anadromous life-history types, with the latter occurring in deltas and estuaries. Adults live to about 14 years of age in lakes, are iteroparous, but likely live to only around 8 years in anadromous populations entering the sea. The details regarding sizes and ages of individuals occurring in marine areas are poorly known; however, it is likely that these are adults making feeding forays. Feeding occurs primarily in shallows. Spawning occurs from fall to early winter, primarily in lakes but occasionally in streams and rivers; similarly to other northern fishes, spawning is not likely to be annual. Generally this species spawns over substrates of gravel and rubble but may also spawn over

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sand and silt in areas with emergent vegetation. Spawning depths may range from 5 m to 10 m but are most often less than 1 m. The eggs are broadcast over the substrate and hatch in spring between March and May. The orange-coloured eggs, up to 20,000 per female, are up to 3.5 mm in diameter before fertilization.

Salmo salar Linnaeus, 1758

Atlantic Salmon, saumon atlantique J. BRIAN DEMPSON

importance:  Round Whitefish are generally not commercially fished in North America but were exploited earlier to a limited extent and also in Siberia where catches are likely higher. Similarly, it does not appear to be targeted by subsistence fishers; however, some by-catch likely occurs in fisheries for co-occurring species, although estimates are very limited. In some areas sport fishers may target this species, although the total catch is very limited.

distribution:  This species is found in Ungava, Hudson, and James Bays, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and the Beaufort Sea. The northwestern group is distributed from central Siberia eastwards through much of Chukotka, Alaska, Northwest Territories, and mainland Kitikmeot of Nunavut south of the Boothia Peninsula, thence southwards throughout the Kivalliq portion of Nunavut to the extreme northern margin of Manitoba. It is not known to occur on the southern margins of the Arctic Archipelago, but this is a possibility. The eastern group is distributed in northern Ontario (southwestern Hudson Bay), northern Québec (southeastern and eastern James and Hudson Bays), and Ungava Bay and Labrador. It is absent from extreme northern Québec.

Distribution of Prosopium cylindraceum

sources:  Mackay & Power (1968); Norden (1970); Jessop (1972);

Jessop & Power (1973); R. Morin et al. (1982); Kemp, Bernatchez, & Dodson (1989); D.B. Stewart, Carmichael, Sawatzky, Reist, & Mochnacz (2007); C. George et al. (2009).



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common names: Local names are Kavisilik, Kumaliq, Saâma, Saama, Saamakutaak, and Saamarug (Inuktitut); Utshashumek (Innu-Aimun); and Atlantikup Kapisilia (Danish/Greenlandic). Other common names are Black Salmon, Grilse, Lake Atlantic Salmon, Landlocked Salmon, Ouananiche, Salmon, Sebago, saumon d’eau douce, and saumon de l’Atlantique. taxonomy:  The genus is the original Latin name of this fish, which, like the species name, comes from the Latin salio (leap). The genus Salmo (or Atlantic salmons and trouts) is a well-defined group within the Salmonidae and consists of about five species distributed in temperate and sub-Arctic waters around the North Atlantic. The greatest natural diversity for the genus is in European waters, although wide-spread transplantation has altered this. Atlantic Salmon are recognized as a single species throughout the North Atlantic region. Previously, freshwater populations were thought to be taxonomically different from anadromous salmon, with some considered as either distinct species or subspecies. This resulted in subspecific designations as S. s. salar for anadromous salmon; and S. s. ouananiche McCarthy, 1894, described from the Saguenay River, Canada (outlet of Lake St John), and neighbouring waters, or S. s. sebago Girard, 1853, described from Lake Sebago, southern Maine, for resident populations. Morphological and genetic differences have been found between sympatric populations of anadromous and resident salmon although subspecific designations are no longer deemed necessary. However, occasional use of S. s. sebago still exists in some recent literature when referring to resident freshwater populations. Genetic analyses have shown that European and North American populations of salmon belong to two deeply divergent phylogeographic groups. description:  This species is distinguished by the teeth in the lower jaw being strong and conical; the lower jaw is long, extending back to or past mid-eye; the scales are small but obvious (19 or more from the dorsal fin origin to the lateral line, 109–124 in the lateral line); anal fin-rays, including small ones at origin, number 8–12; there are 15–20 gill rakers; the dorsal fin usually has dark or light spots; the flank spots are dark brown or black and X-shaped on a light background (except sea-run fish, which are silvery); the pelvic and anal fins lack leading white edges; and teeth are present on the head and the shaft of the vomer bone in the roof of the mouth. Dorsal fin-rays number 10–12; pectoral fin-rays 14–15; pelvic finrays 9–10; branchiostegal rays 10–13, usually 11–12 but slightly higher in Ungava Bay area; pyloric caeca 40–79, with Ungava Bay populations having the higher counts; and vertebrae 58–61. Spawning

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Salmo salar, adult male

males develop a kype. At sea Atlantic Salmon are trout-like in shape with elongated fusiform bodies, their colour being brown, green, or blue on the back, usually silvery along the laterally compressed sides, and white below. When fish return to fresh water for spawning, the silvery colour is lost, and they become darker, such that after spawning they are often referred to as “black salmon,” kelts, or slinks. Often numerous black X- or Y-shaped spots are present along the body, generally above the lateral line, and sometimes on the caudal fin. There are two to three large spots on the gill cover. Spawning males have a bronze to dark-brown colour and may have red, orange, or rust-brown spots on the head and body. The pectoral and caudal fins may darken. Atlantic Salmon parr often have 8–11 narrow parr marks, with a single red spot between the marks along the lateral line. Young fish migrating to the sea, known as smolts, racers, grilts, or fiddlers, are silvery. The closely related Brown Trout (S. trutta Linnaeus, 1758), introduced widely in North America but not recorded from Arctic waters, can be distinguished from Atlantic Salmon by the long maxillary bone that usually extends beyond the eye, the more numerous black spots that spread below the lateral line, and often the orange or orange-red adipose fin. Atlantic Salmon attain 150.0 cm in total length and 35.9 kg in weight.

habitat:  During the freshwater phases of its life history the

Atlantic Salmon is particularly adapted to fluvial habitats, although in some locations (e.g., Newfoundland, Labrador, Norway, Iceland, Finland, and Ireland) juveniles make extensive use also of lacustrine areas. Spawning occurs in fresh water in gravel areas where there is little silt and sand and where water depths are often less than 1 m. Eggs hatch in the spring after overwintering, and emergent fry use areas of relatively slow flow and where they

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can seek shelter. Substrate size, flow, and depth tend to increase with the size of Atlantic Salmon parr as they variously utilize riffle, run, and pool areas in streams. In lacustrine situations greater numbers of older and larger parr are found in deeper benthic and pelagic areas than in the littoral zone. Critical temperatures for survival of parr and smolt range from 22°C to 28°C (incipient survival over 7 days) and 30°C to 33°C (ultimate survival). Juveniles may also utilize estuarine areas. Both nearshore and offshore marine areas are important feeding areas for post-smolt and adult Atlantic Salmon. In the northwest Atlantic these fish are more commonly found in areas where sea-surface temperatures range from 4°C to 10°C. The lethal freezing temperature has been reported to be −0.76°C, and parts of the migration may occur at 1ºC, a lesser temperature than that favoured for feeding and growth. Greater growth has been demonstrated for fish caught between Greenland and Canada as the area bounding 4°C–8°C expanded, but this was overshadowed by mortality in other parts of the life cycle. In Ungava Bay, smolts start to migrate out of rivers in July, but the fish remain in estuaries until the sea warms up to 2°C–4ºC in August, when the adults return. Many smolts remain in estuaries until the thermal barrier between river and sea temperatures disappears in September. The Hudson Bay stock has a limited anadromous migration because smolts cannot swim to the Atlantic feeding grounds with the brief thermal window open to them. Post-smolts and adults at sea spend most of their time near the surface, especially at night, with some deeper excursions presumably related to feeding. Adults home to their natal river, perhaps using the earth’s magnetic field, ocean currents, or the stars to reach coastal waters, where smell directs them to their natal waters. They are able to leap waterfalls over 3.5 m high.

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Salmo salar, male spawner

Fish returning to fresh water after one year at sea are known as grilse and weigh 1.4–2.7 kg. Fish with two sea years are known as salmon and weigh 2.7–6.8 kg.

biology:  Atlantic Salmon display a wide range of phenotypic

plasticity and variability in life-history characteristics. In fresh water the food of small juvenile salmon changes with fish size and consists primarily of invertebrates. Common items include mayflies (Ephemeroptera), stoneflies (Plecoptera), chironomids (Chironomidae), caddisflies (Trichoptera), and black flies (Simuliidae). They also prey upon various terrestrial insects. Both drift and benthic feeding occur. At sea Atlantic Salmon are opportunistic feeders. Prey consists primarily of fish and fish larvae and planktonic crustaceans, with over 40 different species or species groups from at least 19 families documented. However, items such as Sand Lances (Ammodytidae), Capelin (Mallotus villosus), Atlantic Herring (Clupea harengus), cod (Gadidae), and crustaceans (Amphipoda, Euphausiacea) are the most common items found in the diet. Atlantic Salmon, especially as smolts, are fed on by seals, fishes, and various birds such as Gannets. Ungava Bay fish go to sea at four to eight years and 18 cm length. These fish may live up to 11 years, longer than Atlantic coast populations. Generally, Atlantic Salmon spawn in fresh water during the fall or early winter, and juveniles spend one to eight years in fresh water before migrating to sea as smolts (anadromous form) with a body size of 10–25 cm (weighing 10–80 g). Growth in the estuaries of Ungava Bay during summer approaches 0.2 cm/day, and condition increases at the same time. Smolt age has been found to increase with latitude. While some Atlantic Salmon may only spend a few months at sea before returning to fresh water to spawn, most



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spend one to three years (generally 1–10 kg, 45–135 cm) in the ocean. Female 2-sea-winter (i.e., having spent two winters at sea) salmon accounted for 75% of the fresh-run fish in the Leaf River, the most northerly population in Canada. In some north European areas Atlantic Salmon may spend four or five years at sea before returning to fresh water. The timing of return migrations to fresh water is highly variable. It differs among populations as well as within populations over time. In North America Atlantic Salmon return to rivers from May to November, although some runs may also occur in March or April. An Atlantic Salmon that had been tagged at west Greenland in October was caught in the Koksoak River of Ungava Bay in the following August. Atlantic Salmon are iteroparous, meaning they can spawn repeatedly. Some fish spawn as many as eight times, but very few survive the arduous migration and spawning act to spawn three times. Hence, various combinations of consecutive or alternate repeat-spawning individuals may also characterize upstream spawning runs. Fecundity is another trait that varies both among and within populations, with fecundity generally increasing in number with body size. Relative fecundity has been found to vary from about 1,300 to 2,500 eggs per kg. The orange- to amber-coloured eggs are up to 7.0 mm in diameter. The female excavates a redd in gravel bars and riffles that is up to 5.9 m long and 0.9 m wide. The male drives away competitors, and the eggs and sperm are shed as the pair quiver and gape. Sneaker males may rush in and fertilize some eggs. The female dislodges gravel from the upstream edge of the redd to bury the eggs over 25 cm deep. Spawned-out fish are known as kelts and may rest in a pool for several weeks, may overwinter, or may run to sea immediately. Young fish are called alevins and emerge from the gravel in May and

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June. Small salmon are called fingerlings or underyearlings and, when the characteristic flank marks have developed, are called parr. Resident freshwater populations exist in both North America and Europe and may occur in sympatry with anadromous Atlantic Salmon. Resident individuals may grow to over 10 kg in some populations. In addition, “dwarf ” salmon also exist where mature females range in length from 84 mm to 123 mm (Newfoundland) or from 136 mm to 228 mm in a Norwegian watershed (River Mellingselva), with the Newfoundland population having an average fecundity of 33 eggs.

include areas along the east and west coast of Greenland. Instances of Atlantic Salmon occurring in the Svalbard area and in southern Baffin Island, Canada, are becoming more common although spawning populations have not yet been documented. Atlantic Salmon escapees from Pacific coast aquaculture operations have been recently recorded in sub-Arctic waters (i.e., southern Bering Sea). It is unknown whether naturalized populations are becoming established in such areas, and thus the longer-term possibility of Atlantic Salmon occurrences in the marine waters of the western Canadian Arctic is unknown.

importance:  Directed commercial fisheries for Atlantic Salmon

at sea no longer occur in North American waters. Indigenous subsistence sea fisheries, however, still exist. Recreational fisheries are common, this being the quintessential sport fish. There is a voluminous popular literature as well as extensive research found in papers and books. Nominal annual harvests in the North Atlantic over the past decade range from about 1,000 to 3,000 t. In Europe approximately half the catch occurs at sea, the other half in fresh water. In contrast, North American fisheries are mostly riverine. In a number of southern areas, conservation problems persist for Atlantic Salmon such that all harvesting has been terminated. Farmed Atlantic Salmon continues to grow in importance, with upwards of one million tonnes produced in the North Atlantic region in recent years. The west Greenland area is very important for maintaining Canadian stocks. About 40% of Canadian fish visit the Greenland feeding grounds. Some travel as far as the Faroe Islands. This species is now being farmed in Atlantic Canada. Those in the Bay of Fundy were worth $16 million in 1987, with more recent figures exceeding $100 million. Salmon are being genetically manipulated to increase production. Incorporation of the “anti-freeze” gene from Winter Flounder, Pseudopleuronectes americanus (Walbaum, 1792), is aimed at increasing winter survival in nearshore waters, making cage culture more effective. The Committee on the Status of Endangered Wildlife in Canada has assessed it as “Data Deficient” in Nunavik.

distribution:  Atlantic Salmon occur naturally in watersheds that drain into the temperate and sub-Arctic areas of the North Atlantic. In the northwest Atlantic they continue to exist from New England in the south to Ungava Bay and Davis Strait in the north. An isolated anadromous population is known to occur in eastern Hudson Bay in the Nastapoka River, which is thought to have colonized that river from Ungava Bay by headwater exchange with the Koksoak River. General reports for James Bay are not confirmed by specimens. In the northeast Atlantic, Atlantic Salmon range from Portugal to rivers that empty into the Barents and White Sea areas of northern Europe and also rivers throughout the United Kingdom, Iceland, and Ireland. A single spawning population is known to occur in west Greenland. Past evidence suggests that during warm periods of the twentieth century Atlantic Salmon were more abundant in some Arctic locations (e.g., southern Greenland) and perhaps more widely distributed than they are at present. Ocean feeding areas cover a wide range of the North Atlantic and

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Distribution of Salmo salar

sources:  Dymond (1941); Nielsen (1961); Power (1969a, 1976);

Salonius (1971, 1973); Lee & Power (1976); Neilson & Gillis (1979); Power, Power, Dumas, & Gordon (1987); Claytor, MacCrimmon, & Gots (1991); R. Morin (1991); Brodeur & Busby (1998); Klemetsen et al. (2003); Reddin (2006); King et al. (2007); Webb, Verspoor, Aubin-Horth, Romakkaniemi, & Amiro (2007); Department of Fisheries and Oceans, Dartmouth, Nova Scotia & Québec Ministère des Ressources naturelles et de la Faune, Québec (2008); ICES (2010); Finstad, Armstrong, & Nislow (2011); Johansen, Erkinaro, & Amundsen (2011); Rikardsen & Dempson (2011); Friedland & Todd (2012).

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Salvelinus alpinus (Linnaeus, 1758)

Arctic Char, omble chevalier MICHAEL POWER and JAMES D. REIST

Salvelinus alpinus, male

common names:  Local names are Angmalook, Aniak, or

Aniaq (thin char coming from lakes), Aopalayâk, Aoparktulâyoq, Aupalijaat, Eekallûk, Eekalook, Ekalluk, Ekalupik, Ekaluppik, Eqaluk, Equaluk, Erlakukpik, Hiwiterro, Ihkaluk, Ikalopik, Ikalukpik, Iqaluakpak, Iqaluk, Iqalukpiaryuk, Iqalukpik (sea-run char), Iqaluppik, Iqaluupik, Irkaluk, Ivatarak, Ivisaruk, Ivitaaruq (brightred spawning phase), Ivitagok (red or spawning char), Ivitaroq, Ivitaruk, Kaïtilik, Kaloarpok, Nutilliajuk, Nutilliq (land-locked char), Suvaliviniq, Tadlulik, Tisuajuk (Inuktitut); Ekalukpik and Qaluaqpak (land-locked char), Evitaruk, Iqalukpik, Iqalukpiik, Iqalukpiit (sea-run char), and Qalukpik (for both Arctic Char and Dolly Varden) (Inuvialuktun); Shushashu (Innu-Aimun); Eqaluk and Kaporniangaq (Greenlandic); and Sùsàsù (land-locked char) (Cree). Other common names are Alpine Char, Arctic Salmon, Coppermine River Salmon, English Red Trout, Greenland Char, Mountain Char, Red Char, and Silver Char. “Char” is often written as “charr.”

taxonomy:  The genus comes from an old name for “char,” the

same route as the German Saibling. The species name comes from the Latin alpinus (alpine). The following are all synonyms: Salmo alipes Richardson, 1835, and Salmo nitidus Richardson, 1835, both described from lakes in Regent’s Inlet, Boothia Felix (Boothia Peninsula), Northwest Territories; Salmo Rossii Richardson, 1835, from Regent’s Inlet, Boothia Felix (Boothia Peninsula); Salmo arcturus Günther, 1877, from Victoria Lake (82°34' N) and Floeberg Beach (82°28' N), Arctic America; and Salmo Naresii Günther, 1877 (spelled “naresi” on a plate), from lakes near Discovery Bay, Arctic America. Salvelinus salvelinus (Linnaeus, 1758) – the same as S. umbla (Linnaeus, 1758) – described from Lintz, Austria, has been misused for Arctic Char in Canada. Hybrids between this species and Salvelinus namaycush are known from Arctic Canada. Originally described by Linnaeus as Salmo alpinus from high-altitude Swedish lakes, the Arctic Char forms a taxonomically diverse set of salmonid fishes distributed in Canada from latitudes as low as 46° N to as high as 83° N. Arctic Char form what is known by biologists as a “complex” as a result of the variation in



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morphological characteristics and life-history traits found in the species. It has been termed the most variable vertebrate on Earth. Arctic Char are recognized by most as a single species with several separate subspecies, although discussion regarding the existence of several separate species remains as a result of the disagreements between taxonomists with respect to what constitutes a species, and the interrelationships that might exist among the suggested species within the Arctic Char complex. Two subspecies are recognized within Canada. S. alpinus oquassa (Girard, 1854) is native to southern Canada and probably represents the first evolutionary line of Arctic Char in North America. This subspecies is primarily lake dwelling, occupying the profundal zone of a restricted number of freshwater lakes in southern Québec, New Brunswick, and northern New England. S. alpinus oquassa varies in coloration and size by lake, although it is generally less than 40 cm and rarely anadromous. In contrast, S. alpinus erythrinus (Georgi, 1775) is found throughout much of Arctic Canada and has a wide distribution that ranges from the Ob River basin in eastern Arctic Siberia to the eastern Canadian Arctic. The subspecies occupies a variety of lake and river habitats, exhibits a range of life-history types including anadromous, and varies in size depending upon the habitats (freshwater and/or marine) used for feeding and growth. Within freshwater habitats S. alpinus erythrinus also displays considerable variation in colour and size. Phylogeographic distributional patterns inferred from mitochondrial DNA analyses recognize three different lineages in Canada: the Atlantic lineage consisting of populations from Labrador and Newfoundland, the Arctic lineage consisting of populations from the Canadian Arctic, and the Acadian lineage consisting of populations from southern Québec. Within the lineages, S. alpinus erythrinus dominates in the Atlantic and the Arctic, and S. alpinus oquassa dominates in the Acadian area. Additional subspecies are present although the taxonomic relationships and the actual diversity are likely underestimated: S. alpinus taranetzi Kaganowsky, 1955, is present in Chukotka, Kamchatka, and western Alaska; and S. alpinus alpinus and S. alpinus salvelinus (= S. umbla, see above) are distributed throughout northern Europe and Russia and include many populations isolated in deeper lakes in the southerly areas of the range (e.g., Switzerland). Due to the disagreements as to what constitutes a species, many of the Eurasian forms are variously recognized as species, ecotypes, and/or other variants. Closely related, but apparently distinct, species of chars are also found in many areas of Siberia, Chukotka, and Transbaikalia. The notable variation in form, colour, and life-history patterns existing at the taxonomic (species or subspecies) level also exists within these taxa, with much of that variation being in life history and/or ecophenotype. Distinct life-history types are known within populations. After common juvenile residency in freshwater, individuals may become anadromous, migrating to the sea for summer feeding, or remain as freshwater residents. The age at first migration to sea varies by latitude, increasing from approximately two years in the south to as much as nine years in the north. Migration to and from the sea often occurs annually but may be episodic depending on environment and maturation schedules. Migrants typically grow more quickly, mature earlier, and produce a larger number of eggs

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Salvelinus alpinus, resident

than do freshwater residents, but are also likely to suffer increased risks of mortality as a result of migrating. Marine migrants of a wide range of sizes and ages are present in Arctic marine waters for varying periods (likely latitudinally and/or climatically correlated) each summer particularly between the southern range limit and about 75o N in North America. Migratory distances away from natal estuaries, tolerances to sea water, and distances moved offshore appear to be size dependent as well as determined by local environmental situations (e.g., salinities, mixing zones, temperatures). Differences in relative growth rates also have consequences for the body burdens of contaminants found in Arctic Char, with slower growing, often older freshwater residents having higher concentrations of trace metals in their tissues than do anadromous individuals. Variation in size within a single population may arise for reasons other than the use of different life-history strategies. In single-species lakes, piscivory (cannibalism) is thought to play a role in population regulation, with only a few individuals growing very large and the remaining fish growing only to an average (modal) length that is notably smaller. Within lakes, Arctic Char may also display considerable variation. The term “ecophenotype” is used to describe the multiple forms that may co-occur (exist in sympatry) in a lake. The forms will be distinct morphologically, genetically, or ecologically but may also be distinct in any combination of these characteristics. Ecophenotypes typically occur in lakes with restricted fish communities but have been found in lakes with as many as 12 other resident species. When ecophenotypes exist, there is often a small epibenthic form that feeds on benthic invertebrates, and a large pelagic form that feeds on zooplankton. A larger piscivorous form may also exist that predates on the other forms. Different degrees of reproductive separation exist among sympatric ecophenotypes, varying from a high degree of interbreeding to complete isolation. Genetic studies have indicated that in most cases the forms are repeatedly derived from a single original type in the lake, with form occurrence likely

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depending on ecological conditions such as the number of available niches present in the lake. Ecophenotypes generally occur in recently deglaciated lakes (i.e., ages of 10,000 years or less) and are thought by many biologists to represent the initial stages of evolution of new species. Although research has been poor, in some Arctic aquatic ecosystems it appears that both life-history variants (anadromous, resident) and ecophenotypes of resident forms (dwarf, normal) may occur. Moreover, it is unclear whether, or to what degree, switching among strategies may occur. These issues add to the complexity and variety of individuals potentially encountered in marine environments.

description:  As noted above, one of the most important char-

acteristics of Arctic Char is the degree of variation in coloration and morphology within and among populations. Such variation makes it hard to generalize for the purposes of identification. The description provided here, therefore, can at best serve only as a rough guide. The species can be distinguished from its congeners by the caudal fin being emarginate or square when spread out (it can be quite forked in young, cf. S. namaycush); the orange, pink, or red spots on the flank (whitish in S. namaycush), the spots usually being large at about eye diameter and not numerous (the spots are smaller than eye diameter and numerous in S. malma) (note that sea-run Arctic Char are silvery, but spots are still visible); pyloric caeca numbering 13–75 (mostly about 45, more than 80 in S. namaycush, and 13–47 –with means of 32 or less – in S. malma); dorsal and caudal fins without vermiculations, the white leading edge not usually being set off by black behind; the tip of the lower jaw and the roof of the mouth being whitish (cf. S. fontinalis); basibranchial teeth present (absent in S. fontinalis); parr marks that are irregular and not clearly defined (regular in S. fontinalis); and gill rakers numbering on the upper arch usually 7–13, on the lower arch 12–19 (mostly

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Salvelinus alpinus, silver female

about 13) (the upper arch is usually 3–13, the lower arch 8–15, mostly about 10, in S. malma). Arctic Char have an elongate body that is moderately compressed. The body comes to a point at the head, which is small in comparison to the overall body, 20%–25% of total body length. Arctic Char have large eyes that account for 16%–30% of overall head length. The mouth is large and can vary in position depending on the individual but is generally terminal in anadromous individuals. The maxillary extends to the posterior margin of the eye. There is a visible adipose fin and a distinctly forked caudal fin with rounded distal tips. The lateral line curves slightly upward towards the anterior of the fish. Principal or major rays in the dorsal fin number 10-12 with the total number of rays being 12-16, principal anal fin-rays 8-11 (11-15 total), pectoral fin-rays 14–16, pelvic fin-rays 9–11, lateral-line scales 123–152, and vertebrae 61–70. Mature male Arctic Char may develop a kype on the lower jaw, although its presence may vary geographically (e.g., it is rare in Labrador), and its size appears to be age dependent. Anadromous males also develop a dorsal ridge anterior to the dorsal fin, and enlarged teeth. The colour of the species is variable. In anadromous fish the dorsal area varies from dark blue to green, giving way to silver laterally and white ventrally. Mature males develop red coloration over the entire body area with the exception of the leading edges of the pectoral, pelvic, and anal fins, which turn a bright white. This coloration may be pale late in the summer while fish are still in marine areas but brightens upon their entering fresh water. The white of the leading edges may be retained throughout the non-breeding season. It is offset by following black-and-red bands, with the rest of the fin, and other fins, being dusky. Females also develop a red body coloration, although it is limited to the ventral areas. In post-spawning fish the red body coloration fades, and anadromous overwintering fish captured in fresh water will exhibit some lateral spotted coloration ranging from yellow to pink. In some populations the spots may be surrounded by a blue halo, a character that overlaps with Dolly



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Varden (S. malma) and appears to be variably present throughout the Canadian Arctic. The largest spots are usually larger than the pupil. Young fish have 8–17 oval parr marks on the flank. The species attains 101.6 cm in total length and 16.0 kg in weight.

habitat:  The Arctic Char is considered to be a habitat generalist and may be found at appropriate times of the year in a variety of aquatic habitats (lakes, streams, rivers, and the sea). There are fluvial and lacustrine stocks and stocks in which individuals migrate between both habitats. Within a habitat type more than one life-history form (ecophenotype) can live sympatrically with another. Lake occupancy, however, dominates throughout the Arctic distributional range, and north of about 75° latitude Arctic Char is the only fish species found in fresh water. Anadromous populations are numerous, particularly in the mid-range of the distribution where access to productive marine coastal margins is relatively easy. Few entirely riverine populations are known or described. Within lakes, Arctic Char will use all habitat types (e.g., pelagic, littoral, profundal), with usage dependent on age and life stage and the number of co-occurring species in the lake. Arctic Char appear pre-adapted to low aggression and will generally shift habitat usage in the face of interspecific competition. For example, separation between Arctic Char and sympatric Brook Trout (Salvelinus fontinalis) and Lake Trout (Salvelinus namaycush) populations has been observed in northern Québec lakes where feeding competition and predation risks are reduced by non-overlapping habitat use. In sympatry with Lake Trout, Arctic Char show faster growth rates, lower survival, and shorter longevities. High intraspecific competition may also result in life-stage-specific habitat shifts, with adult Arctic Char occupying lake littoral zones and juveniles being confined to deep-water profundal zones, shifting to the pelagic zone because with increasing size they become food limited. Stable isotope studies of feeding patterns in northern lakes have shown that resourceuse separation within and among size or morphotype groupings

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Salvelinus alpinus, anadromous male spawner

of Arctic Char has the effect of lowering resource competition, but it can impose developmental energetic constraints dictating limits on body size, maturation rate, and fecundity. Over most of the southern part of its range the Arctic Char, being a cold-water stenothermic species, will be found only in the cooler, profundal areas of lakes during the summer. Anadromous individuals share a common juvenile history with lake-resident fish but begin to migrate to the marine environment at between two and nine years of age depending on the population. In the marine environment Arctic Char occupy the nearshore areas of coastal bays and fiords for relatively short periods of time (30– 70 days). The extent and duration of migrations are influenced by fish size, maturity, environmental conditions, and prey availability. Individuals have been captured as much as 100 km from shore. Anadromy peaks in frequency in the mid latitudes of the geographic range and may be restricted by unfavourable trade-offs between the costs and the gains of migration (i.e., risks of mortality versus the benefits of greater growth and reproductive potential). It has been suggested that the southern limit of anadromy is controlled by temperature; for example, average coastal water temperatures measured as 10-day means exceeding 14°C at 4–5 m depths provide an effective barrier to anadromy because Arctic Char are the least resistant of salmonids to high temperatures. Studies from southern Canada, however, suggest that thermal barriers to migration do not affect anadromous behaviour, noting that differences in food-resource availability, decreases in predation or disease risks, and favourable abiotic conditions provide better explanations of the occurrence of anadromy. Arctic Char are known to be a cold-adapted species and can survive and feed close to 0°C. Detrimental growth effects of temperatures above 14°C have been noted. Measures of the physiological effects of temperature on specific growth relative to 14°C indicate significant positive correlations between growth and temperature

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up to 14°C, with sharp declines in growth thereafter. Recent studies of growth indicate that an optimum growth temperature of 15°C exists for age 0+ Arctic Char, and within 9°C–20°C there are no significant differences in measured growth rates when fish can feed at will.

biology:  While they are in marine areas, Arctic Char feed on

invertebrates, typically amphipods, or fish such as Sand Lances (Ammodytes spp.), Capelin (Mallotus villosus), resident cod (Gadus spp.) or Sculpins (Triglops spp. and Myoxocephalus spp.). Prey choice varies with fish size: smaller individuals (< 300 mm) tend to feed on invertebrates and small fish, and larger individuals rely to a greater degree on Capelin. Dietary studies of marine migrants have also shown that diets can shift over time as the relative abundance of prey items changes in response to environmental conditions. This species is eaten by Arctic Terns and seals and by Polar Bears in the Union River, Creswell Bay, Nunavut. They are also cannibals. Arctic Char are generally a long-lived fish, with northern lake-dwelling populations showing the greatest age range (e.g., to 30+ years). Anadromous populations generally have shorter lifespans (e.g., 20+ years) than do lacustrine populations, but are similarly long lived in comparison to many southern fishes. The sources and levels of natural mortality acting on Arctic Char populations are generally not well studied, and those studies that exist tend to focus on anadromous populations. The mean annual mortality reported for Greenland Arctic Char aged 3–20 years is in the range of 23%. Estimates of mean annual mortality from selected Canadian anadromous and lacustrine populations for 6- to 15-yearold fish indicate mortality rates in the 30%–45% range. Old age and post-spawning debilitation are thought to be the major causes of death of adult fish. Mortality in early life (i.e., from egg stage to age 3+) is relatively high and is most likely associated with environmental and density-dependent factors. Mortality during transition

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Salvelinus alpinus/malma

to migratory habitats for anadromous individuals up to first sexual maturity is similarly high. Mortality resulting from subsistence or commercial fisheries can be moderately high and for many populations remains the most significant single source of mortality. Changes in size structure are evident during the migration period, with larger maturing and non-maturing fish typically beginning to enter the sea first, followed by smaller adults and juveniles. Return migrations begin in mid-July and peak in August. Maturing fish, predominantly females, tend to leave the ocean first, followed by the smaller and non-maturing adults, and juveniles. The maximum duration of the sea-water phase observed in Labrador was 13 weeks, with averages ranging between 7.5 and 8.2 weeks depending on the population. Further north, the period of marine residency is reduced as a result of later ice-out and a reduced period of suitable marine conditions. The age at maturity in Arctic Char varies with latitude, increasing from three years in the south to ten years in the north, with females tending to mature one to two years later than males. Fecundity is correlated with length, which explains most of the observed variation in the fecundity of lacustrine and anadromous populations. Fecundity also declines as latitude increases. Lake



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environments appear to buffer some of the negative effect of latitude, with declines in the fecundity among lake-resident Arctic Char being less severe than among anadromous fish. Arctic Char are iteroparous, potentially reproducing several times during their life. Spawning occurs in fresh water in September and October, usually in lakes or in streams and rivers. Arctic Char may excavate redds, in which case they require an unconsolidated gravelly substrate. The eggs are often deposited in shallow pools below rapids in river gravel beds. Arctic Char may also broadcast spawn when the substrate is unsuitable for redd excavation. In streams and rivers, groundwater upwellings are probably required to protect the eggs from freezing; for example, upwelling ground water was present in sites used by anadromous Arctic Char in rivers in northern Québec. Where lakes are accessible and rivers are unsuitable, anadromous Arctic Char will spawn in lakes. The use of lakes may involve a two-year cycle of migration and maturation marked by entry into fresh water after marine feeding, overwintering in the lake, and maturing and spawning in the subsequent summer and autumn, such as that first described for the Nauyuk Lake–Willow Lake system in the Canadian central Arctic. In the Fraser River (Labrador), however, anadromous Arctic Char use both lake and

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stream habitats without first overwintering in fresh water. Anadromous Arctic Char do not necessarily spawn every year, with cycles of two to four years being known. Spawning anadromous Arctic Char tend to have a high fidelity rate to their natal systems, but there is also some straying of fish between river and lake systems. Unlike many other salmonids, there is no evidence of an effect of egg size on Arctic Char fecundity, which may result from the ability of Arctic Char to vary reproductive periodicity by “resting” for a year or more to gather sufficient energy for reproduction. The tactic is more typical of females as a result of the greater energy required for egg production. Despite the Arctic Char’s apparently long lifespans, the limitations imposed by short feeding seasons on acquiring surplus energy for gonadal development reduce the overall lifetime reproductive potential of the fish and may have significant implications for the exploitation and sustainability of many populations. The eggs hatch in the spring (April), but alevins remain in the gravel and continue to develop to emerge as free-swimming fry in July when planktonic prey normally become abundant. Arctic Char young of the year appear to be opportunistic users of available habitat, with the habitat use varying by locality. Juveniles may use profundal areas of lakes, the shallow margins of lakes, and streams and rivers where water velocities do not present substantive barriers to movement. Older juveniles are often precluded from using shallow-water habitats because of avian-predation risks and will migrate to deeper water habitats. In populations where anadromy occurs, physiological changes occur that facilitate the use of marine environments. Such changes are often accompanied by a silvering of body colour. Anadromy is facultative and varies depending on local conditions. Where anadromy does occur, the transition to marine feeding begins generally between the ages of two and nine years, with individuals feeding annually in the sea for a six- to eightweek period.

importance:  Arctic Char are the numerically most abundant

of the anadromous salmonids found in Canadian sub-Arctic and Arctic marine waters. As a result, Arctic Char form the basis of important subsistence fisheries throughout their range, particularly in the communities that border the waters of the Labrador Strait, Ungava and Hudson Bays, coastal Baffin Island, Victoria and Banks Islands, and the coastal areas of the northern continental coast west of Hudson Bay and east of the Mackenzie Delta. In northern Québec, Arctic Char are second only to Caribou in weight of country food consumed by Inuit, and at Kangiqsualujjuaq, Ungava Bay, they accounted for over 95% of fish captured. Archaeological evidence also points to the importance of anadromous Arctic Char fisheries to human populations throughout the history of human occupancy of the Arctic. Although Arctic Char have and continue to form the basis of important food fisheries, long-term investigations into the effects of exploitation on Arctic Char are generally lacking. Exceptions include longer-term (but episodic) studies at Nauyak Lake (Nunavut), in northern Labrador, and at Cambridge Bay (Nunavut). The importance of Arctic Char is matched by the extensive research effort and the accompanying literature, only a small part of which is cited in the “Sources” section

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of this account, although most literature refers to the freshwater part of the life cycle. Commercial exploitation of Arctic Char is primarily focused in the areas around Cambridge Bay and Pangnirtung in the central and eastern Arctic, respectively. Additional commercial fisheries are scattered throughout the Arctic. The best studies of the effects of exploitation are those available for the fishery in northern Labrador where Arctic Char have been commercially fished for over a century. Although Arctic Char have traditionally been characterized as vulnerable to exploitation owing to their relatively slow growth rates and late sexual maturity, evidence from the fishery at Nain provides a factual counterpoint. Peak annual catches during the late 1970s and early 1980s represented between 50,000 and 90,000 Arctic Char per year in the fisheries on the Voisey, Nain, and Okak stock complexes. Those exploitation levels have not resulted in the short-term population collapses traditionally predicted for intensely fished stocks. Over the past three decades Arctic Char catches in the dominant age classes (7–10 years) have shown little variation in mean length at capture. The similarity in length and age–composition data reported for 1953 indicate that over the span of a half-century the age-and-length composition of Arctic Char taken in the commercial fishery of north Labrador has remained relatively constant. Constancy in key biological metrics has been interpreted as indicative of the capacity of the north Labrador region to produce and sustain substantive anadromous Arctic Char fisheries over significant periods of time. There are some sport fisheries for Arctic Char, but they are expensive because of travel costs and are easily fished out. The trophy sport fishery of Tree River (Kogluktualuk), Nunavut, produced 0.4 char per angler hour. Arctic Char are strong fighters, and smaller fish may leap. They can be caught on streamers, spoons, and dry flies. The management of Arctic Char populations remains challenging in the face of incomplete harvest information and a lack of sufficient allocated scientific resources to explore alternative management initiatives. Variable year-class strength, the noted lack of directional change in length or age structure in well-studied populations that have been fished, the absence of established biological reference points, and the complexity of Arctic Char migratory patterns that see fish move among rivers in local areas further complicate management of anadromous stocks. Human-population-driven pressures for increased access to Arctic Char in many northern communities and the likely impacts of climate change on Arctic Char will require the development of research programs aimed at better understanding the long-term dynamics of Arctic Char populations.

distribution:  Arctic Char are wide spread in the Canadian Arctic Archipelago and in the rivers and lakes draining the Labrador and Ungava peninsulas, in Hudson Bay drainages as far south as Kuujjuarapik in the east and the Nelson River in the west, and in the northward-draining watersheds that discharge along the northern continental margin of Canada east of the Mackenzie River. Only a few isolated populations (land-locked) occur west of the

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Mackenzie River estuary; however, much of the earlier literature on chars from this area confuses Arctic Char with Dolly Varden. Throughout most of the Canadian range anadromous populations predominate where access to the sea is possible. In the High Arctic, particularly on Ellesmere Island, anadromy is less common and may occur only intermittently as environmental conditions allow. The northernmost established case of anadromy occurs from Clements Markham Lake on northern Ellesmere Island (83° N). Where access to the sea is limited, lake populations dominate, and numerous lake-dwelling populations are found throughout the Arctic Archipelago, Labrador, and northern Québec and across the continental portions of Nunavut and the Northwest Territories east of the Mackenzie River. Southern populations of Arctic Char on insular Newfoundland and in southern Québec watersheds bordering the St Lawrence River or lakes in New Brunswick are generally lake dwelling, although anadromy has been established for populations on insular Newfoundland (Parkers Pond) and on the north shore of the Gulf of St Lawrence (Rivière de la Trinité). Arctic Char are also found on all coasts of Greenland.

Baroudy & Elliott (1994); Marshall, Heuring, & Babaluk (1994); Dempson (1995); C.C. Wilson, Hebert, Reist, & Dempson (1996); Reist, Johnson, & Carmichael (1997); Beddow, Deary, & McKinley (1998); Larsson & Berglund (1998); Marshall, Layton, & Babaluk (1998); Doucett, Power, Power, Caron, & Reist (1999); Hammar (2000); Brunner, Douglas, Osinov, Wilson, & Bernatchez (2001); Jonsson & Jonsson (2001); Dempson, Shears, & Bloom (2002); Guiguer, Reist, Power, & Babaluk (2002); Johnson (2002); Klemetsen et al. (2003); Day & de March (2004); Power, Dempson, Reist, Schwarz, & Power (2005); Reist, Power, & Dempson (2006); Reist, Wrona, Prowse, Power, Dempson, King, & Beamish (2006); Dyck & Romberg (2007); Dempson, Shears, Furey, & Bloom (2008); Power, Reist, & Dempson (2008); Power, Power, Reist, & Bajno (2009); Knopp (2010); Reist & Sawatzky (2010); Reist, Majewski, Atchison, Geoffroy, Loseto, & Young (2012); Klemetsen (2013); Reist, Power, & Dempson (2013).

Salvelinus fontinalis (Mitchell, 1814)

Brook Trout, omble de fontaine MICHAEL POWER

common names: Local names are Âait, Aanak, Âna, Anâtlik, Ânna, Anokik, Anuk, and Nutilliq (Inuktitut); Màsimàkus, Màsimàkush, Màsimekush, and Màsimekw (Cree); and Matameku (Innu-Aimun). Other common names are Brook Char(r), Brookie, Coaster, Eastern Brook Trout, Mud Trout, Redspotted Trout, Salter, Sea Trout, Slob, Speckled Trout, Squaretail, truite, truite de mer, and truite mouchetée.

Distribution of Salvelinus alpinus

sources:  Sprules (1952); Grainger (1953); Andrews & Lear (1956); Backus (1957); McPhail (1961); Swift (1964); Moore & Moore (1974c); Qadri (1974); Marshal (1977, 1981); Power (1978, 2002); Behnke (1980, 1984, 1989); Johnson (1980, 1983a, 1989); Jensen (1981); Jobling (1983); Gyselman (1984); Johnson & Burns (1984); Marshall & Layton (1984); Dempson & Green (1985); Marshall & Layton (1985a); Cunjak, Power, & Barton (1986); Dutil (1986); Dempson & Kristofferson (1987); Power & Barton (1987); Berg & Berg (1989); Boivin, Power, & Barton (1989); Fraser & Power (1989); Macmillan (1989); Sigurjónsdóttir & Gunnarsson (1989); Heuring, Babaluk, & Marshall (1991); Brännäs & Wiklund (1992); Wilson & Hebert (1993);



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taxonomy:  The species name comes from the Latin fontinalis (living in springs). Salmo hudsonicus Suckley, 1861, described from Hudson Bay and tributaries, New York U.S.A (sic, after Catalog of Fishes and presumably not Hudson Bay in Arctic Canada), Labrador and Newfoundland, Canada, is a synonym. This taxon was used formerly for sea-run Brook Trout. Brook Trout is recognized as a single, stable species throughout its North American distribution, and compared to other related species there is little controversy associated with its classification. Nevertheless, Brook Trout display variability in growth rate, colour, and meristic characteristics throughout the range. Although sea-run Brook Trout were initially considered to be a separate species, modern taxonomic and genetic investigations have not supported the suggestion, and freshwater resident and anadromous forms should be regarded as life-history variants of a single species. Genetic evidence of differences among stocks exists, with experimentation having shown that northern populations have an

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Salvelinus fontinalis, parr (bottom right)

enhanced genetic capacity for growth and longevity compared to southern populations.

description:  Distinguishing characters from congeners are given under S. alpinus. Brook Trout have a laterally compressed body and a comparatively large head. The body depth is greatest anterior of the dorsal fin. The snout is rounded, and the mouth terminal. A large lower jaw is evident, with the maxillary extending beyond the posterior margin of the eye. The scales are cycloid, small, and readily evident above the straight lateral line. Dorsal fin-rays number 9–14, anal finrays 8–13, pectoral fin-rays 10–15, pelvic fin-rays 7–10, lateral-line scales 109–132 (195–243 in mid-lateral series), gill rakers 13–22, pyloric caeca 20–55, and vertebrae 57–62. In fresh water Brook Trout have a dark olive-green to brown dorsal area dominated by an intricate pattern of cream-coloured wavy lines (vermiculations) that change laterally to a pattern of red spots surrounded by blue halos. Vermiculations are also evident on the dorsal fin. The ventral area is white. The jaw tips and the roof of the mouth are blackish. The pelvic, pectoral, and anal fins are dark in colour but may retain some of the orange pigmentation distinctive of the juvenile phase. As with other char species (e.g., Arctic Char, S. alpinus) the ventral fins of the Brook Trout are notable for having a distinctive leading white edge. The caudal peduncle is relatively thick, with the largely square caudal fin forming only a shallow fork. In marine environments Brook Trout take on a distinctive, overall silvery colour with a blue-green back and sometimes purplish flank tinges. Only the red spotting of their freshwater coloration is visible. Marine colouring is lost quickly on freshwater re-entry. Spawning males are much brighter and have orange-red lower flanks and upper belly, bordered

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below by black on each side, which delimits the white belly. Young have 6–12 brown parr marks, the widest being equal to the eye diameter, and small red, yellow, or blue flank spots. The species attains 70.0 cm in total length and 6.57 kg in weight.

habitat:  Although Brook Trout commonly occur in fully saline

waters (and are called “salters”), the distances travelled in the marine environment remain uncertain, and the evidence suggests that they do not reside for extended periods of time in full-strength sea water. Congregations of anadromous Brook Trout are typically found at river mouths along the Hudson and Ungava Bay coasts and down the coast of Labrador. Two forms of sea-run Brook Trout along the Labrador coast have been distinguished: an estuarine form that inhabits the brackish waters of river mouths and estuaries, and a sea-run form that makes extended migrations away from rivers. There is no evidence of long-distance movements in sea water (e.g., in Richmond Gulf). Anadromous Brook Trout appear to remain in coastal areas influenced by river discharge in warm surface waters of salinities of approximately 20‰. In the marine environment Brook Trout remain in the mixing zone of river estuaries or within the influence of the freshwater plume of river discharge in nearshore coastal environments. Upstream return migrations in the north begin in July, with peak movement occurring in August. Nearly all fish will re-enter fresh water by the end of August, with immature individuals remaining in marine-influenced environments until early September. Brook Trout largely return to their natal river, although studies have shown that natal homing is not complete. All Brook Trout initially live and overwinter in fresh water, using both fluvial and lacustrine habitats, feeding on a variety of

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Salvelinus fontinalis, male

invertebrates, and switching increasingly to fish as they grow. Temperatures are usually less than 20ºC. Brook Trout may be either sedentary or migratory. Sedentary strategists will spend their whole lives within relatively confined areas, some as small as 100 sq m. Migratory strategists will make regular movements between feeding and overwintering habitats, particularly as older individuals. Movement from stream or river habitat to lakes occurs in the second or third summer when lengths of 80–150 mm have been attained. Estuarine or marine migrations occur in May as soon as the river ice clears, and are undertaken for feeding.

biology:  Brook Trout display an extreme degree of morphological plasticity shaped by the glacial history of the species and its subsequent dispersal and separation into isolated breeding populations in a wide variety of habitats. Anadromous Brook Trout are largely piscivorous, feeding on available smaller marine fish species such as Capelin (Mallotus villosus), Sand Lances (Ammodytes spp.), and Sculpins (Myoxocephalus spp.). Anadromous Brook Trout also feed on amphipods, isopods, and nereid worms, with the latter prey being most prevalent in August when the worms are engaged in mating activities. Brook Trout are cannibals on their own eggs and young and are eaten by a wide variety of other fishes, water snakes, turtles, various birds, and otters. Brook Trout may complete their life cycles entirely within fresh water in as little as three years in the south. They can adopt freshwater resident or anadromous life-history tactics in the north and live for 11+ years. Sea-run fish have growth rates more than twice those of fish in fresh water. The maximum lifespan is over 20 years.



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Brook Trout spawn in the autumn to early winter (August to December) in well-oxygenated gravels, often at the downstream ends of pools. Lake spawning may also occur in some southern populations. Sea-run Brook Trout enter their natal stream in spring and summer, even though spawning occurs in autumn. A female cleans a redd of debris by turning on her side and lashing her tail. Redd depth between stones is tested by inserting the anal fin. Redd construction may take two full days. Courtship involves gentle pushes, touches, and strokes of the female by the male. The female is ready to spawn when she crouches in the redd with her genital area between the stones. The male arches and may press the female against the redd bottom. Both fish vibrate, and eggs and sperm are shed. Sneaker males may rush in and fertilize some eggs. The female lashes her tail to push eggs into the gravel and then dislodges gravel with her anal fin to cover them to depths as great as 20 cm. The yellow-orange-coloured eggs are up to 5.0 mm in diameter and number up to about 17,000 per female, although averages range from the low hundreds to a few thousand. Both sexes may spawn again with other fish. The eggs develop over winter, taking 165 days at 2.8ºC, but only 47 days at 10ºC. Temperatures above 11ºC will kill the eggs.

importance:  There are no commercial fisheries for Brook Trout,

although there are substantial recreational and subsistence fisheries for the species. They can be caught on a wide variety of lures and flies, fighting well but not leaping, and are often quite small. Inland southern populations are often supplemented with hatchery programs or stocked specifically for fishing access. Populations in the north are natural and are fished most intensively by recreational sport fishing in freshwater environments. Anadromous Brook Trout in the north are taken in subsistence fisheries but are

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less important as a food resource than either Arctic Char or Atlantic Salmon. Northern anadromous stocks generally appear to be numerically small. The Koksoak River (58°31' N, 68°10' W), however, is an exception, where the Indigenous harvest has been estimated at 17,250 fish, or 15,500 kg/yr. As a result of generally small numbers the risk of over-exploitation of anadromous populations remains.

distribution:  The Brook Trout is found in waters bordering Ungava, Hudson, and James Bays, the southern Hudson Strait, and the Labrador coast both as freshwater and anadromous life-history types. In the west anadromy does not extend north of 60° N. In the east the northern portion of the Ungava Peninsula north of 60° N is also largely unoccupied by Brook Trout. Incomplete natal homing, coupled with the predicted impacts of climate change, may result in the northward colonizations along both Hudson Bay coasts and perhaps into the southern archipelago in the years to come. The Brook Trout is indigenous to eastern North America and may be found in abundance in the Canadian provinces of Ontario, Québec, Newfoundland and Labrador, New Brunswick, and Nova Scotia. Brook Trout are also prevalent in the northern states of the Unites States westward as far as Minnesota. Extensive transplanting has greatly extended the westward ranges in southern latitudes. It has also been introduced to South America, Europe, Asia, and Australasia.

Distribution of Salvelinus fontinalis

sources:  Backus (1957); Weir (1979); Dutil & Power (1980); Power (1980a).

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Salvelinus malma (Walbaum, 1792)

Dolly Varden, omble malma JAMES D. REIST

common names:  Local names are Evitaruk, Iqaluakpak, Iqaluqpik, Ivitaaruq, Qaluaqpaq, Qalukpik (Inuvialuktun); and Dhik’ii (Gwich’in). Other common names are Arctic Char, Bull Char, Pacific Brook Trout, Red Char, Red-spotted Rocky Mountain Char, Salmon-trout, Sea-trout, and Western Brook Trout; and Dolly Varden (in French). taxonomy:  The species name comes from a vernacular name in Kamchatka. Salmo Hearnii Richardson, 1823, described from Bloody Fall, Coppermine River, 67° N, is a possible synonym (see the “Distribution” section). “Dolly Varden” is the name of a Dickensian character in Barnaby Rudge who was colourfully dressed. A pink-spotted calico was called Dolly Varden during Charles Dickens’s visit to North America, and the char was likened to the material. Two subspecies of Dolly Varden in North America have been formally differentiated on the basis of allopatric distributions and taxonomic characters: S. m. lordi (Günther, 1866), the southern form, has lower gill-raker and vertebral numbers and a karyotype of 2n = 82 and occupies Pacific drainages south of the Alaska Peninsula to Washington, well outside of our area of consideration; and S. m. malma, the northern form, has higher gill-raker and vertebral numbers and a karyotype of 2n = 78 and occupies Bering, Chukchi, and Beaufort sea drainages north of the Alaska Peninsula and east to the Mackenzie River. However, recent evidence suggests that a third form is present in the Bering Sea drainages of the Alaska Peninsula north to the Norton Sound drainages of the Seward Peninsula of western Alaska; thus, the true northern form appears to occur in drainages from Kotzebue Sound north, then east to the western Canadian Arctic. The southern subspecies has been confused with Bull Trout – S. confluentus (Suckley, 1859), a freshwater taxon from western North America – but these are now known to be distinct. The northern subspecies and the intermediate form have been confused with Arctic Char (S. alpinus) although recognized as distinct within that taxon. Recent scientific work has clearly differentiated northern Dolly Varden from Arctic Char, but the earlier scientific literature confused them. These species have also not been differentiated, until recently, by local fishery managers or generally by resource users. This taxon is closely related to Arctic Char and is viewed by many, especially Russian workers, as simply a member of that species complex (sensu lato). description:  Distinguishing characters from congeners are given under S. alpinus. Canadian populations of the northern subspecies exhibit the following counts: lower gill rakers, range 8–15, means from 11.7 to 12.9; upper gill rakers, range 3–13, means from 8.4 to 9.8; pyloric caeca,

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Salvelinus malma, residual male

Salvelinus malma, silver

range 13–47, means from 20.7 to 31.7; and vertebrae 60–71. Isolated and residual forms tend to exhibit lower counts within the above values. These values are low in comparison to and distinguish this taxon from Arctic Char found to the east of the Mackenzie River. Generally, dorsal fin-rays number 10–12, anal fin-rays 9–13, pectoral fin-rays 14–16, pelvic fin-rays 8–11, and lateral-line scales 105–142. Other characteristics such as colour and body form, though highly variable, also tend to distinguish these species. Northern form, anadromous Dolly Varden are somewhat laterally compressed (more so in large males and at spawning time) and have relatively short and broad caudal peduncles, large caudal fins typically with little or no fork and with acute tips, and pinkish or light spots over the body that are never larger than the pupil and are surrounded by pale light-coloured halos. The body colour of non-spawning anadromous fish (large juveniles, adults of both sexes while resting and at sea) tends to be olive green on the back, grading to silvery sides and silver or white below. Spots are very pale and generally lost in the silvery colour. Larger spawning anadromous males develop strong colours: dark green and black on the body dorsally, grading to green laterally, then to bright red ventrally; the head is black overall with green tinges, bright white highlights are present in the angle of the



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mouth, the gular region, between the branchiostegals, and along the anterior edge of the pectoral girdle, a patch of red develops on the snout around the notch formed to receive the kype, and the kype is well developed in only the larger males; spots on the body are bright red to pink and surrounded by strong blue halos, those near and below the lateral line tend to be larger and may exceed the diameter of the pupil but never are eye-sized; and paired fins, the anal fin, and the lower border of the caudal fin tend to be black suffused by red, with a strong white leading edge followed by a strong black line. Anadromous females develop similar strong coloration, but this is never as vivid as in the males. The back and sides are olive green, grading to white below the lateral line and on the belly. The small pink spots become more vivid, and the blue of the halos deepens, as spawning time approaches. Paired fins and the anal fin develop a white leading edge that tends to be followed by a distinct line or blackish smudge over the fin, and the trailing edges tend to be suffused by a pale orange or reddish colour. The caudal fin tends to be uniformly dark grey with greenish highlights, a whitish lower leading edge may develop, and, a tendency to orange suffusion may occur on the trailing edge. The head is olive green dorsally and laterally, grading to white ventrally. Highlights on the head tend to be

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Salvelinus malma, silver male

Salvelinus malma, silver female

blackish smudges on the operculum and branchiostegals with intervening white patches. The kype may be present but is typically not well developed in females, although a patch of orange is usually present on the tip of the snout and possibly also the tip of the lower jaw. Pre-spawners for that year of both sexes may initiate development of the above colour patterns during the late summer while they are still at sea. Young fish have 8–12 dark parr marks, wider than the spaces between them. The species attains 128.0 cm in total length and 18.3 kg in weight.

habitat:  During the freshwater phases of its life history the

northern form of Dolly Varden is a taxon adapted primarily to high-gradient, relatively pristine rivers with low turbidity, although lake residency is suspected in at least one Alaskan population. Critical riverine freshwater habitats are associated with perennial springs located in headwater areas that provide spawning and overwintering requirements. Riverine freshwater habitats down to and including freshened estuarine zones provide summer nursery

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habitat for the first few years of life. Estuarine and nearshore marine habitats provide summer feeding areas for larger (post-smolt) juveniles and adults. Some offshore marine habitats are likely used for feeding, especially the warmer, freshened surface waters, but there is little documentation of this. Preferred marine habitats seem to be those of lower salinity and higher temperature than the surrounding, more marine waters. The northern form primarily exhibits seasonal anadromy in the western Arctic. Due to hypersaline conditions and marine water temperatures of below 0°C in winter, anadromy is seasonally exhibited only during the summer (more properly termed “amphidromy”). Thus, concerted downstream migrations out of the rivers are seen in the spring, and upstream migrations back into rivers are seen in the autumn. Coastal migrations along the nearshore are seen throughout the summer, with movements of up to 3–6 km/day. In addition to the anadromous life history, two additional life-history types are known to be associated with fresh water: a residual or resident type consisting of dwarf males that never go to sea but

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Salvelinus malma, anadromous male spawner

Salvelinus malma, anadromous female spawner

participate in spawning with anadromous pairs; and isolated types in some rivers above barriers to migration. Isolated or semi-isolated types also exist in some inland drainages.

biology:  The food of young and small juveniles in fresh water

consists primarily of insect larvae (chironomids, plecopterans, and trichopterans). Large juveniles and adults are opportunistic predators in marine areas, consuming amphipods, mysids, and fish. Larger adults primarily feed upon fish that include nearshore marine species such as Sculpins and Arctic Cod, as well as anadromous species such as Arctic Cisco, Broad Whitefish, and smaller members of their own species. They are eaten by larger fish, various birds, and seals and by various mammals such as Grizzly Bears, Grey Wolves, and American Mink at spawning and overwintering sites. They are also parasitized by Arctic Lamprey. Anadromous fish in Arctic Canada live to approximately 15 years of age, attaining a length of about 70 cm. Generally the first few years of life occur in the natal river, likely characterized by the



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fish’s dispersal throughout the system in the summer to feed. All the rivers in this area freeze to the bottom throughout most of their lengths during winter, so the return movements by the young of the year and small (pre-smolt) juveniles to spring-fed overwintering sites occurs in the autumn. Smoltification generally occurs between three and five years of age, and large (post-smolt) juveniles forage during the summer in nearshore environments for the next several years. These individuals return to fresh water each autumn to overwinter in upstream spring-fed areas. Sexual maturity is attained between six and eight years of age. Spawning occurs in the same spring-fed upstream areas that are used for overwintering, in areas with suitably sized gravel substrate. Redds are likely constructed and defended primarily by females, and spawning anadromous pairs are closely attended by several resident males that likely follow a sneaker strategy to fertilize a significant portion of the eggs. Fecundity is correlated with body length and differs among females from different rivers, ranging from 1,100 to 5,000 eggs in Canadian rivers. The egg size is approximately 4.2–4.8

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Salvelinus malma, large male

mm. These are iteroparous fish capable of spawning several times during their lifetime. In the Canadian Arctic, generally the anadromous fish do not spawn in consecutive years, presumably due to energy constraints. This is likely especially true for females. In contrast, the residual males as well as the isolated-life-history type appear to spawn consecutively in many cases.

were traditionally smoked, dried, and frozen for later use, highly valued, and used mostly as human food. Sport fishing is limited primarily due to the remote nature of the area, although potential exists. The Committee on the Status of Endangered Wildlife in Canada has assessed freshwater western Arctic populations as “Special Concern.”

importance:  The northern form of this species is the only char

distribution:  This species is primarily distributed in freshwater drainages of the North Pacific from Washington State to Alaska and from Chukotka to Kamchatka and in mainland drainages to the Sea of Okhotsk, as well as on islands (Aleutians, Kurils, Sakhalin, etc.) south to Hokkaido, Japan. The northern form is confirmed as only occurring in Arctic Canada in rivers that drain into the west side of the lower Mackenzie River (Rat, Vittrekwa, and Peel Rivers) and the Beaufort Sea (Big Fish, Babbage, and Firth Rivers), as well as seasonally in the nearshore Beaufort Sea. This form also occurs in suitable North Slope Alaskan rivers west and south at least to the north side of the Seward Peninsula. Other subspecies comprise the remaining North American distribution, which includes both coastal and interior (isolated?) taxa. So far as is known, this species does not occur

in the coastal areas of the Arctic west of the Mackenzie River and supports local Indigenous subsistence fisheries in this area. Such fisheries are conducted at traditional coastal locations during the summer, on upstream migrants at freshwater locations during the autumn, and occasionally on overwintering populations during the winter. Canadian harvests are typically about 2,000–5,000 fish, but this is highly variable and has been much lower in recent years. Coastal migrants that originate in Canadian rivers are also domestically fished to some extent in Alaska, and the reverse likely happens to some degree as well. No current commercial exploitation occurs, although some attempts have been made. The lack of success has been due in part to the high transportation costs to market and to the small population sizes, hence limited quotas. The fish

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in areas of the Canadian Arctic to the east of the Mackenzie River. However, two situations require further study: occasional char captured in fresh waters from easterly locations (e.g., Prince Albert Sound area of Victoria Island) that appear to represent this species; and individuals captured in several mainland rivers in the Coronation Gulf area (e.g., Coppermine, Kugaryuak, and Tree Rivers) that exhibit morphotypes of either Arctic Char or Dolly Varden. Genetically the situation is similarly complex, with genotypes not matching the expected phenotypes. This may represent co-occurrence of both species with interspecific hybridization and introgression, or some other situation, the resolution of which requires more research.

Salvelinus namaycush (Walbaum, 1792)

Lake Trout, touladi HEIDI K. SWANSON

common names: Local names are Tahiqmi Ihugaqyut (Innuinaqtun); Col-lic-puk, Idlôk, Iluuraq, and Isiuralittaaq (Inuktitut); Ehohok, Ehok, Ihok, and Iqaluakpak (Inuvialuktun); Singayuriaq (Inuvialuktun – Paulatuk); Vit (Gwich’in); Higayuriaq, Kùkamàs, Kùkamàsh, Kùkamàw, Kùkamesh, and Namekush (Cree); and Kukumes (Innu-Aimun). Other common names are Forktail Trout, Grey Trout, Lake Char(r), Laker, Mackinaw, Mountain Trout, Salmon Trout, Siscowet, Taque, Togue, and truite grise. taxonomy:  The species name comes from the Cree and Ojibway names for this fish. Salmo namaycush was described from Hudson Bay lakes, Canada. The Lake Trout has been placed in the genera Salmo Linnaeus, 1758, and Cristivomer Gill and Jordan, 1878. The variation in generic associations primarily reflects earlier uncertainty in taxonomy and also an interpretation that Lake Trout appear to be distinct from other chars. description:  This species is distinguished from its congeners by

Distribution of Salvelinus malma

sources:  McPhail (1961); Cavender (1978); Behnke (1980, 1984); McCart (1980); Morrow (1980a); Reist (1989a, 1989b); Haas & McPhail (1991); Reist, Johnson, & Carmichael (1991, 1997); Phillips, Gudex, Westrich, & DeCicco (1999); Reist, Low, Johnson, & McDowell (2002); COSEWIC (2010a); Kowalchuk, Reist, Bajno, & Sawatzky (2010); Kowalchuk, Sawatzky, & Reist (2010); Mochnacz, Ghamry, Enders, & Reist (2010).



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having a deeply forked caudal fin, more than 80 pyloric caeca, and irregular light (whitish, rarely orange) markings on the flank and back (see also under S. alpinus). An anadromous form of Lake Trout has only recently been described (thus explaining the inclusion of this primarily freshwater species in this marine fishes guide), and as yet there are no external characteristics that can be used to differentiate between anadromous and freshwater forms. Lake Trout display considerable variation in external morphology and colour both among and within lake systems. This variation can be considerable within some lakes (e.g., Great Bear Lake) and is poorly explored in the north; however, to date the anadromous forms from different systems appear to all exhibit similar morphology. In general, Lake Trout colour ranges from silver to olive to a greenish grey or a black or brownish grey. The dorsum is usually darkest and, along with the top of the head, may have light-coloured vermiculations. Pale roundish spots are present over the whole body with the exception of the throat, belly, and paired fins. Red spots are absent. The belly is a dirty white to yellow. The pectoral, pelvic, anal, and caudal fins can show some orange coloration, and the pectoral, pelvic, and anal fins can have whitish leading edges that are less pronounced than those found in congeneric Brook Char and Arctic Char and lack a contrasting black bar behind. Breeding males develop a black flank stripe, a more reddish-brown body, and reddish lower fins. The jaw tips and the roof of the mouth become whitish. Young have 5–12 parr marks, with the spaces between being equal or greater in width.

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Salvelinus namaycush, oceanic

Salvelinus namaycush, Hood River estuary

As with most Salvelinus species, the coloration of the anadromous form during migrations and while in the sea tends to be muted, only becoming more vivid in mature adults in fresh waters near to spawning times. Lake Trout are generally elongate and quite slender, but large fish can be deep bodied. Anecdotal evidence suggests that anadromous Lake Trout are more deep bodied than their freshwater counterparts, but further research is required. The head is large, dorsally broad, and approximately 20%–28% of total body length. The eye, near the top of the head, is approximately 12%–20% of head length. Lake Trout have a large terminal mouth with strong teeth on the tongue, the roof of the mouth, and both upper and lower jaws. When their mouth is closed, the snout may be slightly longer than the lower jaw. Fin size varies greatly among systems. The dorsal fin of Lake Trout is placed further back than on other Salvelinus species. Males and females have tiny nuptial tubercles around the anus.

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Meristic characters for the species are dorsal fin-rays numbering 8–10, anal fin-rays 8–10, pectoral fin-rays 12–17, pelvic fin-rays 8–11, lateral-line scales 116–138, gill rakers 16–26, pyloric caeca 81–210, and vertebrae 61–69. The species attains 126.0 cm in total length and 46.3 kg in weight.

habitat:  Marine habitats used by anadromous Lake Trout are not

well characterized. It appears likely, however, that Lake Trout are more common in brackish coastal waters than originally thought and that they prefer such habitats rather than fully saline areas. In Arctic coastal waters, summer migrations to sea appear to last for approximately three to six weeks during July–September. A review of Arctic field observations suggests that anadromous Lake Trout occur most commonly in waters with 6‰–9‰ salinities, are rare in waters with 10‰–12‰ salinities, and are not generally observed in waters with salinity greater than 13‰. Laboratory studies indicate,

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however, that Lake Trout are capable of surviving full-strength sea water (at least for short periods of time).

biology:  Populations composed of wholly anadromous life history types of Lake Trout have yet to be discovered. Anadromous Lake Trout have been found in populations that are partially anadromous; that is, some individuals within the population migrate to sea whereas others do not. This is similar to other char species distributed coastally in the Arctic. Compared to freshwater residents, anadromous Lake Trout from the same populations are in better condition and exhibit a greater degree of piscivory. Common marine prey items for anadromous Lake Trout include Saffron Cod (Eleginus gracilis), Capelin (Mallotus villosus), and Pacific Herring (Clupea pallasii). Recent stable isotope evidence suggests that marine prey items constitute approximately 66% of anadromous Lake Trout diet, with freshwater prey items constituting the remainder. Anadromous Lake Trout have only been found in populations that are at the extreme northern end of the species’ range. Along with the fact that Lake Trout tend to occupy cold, oligotrophic systems when they are in fresh waters, this leads to relatively slow growth. The age of maturity has yet to be examined for anadromous Lake Trout. In freshwater Lake Trout the age at maturity varies from about 4 to 13 years. The largest confirmed anadromous Lake Trout to date is 915 mm (fork length) and 9.5 kg, but larger sizes are possible. Anadromous Lake Trout inhabit fresh water for the majority of their life cycle, and, similar to congeneric Arctic Char, they use marine habitats only as feeding grounds during summer. Spawning is intermittent and occurs in freshwater boulder-and-rubble habitat. No redd is built although the spawning ground is cleaned with body and tail brushes or by rubbing with the snout. Late September to late October is the most likely time for spawning by anadromous Lake Trout, but further research is required. The emergence of fry occurs in spring. It appears that Lake Trout require significant rearing time in fresh water before anadromous migrations begin. A recent study from the Canadian Arctic found that the age at first sea migration of anadromous Lake Trout varied from 3 to 29 years. importance:  The importance of anadromous Lake Trout is difficult to assess because (1) they have only recently been discovered, (2) they are not externally distinguishable from their freshwater counterparts, and (3) information summarized in reports and literature likely represents both anadromous and freshwater forms. In general, Lake Trout are very important to commercial, sport, and subsistence fisheries. Given that anadromous Lake Trout appear to be limited to coastal Arctic regions, it is likely that they are harvested in subsistence fisheries. Anadromous Lake Trout no doubt contribute to the overall productivity of partially anadromous populations. Researchers have speculated that partial anadromy in fish populations is a form of “bet-hedging” that allows populations to persist and/or colonize extreme and variable environments. The contribution of anadromous Lake Trout to the species’ persistence and distribution in the Arctic has not been quantified and is likely variable. However, in four lakes in the central Canadian Arctic it was found that



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anadromous individuals comprised about 13%–40% of partially anadromous populations. In fresh water, Lake Trout are important sport fish taken with flies and lures in cool seasons and by deep trolling in summer. They may also be caught by ice fishing. Commercial freshwater catches are particularly important in the Northwest Territories and, with Lake Whitefish, make up 95% of the total catch there. The total Canadian catch in 1988 was 1,050 t.

distribution:  Research on the distribution of anadromous Lake Trout is ongoing. Anecdotal evidence suggests that the range of anadromous Lake Trout extends along the North American Arctic coast from Bristol Bay, Alaska (57° N, 160° W), to Nain, Labrador (57° N, 61° W). Scientific studies have confirmed the presence of Lake Trout in the central Canadian Arctic (West Kitikmeot region, Nunavut, 68° N, 107° W). Preliminary findings from follow-up research indicate that anadromous Lake Trout are present in several other coastal systems ranging from Mackenzie Bay, Northwest Territories (70° N, 135° W), to Rankin Inlet, Nunavut (63° N, 92° W). The distribution map shows estuarine occurrences, only some of which may eventually prove to be anadromous.

Distribution of Salvelinus namaycush

sources:  Boulva & Simard (1968); Marshall & Keleher (1970); Marshall (1978, 1999b); Martin & Olver, in Balon (1980); Olver & Martin (1984); Marshall & Layton (1985b, 1995); Nelson & Paetz (1992); Hiroi & McCormick (2007); Swanson (2009); Swanson, Kidd, Babaluk, et al. (2010); Swanson, Kidd, & Reist (2011).

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Stenodus leucichthys (Güldenstädt, 1772) Inconnu, inconnu JAMES D. REIST

common names:  Local names are Si-airryuk, Sierak, Siiraq,

Teirark, and Tiktalerk (Inuktitut); Higaq, Sigaq, Siirgarq, and Siiraq (Inuvialuktun); and Shruh, Shryuh, and Sruh (Gwich’in). Other common names are Coney or Connie, locally in the western Arctic; and Eskimo Tarpon, Sheefish, and Shovelnose Whitefish. The common name in French recognizes the unknown aspect of this fish when it was encountered by the early European explorers.

taxonomy:  The genus comes from the Greek stenos (narrow) and

odous (tooth), in recognition of the fine dentition retained as adults (i.e., the small teeth of juveniles of other coregonines are resorbed and absent in the adults). The species name comes from the Greek leucos (white) and ichthys (fish), in reference to the overall colour. Salmo leucichthys Güldenstädt, 1772, described from the Volga and Ural Rivers of the Caspian Sea basin, is the taxon used in most modern literature for Canadian populations, sometimes as a subspecies based on Salmo nelma Pallas, 1773, described from larger Siberian rivers. Both species have been recognized as distinct, with Stenodus leucichthys being restricted to the Caspian Sea basin and Stenodus nelma being the North American species. However the American Fisheries Society list retains Stenodus leucichthys until data are presented to confirm distinction. Life-history variants and some disjunct distributions occur, so the potential exists for further taxonomic differences. Salmo Mackenzii Richardson, 1823, described from the Mackenzie River system including Salt River, is a synonym.

description:  This species is distinguished by its pike-like body and the presence of teeth in the lower jaw; the mouth is very large; the maxilla is long and relatively narrow (extending to the rear margin of the pupil); and the lower jaw obviously projects beyond the upper jaw. Inconnu are very large, cisco-like fishes with an elongate body that is relatively deep but laterally compressed. The head is long, about 25% of standard length, with moderately sized eyes. Teeth are present on the lower jaw, premaxillae, maxillae, tongue, vomer, and palatines. Dorsal fin-rays number 11–19, anal fin-rays 14–19, pectoral fin-rays 14–17, and pelvic fin-rays 11–12. There are 90–115 lateral-line scales. Gill rakers number 17–24, and pyloric caeca 144–211. Its colour is dark green to brownish on the dorsal surface, grading to silver and white below, but overall lightly coloured. The dorsal and caudal fins are typically dusky or tipped with black; other fins are immaculate. The young lack parr marks, and generally their colour is similar to that of adults. The species attains 100.0 cm in length and over 28.5 kg in weight, and 150.0 cm and 40.0 kg in Eurasia.

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habitat:  Similar to other coregonines of northwestern North

America, Inconnu appear to prefer river systems with high turbidities for much of their life history. They also appear to be preferentially associated with very large, slow-moving rivers. Turbidity, in turn, is associated with both the size of the system and the nature of the landscape drained. In this case the Mackenzie and Anderson Rivers and their tributaries drain large areas of sedimentary landscapes, appearing ideal for these fishes. Migratory pathways along slow-moving rivers appear to be important to life history. The spawners of the year leave the downstream overwintering sites prior to spring break-up and appear to make summer-long upstream migrations to spawning areas. They spawn in late summer or early fall in upstream reaches of rivers tributary to lakes (e.g., Great Slave Lake) or the Mackenzie River (e.g., Arctic Red, Peel, and Liard Rivers), likely in habitats similar to those of other coregonines. Egg deposition, similarly to other coregonines, appears to coincide with the decreasing local turbidities resulting from decreased autumn precipitation and bank stabilization after early freeze-up. Spent fish move downstream near freeze-up (October throughout much of the lower Mackenzie and its tributaries) to overwinter in the outer delta and perhaps the freshened areas nearshore. Although they are defined as anadromous, the salinities at capture sites along the coast are generally lower (0–15 psu) than those for other coregonines, suggesting a strictly coastal association with highly freshened waters. This species thus is perhaps best defined as amphidromous. The details of spawning locations, habitats used during early life history, and nursery areas for young fish are poorly known. Fish of about 10.0 cm in length or greater are present in low abundances and scattered distributions in the Mackenzie Delta and coastal nearshore areas.

biology:  The diet of non-anadromous fish varies with age from

insects and crustaceans in young to a wide range of co-occurring fishes in adults. The diets of anadromous fish in Canada are poorly known; those fish from inland Alaskan locations consumed both freshwater and anadromous fishes. Whitefishes, Sculpins, flounders, Pacific Herrings, and Arctic Lampreys are diet items. Incidents of cannibalism have been documented for the non-anadromous form. At least two, and perhaps three, life-history forms exist. Those in the Mackenzie River appear to be primarily anadromous. Fish in Great Slave Lake and associated rivers entering the southern shore of the lake are adfluvial, that is, utilize the rivers for spawning and the lake for feeding. A wholly lacustrine form and/or a wholly riverine form associated with larger rivers such as the Liard River may also exist. These likely intermix during life histories. The anadromous migratory form makes long-distance journeys (about 1,800 km) between the Liard River (northern British Columbia), other rivers tributary to the Mackenzie basin, and the outer Mackenzie Delta and Tuktoyaktuk Peninsula. It is unknown what proportion of these anadromous populations undertake migrations, or how many distinct populations may be present in the Mackenzie River and its tributaries. Late-summer upstream migrations appear to be protracted, and therefore no concerted “pulse” of migrating pre-spawning fish has been observed (unlike most other coregonines in the area).

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Stenodus leucichthys

Anadromous fish mature at around age six, with males doing so earlier than females (e.g., 7 versus 11 years in Arctic Red River). Ages up to 22 years have been estimated from scales for fish in the Mackenzie River although it is likely that Inconnu live much longer. Spawning occurs in autumn, and concerted downstream migrations of spent adults are observed, especially of the adfluvial form entering Great Slave Lake. Spawning temperatures are 1.5ºC–4.6ºC. Spawning intervals are two to four years for individuals, with up to 420,000 eggs per female. The ecology of young Inconnu is particularly problematic, and it is unclear how long the emerging young remain associated with their natal areas, although some have been captured in coastal locations. Larger individuals appear to seasonally occupy coastal areas along the outer Mackenzie Delta.

abundant again in the Mackenzie River basin. It is not present on the Arctic Archipelago islands. It occurs throughout the Mackenzie and Liard systems to northern British Columbia and in many of the rivers flowing to the southern shore of Great Slave Lake including the Slave River, but it is not known to occur in Alberta.

importance:  Inconnu have oily flesh, which contributes to

wide perceptions regarding their quality and suitability for human food. This is likely seasonally dependent and may also vary with life-history type. Similar to other coregonines, Inconnu were and are an important component of subsistence and domestic (household) fisheries wherever they occur. Split and dried or smoked fish are used for human and dog food throughout the area. Recent estimates suggest that subsistence fisheries for the anadromous type are greater than 10,000 individuals annually (1988–97 figures), that is, about 25,000 kg, using an average estimate of 2.5 kg per fish. At that time commercial fisheries were estimated to capture an additional 1,000 individuals (2,500 kg) annually. Limited sport fishing also occurs, given the size attained by some fish. The growth rate, size, diet, local abundance, and migratory characteristics all suggest that it is a key component of aquatic ecosystems; however, the dearth of information contributes to under-estimation of its importance.

distribution:  This species is found in the Beaufort Sea, east to the Anderson River, and is wide spread from the White Sea of northern Russia across Siberia and Chukotka to southern Kamchatka. It is abundant in Alaskan Bering Sea drainages (e.g., Kuskokwim and Yukon Rivers) but less so in Chukchi Sea drainages. The Inconnu occurs occasionally in Alaskan north-coast drainages, becoming



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Distribution of Stenodus leucichthys

sources:  Melville (1914); Johansen (1926); Alt (1965, 1969, 1973a, 1977, 1981b, 1987, 1988); Reist & Bond (1988); Howland (1996); Department of Fisheries and Oceans (1998a, 1998b); Fisheries Joint Management Committee (2000a); Howland, Tallman, & Tonn (2000); Howland, Tonn, Babaluk, & Tallman (2001), Howland, Chiperzak, Tallman, Tonn, & Gendron (2003); S.A. Stephenson, Burrows, & Babaluk (2005); C. George et al. (2009).

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Family Gonostomatidae Bristlemouths, Cyclothones

Brian W. Coad

Bristlemouths, Lightfishes, and Anglemouths are found in all seas including the Arctic and Antarctic Oceans. There are about 26 species, including 17 reported from Canada, of which 2 are found in Arctic waters. The maximum size is about 36 cm although most species are less than 8 cm. These fishes have a delicate, elongate body, with or without an adipose fin. Photophores, or light organs, are present in a regular row along the belly, on the branchiostegals (not shown in the illustration), and variably on the isthmus. These are black rimmed and emit red or green light. The mouth is large, with long jaws reaching back behind the eye at an angle, and it is armed with bristle-like teeth. The nostrils are high on the head and prominent in a dorsal view. The gill rakers are usually long. There is no chin barbel. The black skin and cycloid scales are often rubbed off during capture. It is rare that a bristlemouth is captured in good condition because

it is so fragile. Identification of individuals is often difficult as a result, but systematists usually have very large samples to study and can describe species by filling in the gaps. Some Bristlemouths are almost transparent. Males have larger olfactory organs than those of females. The overall pigmentation varies from colourless to brown or black. Bristlemouths are probably the most abundant fishes in the world in terms of numbers of individuals. However, they are deepsea fishes, mostly meso- and bathypelagic, and are usually seen only by scientists. Adults and young are found at between 200 m and 300 m by day. Small fish are found in shallower water than larger fish. Larvae are near the surface but sink as they develop. Some Bristlemouths do not migrate, and their gas bladder is poorly developed; others are partial or daily vertical migrants. They feed on various crustaceans, fry of other fishes, and their own family members and are an important food for other fishes.

sources:  Grey (1960, 1964); Mukhacheva (1972, 1974); Miya & Nishida (2000).

Cyclothone microdon (Günther, 1878)

Veiled Anglemouth, cyclothone à petites dents

common names: A local name is Småtandet Rundflab (Danish/Greenlandic). Other common names are Scaled Eye-nosed Fish, Small-toothed Bristlemouth, Small-toothed Lanternfish, Veiled Bristlemouth, cyclothone jaune, and cyclothone brun.

Abbreviations for photophores

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Cyclothone microdon

taxonomy:  The genus comes from the Greek kyklos (round) and

with each female producing up to 10,000 eggs with a diameter of 0.5 mm. The number of eggs increases in higher latitudes. Larvae metamorphose at 11–14 mm standard length.

description:  This species is distinguished by not having a white mass in the fold of skin at the rear of the upper jaw and by having gill rakers numbering 19–23 and anal fin-rays 16–20, usually 19. The eyes are minute. The anus is nearer to the pelvic fin bases than to the anal fin origin. Photophores are ORB 1, OP 2, BR 9–11, IV 2–3 + 10, VAV 4–5, AC 13–15, OA 6–10, and IC 29–33. Scales are present, about 27–31 or more along the flank. There is no lateral line. Gill rakers are visible in the open mouth, and gill filaments are fused. Dorsal fin-rays number 12–15, usually 14, pectoral fin-rays 8–10, and pelvic fin-rays 5–6. The adipose fin is absent. The overall colour is brown to brown black or jet-black with dense, star-shaped pigment cells on the head, body, and fins. The snout is pigmented diffusely, and the branchiostegal membranes are uniformly pigmented. Females reach 7.6 cm in total length, but males only reach 4.9 cm.

importance:  It is not economically important.

othone (veil) or kyklothen (around, in a circle). The species name comes from the Greek mikros (small) and odous (tooth).

distribution:  It is found in Baffin Bay and Davis Strait; northwest, southwest, and southeast Greenland; and in the Atlantic, Indian, and Pacific Oceans including off Atlantic Canada.

habitat:  This species is widely distributed off eastern Canada

including as shallow as 105 m in the Davis Strait. Adults and young are mesopelagic to bathypelagic, found generally at 200–2,700 m, but precise depth varies with age, season, and latitude. Some individuals have been caught down to 5,301 m off Bermuda. Larvae are in the upper 50 m. The Veiled Anglemouth has been caught at 330– 1,510 m and 0.0°C–3.9°C in the Davis Strait and southern Baffin Bay. There is no daily vertical migration. This is an extremely abundant species found in large, mid-water schools. Over 82.2% of all fishes from the deep sea in the Atlantic Ocean, and nearly 55% in the Pacific Ocean, are this species. Distribution of Cyclothone microdon

biology:  Its food is mainly copepods and euphausiids. It is a major food item for many other fish species and is eaten by Rock Grenadiers in the Davis Strait. In this species sex reversal occurs at 22–42 mm in standard length. Males become females (protandrous hermaphroditism). Spawning occurs in the summer and autumn,



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sources:  Hildebrand (1948); Karrer (1973); McKelvie (1989);

Jørgensen (1996); Treble, Brodie, Bowering, & Jørgensen (2000); Jørgensen et al. (2005).

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Sigmops bathyphilus (Vaillant, 1884)

Spark Anglemouth, gonostome étincelé

common names: A local name is Stor Laksesilding (Danish/

Greenlandic). Other common names are Deepsea Fangjaw and Deepsea Lightfish.

taxonomy:  The genus comes from the Greek letter sigma (S) and

-ops (having an eye or face indicated by the stem word). The species name comes from the Greek bathys (deep) and phileo (to love), meaning deep loving. It was formerly in the genus Gonostoma Rafinesque, 1810.

description:  This species is distinguished by having a white mass in a fold of skin at the rear of the upper jaw and by having gill rakers numbering 24–28 and anal fin-rays 21–26. The eyes are moderate in size. The anus is nearer to the anal fin than to the pelvic fin bases. Photophores are minute and usually obsolete in adults: SO 1, ORB 1, OP 3, BR 9–11, IV 11–13, VAV 4–5, OA 14–16, AC 18–22, and IC 32–38. Dorsal fin-rays number 11–15, pectoral fin-rays 7–14, and

pelvic fin-rays 7–9. The adipose fin is present. The overall colour is black. Ripe males have an excessively developed olfactory structure. Females of the species attain 20.0 cm in total length, and males 15.0 cm.

habitat:  The young and adults are bathypelagic and are generally

found at 700–2,700 m, depending on size, over the whole species range. This species is said to be more common below 2,000 m than above this depth. There is no daily vertical migration. A Canadian Arctic record was taken in Davis Strait over a bottom depth of 1,019 m and a bottom temperature of 3.6°C, and a second record at 1,407 m and 3.0°C. Another was caught at 754 m in Davis Strait.

biology:  Its food is crustaceans. Sex reversal occurs at 50–100

mm in standard length although some males are supermales and do not change to females. These supermales are the principal spawners and are mature at 10 cm or more. Females spawn first at 11–12 cm.

importance:  It is not economically important. distribution:  This species has been found in Davis Strait off southern Baffin Island as five records from cruise data and ARC 8705027, southwest and southeast Greenland, and in all temperate to tropical oceans.

Sigmops bathyphilus

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Family Sternoptychidae Marine Hatchetfishes, haches d’argent

Brian W. Coad

Distribution of Sigmops bathyphilus

sources:  See the family sources and the bibliography.

The Marine or Silver Hatchetfish Family comprises small, deep-sea fishes found in all oceans. There are about 67 species, including 12 found in Canada, of which one is found in the Arctic. The family is divided into two subfamilies, the Maurolicinae having an elongate body, 19–38 anal fin-rays, and light organs on the isthmus, with six on the branchiostegal membrane (not in Arctic Canada); and the Sternoptychinae having an eponymous hatchet-shaped, deep, and very compressed body, an almost vertical mouth, eyes that are sometimes telescopic, an unusual blade in front of the dorsal fin that is formed from fused fin-spine bases, 11–19 anal fin-rays, and a keel-like abdomen. The scales are easily detached on capture. The light organs are large, black, and silvery, vertically elongate, and arranged in clusters and have species distinctive patterns. An adipose fin is present, rarely absent. There are 6–10 branchiostegal rays, three of which are on the epihyal, and a pseudobranch, and the gas bladder is well developed. These fishes, being mesopelagic, are numerous in oceanic waters, although some are bathypelagic or even benthic. They are found as deep as possibly 3,658 m, although most are found higher up in the water column, the youngest at the highest. Some species are vertical migrators, and others do not migrate at all. The light organs serve for species recognition by their own species and to merge their image with light from the ocean surface so that a predator below cannot separate them from the diffuse light above. Hatchetfishes are an important food for many larger fishes.

source:  Baird (1971).



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Argyropelecus gigas Norman, 1930

Greater Silver Hatchetfish, grande hache d’argent

common names: A local name is Stor Sølvøkse (Danish/Greenlandic). Another common name is Giant Hatchetfish.

taxonomy:  The genus comes from the Greek argyros (silver) and pelekys (hatchet, axe). The species name comes from the Greek gigas (giant). description:  This species is distinguished by its hatchet-shaped and very compressed body; an unusual blade in front of the dorsal fin formed from fused fin spines (its height being more than one third of its length); 12 abdominal light organs; and telescopic eyes (directed upwards). Dorsal fin-rays number 9–10, anal fin-rays 12–13, pectoral fin-rays 10–11, and total gill rakers 18–21. The lower light organs (supra-abdominal, pre-anal, anal, and subcaudal) are arranged in a nearly continuous horizontal row (not broken into groups at different levels). The body profile below the third and fourth dorsal spines

is raised. There is a prominent sphenotic spine, directed laterally, near the dorso-posterior corner of the eye. The ventral keel scales form an evident flap below the photophores. This fish often produces heavy mucus secretions. The overall colour is silvery. The flank in preserved fish is light brown with small mid-flank spots, partially outlining muscle blocks. The light organs are silvery and edged with black. A large Marine Hatchetfish, it exceeds 12.0 cm in standard length.

habitat:  The species is found mesopelagically at about 300–1,000

m, mostly between 400 m and 600 m without any evidence of vertical migration. The single Arctic Canadian specimen was caught at 1,096–1,104 m and 3.4ºC.

biology:  Its biology is unknown in detail. Adults acquire a full complement of photophores at 1.4 cm standard length.

importance:  It is not economically important. distribution:  The species has been found in Davis Strait as a single record (CMNFI 2002-0039 at 62°14'24" N, 58°34'18" W), presumably an expatriate, and in southwest Greenland. It is also found in all oceans, except the northeastern Pacific.

Argyropelecus gigas

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Family Stomiidae Dragonfishes, Dragons à écailles

Brian W. Coad

Distribution of Argyropelecus gigas

sources:  See the family source and the bibliography.

The deep-sea Family Stomiidae is found in all oceans. There are 228 species, including 64 reported from Canada, of which 5 reach Arctic waters. The family includes the black dragonfishes, loosejaws, scaleless black dragonfishes, snaggletooths, viperfishes, and the scaly dragonfishes that were all once placed in separate families. Its maximum size is about 41 cm in length. They are characterized by various internal characters such as a lack of gill rakers in adults (some have teeth on the gill arch); a reduced or absent mesopterygoid bone; only one infra-orbital bone (related families have two to six); one or no supramaxillae; and light organs that are without a lumen or ducts. Most have a chin barbel that is longer than the head, and large, pointed teeth that are developed as fangs in some. Scales are present in the viperfishes and scaly dragonfishes (Chauliodus, Stomias) but absent in other family members. The scales are easily lost or dissolved in preservative, leaving only a typical hexagonal pattern along the flank. There are light organs on the lower part of the body in rows, behind the eye, and, in some species, on the hexagonal areas in various patterns. The photophores in some species are in two obvious rows low on the body – the lateral or OA row (made up of pre-pelvic [OV] and post-pelvic [VAL]), the ventral or IC row (made up of pre-pectoral [IP], pre-pelvic [PV], and post-pelvic [VAV]) – and a row between these two ending at the tail (AC). The dorsal fin may be centrally placed on the back, far anterior, or far posterior. A dorsal adipose fin is present in some species and absent in others, and some species also have a ventral adipose fin. These fishes are quite common in mesopelagic to bathypelagic waters, down to about 4,000 m. They are only rarely seen alive, and, like most deep-sea fishes, their delicate bodies are damaged in the long haul to the surface in nets. Dragonfishes are predators, swallowing their prey whole. They have no commercial importance.

sources:  Gibbs (1964); Morrow (1964a, 1964b); Parin & Novikova (1974).



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Astronesthes cf. richardsoni (Poey, 1852)

Richardson’s Snaggletooth, dragon-saumon de Richardson

common names: None.

biology:  Its biology is unknown. importance:  It is not economically important. distribution:  The species has been reported as a single record from Davis Strait at 61.5558° N, 60.7475° W, off southern Baffin Island, based on cruise data, and in the North and South Atlantic Oceans and the Gulf of Mexico.

taxonomy:  The genus comes from the Greek astroy (star) and esthio (to eat). This species is named after Sir John Richardson (1787–1865), a British naturalist, an Arctic explorer, and the describer of the genus Astronesthes. The material collected in Canadian waters is believed to be this species but was not identified with certainty. description:  Astronesthes cf. richardsoni is distinguished from other snaggletooths by a lack of scales; no true gill rakers; teeth present on the gill arch in groups of 2–3 (13–17 total); and 8–24 minute teeth on the rear part of the upper jaw (maxillary bone) that are close together (comb-like) and slanting rearwards. Dorsal fin-rays number 12–15, anal fin-rays 13–18, pectoral finrays 6–9, and pelvic fin-rays 7. The dorsal and anal fin bases are of similar length. The palatine teeth are weak with up to four on the head of each bone. The photophores are IP 10–11, PV 15–17, VAV 19–22, OV 14–16, VAL 19–23, and AC 10–13. The IP and PV rows are continuous, as is the AC. The postocular light organ is very close to and smaller than the eye. There is a ring of luminous spots in the cheek region. The barbel tapers slightly to its tip, which has no bulb. Older fish have four small white spots between the postocular light organ and the preopercle, and three to four white spots on the preopercle. The barbel is white. The species attains 15.9 cm in standard length. habitat:  This is a mesopelagic species found down to about 1,000

Distribution of Astronesthes cf. richardsoni

sources:  Gibbs (1964); Parin & Borodulina (2003).

m. A southern Davis Strait capture was found at 577 m.

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biology:  Its food is crustaceans and mesopelagic fishes.

Borostomias antarcticus (Lönnberg, 1905)

importance:  It is not economically important.

Large-eye Snaggletooth, dragon-saumon à grands yeux

common names: A local name is Antarktisk Ulvekæft (Danish/

Greenlandic).

distribution:  It is found in Davis Strait off southern Baffin Island, in southwest and southeast Greenland, along the Atlantic coast of Canada, and in all major oceans.

taxonomy:  The genus comes from the Greek boros (gluttonous) and the Latin stomias (mouthy). The species name comes from the Latin antarcticus (southern). description:  This species is distinguished by the dorsal fin being at mid-body; a dorsal adipose fin being present; and the lower caudal fin-rays not being elongated. The teeth on the maxilla bone of the upper jaw are clearly separated and do not slant rearward. There is no adipose fin ventrally, and there are fang-like teeth in the jaws. There is a double light organ behind the eye, the anterior part being small. Lateral photophores (OA) number 40–47, and ventral row photophores (IC) 64–69. Photophores number IP 9–11, PV 20–26, VAV 19–25, OV 22–25, VAL 19–24, and AC 10–13. Dorsal fin-rays number 9–13, anal finrays 12–17, pectoral fin-rays 7–9, and pelvic fin-rays 7. Gill rakers number 12–16 on the lower arch. The head and body are black, but the skin often becomes detached after the fish has been hauled up from great depths. The single chin barbel has a bulb with one to two filaments and is pale distally. The species reaches more than 35.0 cm in standard length. Distribution of Borostomias antarcticus

habitat:  This species is bathydemersal and oceanodromous. It is

reported from Davis Strait at 900–1,402 m, and in Canadian waters off Baffin Island at 477–1,461 m and 2.9°C–3.8°C. Elsewhere it is oceanic and mesopelagic to bathypelagic at 300–2,630 m.

sources:  Karrer (1973); Sardou (1980).

Borostomias antarcticus



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Chauliodus sloani

Bloch and Schneider, 1801 Manylight Viperfish, chauliode très-lumineux

common names: A local name is Slones Segltandfisk (Danish/ Greenlandic). Other common names are Sloan’s Fangfish, Sloane’s Viperfish, chauliode de Sloane, and grandcroc petit fil. taxonomy:  The genus comes from the Greek chaulios (exserted,

or mouth opened) and odous (tooth). The species is named after Sir Hans Sloane (1660–1753), a naturalist whose donation of collections founded the British Museum. The species may also appear incorrectly as sloanei.

description:  This viperfish is distinguished by the postorbital photophore (light organ behind the eye) being rounded posteriorly, and the dorsal fin origin being over photophores four to eight in the lateral series (over the pectoral fin level). The lateral photophore row is ventral to the fifth “scale” row. The third “scale” row has only two photophores. The photophores emit a bluish light in life. They are IC 62–72, OA 41–49, IP 8–11, PV 17–23, VAV 23–30, AC 9–13, OV 17–22, and VAL 23–29. The dorsal fin-rays number 5–8, usually 6–7 with the first one elongate; anal fin-rays 10–13, usually 11–12; pectoral fin-rays 9–14, usually 13; and pelvic fin-rays 6–8, usually 7. The dorsal adipose and anal fins are far posterior on the body. The largest lower jaw fang protrudes over the top of the eye when the mouth is closed. A slender, flexible chin barbel is present in fish of up to about 100 mm standard length and is lost during the process of transformation from larva to adult. The

body is covered in a clear gelatinous coat. The colour is an iridescent silver blue on a black to dark-brown background and is greenish, black, or dark brown above. The fins are lighter than the body, and the membranes are mostly transparent. This species reaches 35.0 cm in standard length.

habitat:  This species is mesopelagic and bathypelagic. In the North Atlantic it is fairly common and has been caught at less than 500 m down to 2,800 m during the day, ascending to the upper 800 m at night – as shallow as 45 m. Its maximum depth is 4,700 m. In the Davis Strait and southern Baffin Bay it has been caught at 680– 1,451 m and at 3.1°C–4.2°C, and as shallow as 276 m in Hudson Strait.

biology:  Its food is small fishes, particularly Lanternfishes, and

crustaceans, with some algae. It hovers head up, fins extended, with the tip of the elongate second dorsal fin-ray dangling in front of the head to attract prey. Atlantic Cod eat this viperfish. The photophores have been studied in the laboratory and found to flash or emit continuous luminescence when stimulated electrically. They serve to obliterate the shadows cast by daylight from above and the fish’s body. It spawns year round, with a peak in late winter to early spring. Larvae are about 7 mm long at hatching, grow to about 44 mm, and then shrink to 27 mm before resuming growth to transform into adults.

importance:  It is not economically important. distribution:  It is found in Baffin Bay, Davis Strait, and eastern Hudson Strait. Worldwide it is found in tropical to temperate seas including southwest and southeast Greenland and the Atlantic coast of Canada.

Chauliodus sloani

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Malacosteus niger Ayres, 1848

Stoplight Loosejaw, drague rouge-verte

common names: A local name is Sort Smalkæbefisk (Danish/ Greenlandic). Other common names are Black Loosejaw, Lightless Loosejaw, Northern Stoplight Loosejaw, drague à godet, and drague sans lampe.

taxonomy:  The genus comes from the Greek malakos (soft) and osteon (bone). The species name comes from the Latin niger (black).

Distribution of Chauliodus sloani

sources:  Tchernavin (1953); Karrer (1973); Parin & Novikova (1974); Van Utrecht, Van Utrecht-Cock, & de Graaf (1987); Hudon (1990a); Jørgensen et al. (2005).

description:  There is no floor to the mouth, an obvious and unique character. The absence of a barbel, the presence of poorly developed flank light organs, and only one pair of nostrils also distinguish this species from other loosejaws in Canada. Dorsal fin-rays number 14–21, anal fin-rays 17–24, pectoral finrays 2–6, and pelvic fin-rays 6–7. The dorsal and anal fins are near the caudal fin. The photophores are OA 6–15 and IC 12–22, often not countable. There is a large, tear-drop-shaped, dark-red light organ under the eye and a smaller bright green one behind it. The diameter of this latter light organ is more than 25% of the size of the eye. Other light organs are white or violet. The head and body are black. This species attains 25.6 cm in total length but can be as large as 30.0 cm.

Malacosteus niger



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habitat:  This species is not uncommon compared with other loosejaws and is caught generally at 915–1,830 m depth, rarely at the surface, and as deep as 3,886 m. It is a mesopelagic and bathypelagic species. Canadian Arctic records are from Davis Strait at 1,019–1,463 m and a bottom temperature of 3.1°C–3.6°C, and from 943.5–1,451 m and 3.4°C–3.6ºC. The shallowest record in Canadian waters is 880 m. biology:  Its food is crustaceans and fishes. The red fluores-

cent light emitted from the organ under the eye in ultraviolet light is rare among sea creatures, blue or green being more common. The red-shifted light is undetectable by other mesopelagic fishes. Other members of this species use a bacteriochlorophyll-based photosensitizer to perceive the red light, sequestering the bacteriochlorophyll from copepods for long-wave sensitivity. This fish may use red-shifted bioluminescence from the AO photophores to search for prey such as these copepods, sustaining itself on these crustaceans between rarer encounters with larger myctophid and gonostomatid fish prey.

importance:  It is not economically important. distribution:  The species is found in Davis Strait and eastern Hudson Strait, southwest and southeast Greenland, and all tropical and subarctic oceans.

Rhadinesthes decimus (Zugmayer, 1911)

Slender Snaggletooth, dragon-saumon élancé

common names:  A local name is Slank Ulvekæft (Danish/

Greenlandic). The English common name is taken from the Danish name.

taxonomy:  The genus comes from the Greek rhadinos (slender)

and esthes (garment). The species name comes from the Latin decimus (tenth).

description:  This species is distinguished by a line of luminous tissue originating on the head behind the postorbital organ and extending onto the flank; elongated lower caudal fin-rays; and no distinctive dorsal adipose fin. The postorbital organ is smaller than the eye. The bulb of the barbel has one to four filaments. The gill teeth are small, equal to or shorter than the gill-bar width. Maxillary teeth are short, not slanting rearward, and widely spaced. There are no fangs in the jaws. Dorsal fin-rays number 11–14, anal fin-rays 17–21, and pectoral fin-rays 6–8. Photophores are IP 6–10, PV 25–26, VAV 20–23, OV 22–24, VAL 20–27, IC 66–74, OA 44–49, and AC 15–16. The head and body are black. This species reaches 41.0 cm in standard length. habitat:  A single specimen was caught in Davis Strait at an

unknown depth over a bottom temperature of 4°C. This species is oceanic and meso-bathypelagic in habitat, usually below 500 m during the day.

biology:  Its food is mid-water fishes and crustaceans. importance:  It is not economically important. distribution:  This species is found in Davis Strait off southern Baffin Island, from a single record at 63º06' N, 59º47' W (ARC 8704111); southeast Greenland; and in the North Atlantic, Indian, and Pacific Oceans.

Distribution of Malacosteus niger

source:  Kenaley (2007).

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Rhadinesthes decimus

Stomias boa (Risso, 1810)

Boa Dragonfish, dragon-boa

common names:  A local name is Boafisk (Danish/Green-

landic). Other common names are Boa Scaly Dragonfish, Scaly Dragonfish, poisson-drague commun, and stomie commune.

taxonomy:  The genus comes from the Latin stomias (mouthy).

The species name comes from the Latin boa (a large water snake). The subspecies Stomias boa ferox Reinhardt, 1842, described from Greenland, is the one present in Canadian waters. Some authors recognize this subspecies as a distinct species.

Distribution of Rhadinesthes decimus

sources:  See the family sources and the bibliography.



UTP Fishes Book 5pp04.indb 313

description:  This species is distinguished by having six rows of hexagonal areas above the belly’s light-organ rows; dorsal and anal fins near the caudal fin; and two pairs of nostrils. The barbel is about as long as the head and bears two to four short dark filaments on the bulb at its tip. There are two to three palatine teeth on each side and two small teeth on each side of the lower jaw symphysis. Dorsal fin-rays number 17–22, anal fin-rays 18–23, pectoral fin-rays 6 (rarely 7), and pelvic fin-rays 5. There are 82–91, usually 87–89, light organs in the ventral series, more than in the nominate subspecies (Stomias boa boa). Its colour is iridescent silver on the sides and black to dark brown underneath and on the belly and back. The hexagonal areas form a distinct pattern. It attains 32.2 cm in standard length.

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Stomias boa

Stomias boa, x-ray

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habitat:  It is a bathypelagic species that descends to about 1,500

m; most are caught above 300 m, and it may migrate near the surface at night. It has been caught at 489–1,395 m in Davis Strait at 1.2°C–3.6°C, and at 556.5–1,451.0 m and 0.1°C–4.6ºC in southern Baffin Bay and Davis Strait generally.

biology:  This species feeds on fish and crustaceans, and a vertical

migration occurs. It positions the body horizontally with pectoral and pelvic fins spread out and the barbel pointing forward. This posture, coupled with the dorsal and anal fin position at the rear of the body, suggests an ambush predator. This species may be eaten by Atlantic Cod in Ungava Bay. Spawning occurs in the spring. Larvae grow to about 44 mm, shrink to about 23 mm, and then resume growing to transform into adults.

Family Notosudidae Waryfishes, Guetteurs

Brian W. Coad

importance:  It is not economically important. distribution:  The species is found in Baffin Bay, Davis Strait, Frobisher Bay, Hudson Strait, and Ungava Bay. It is also found in the North Atlantic Ocean including northwest, southwest, and southeast Greenland, in the Atlantic waters off Canada, and in the Indian and Pacific Oceans.

The Family Notosudidae is widely found, though not common, from sub-Arctic to sub-Antarctic waters. There are 19 species, including 6 reported from Canada, 1 of which is from Arctic waters. The maximum size is about 50 cm. Waryfishes or Paperbone Fishes are found from surface layers to the bottom in oceanic waters. Their bones are thin and delicate, as their other common name suggests. Their snout is spatulate, and they have large oval eyes with an elliptical pupil. Waryfishes lack a gas bladder and light organs. The scales are large, cycloid, easily detached, and number 44–65 in an obvious lateral line. An adipose fin is present over the anal fin. Anal fin-rays number 15–21, and the dorsal fin is at the middle of the body with 9–14 rays. Pectoral finrays number 10–15, and pelvic fin-rays 8–10. They are unique among their relatives in that their larvae have teeth on the maxillary bone. The fish lose their teeth and gill rakers at maturity. They are very active and dart from a hovering position to catch such food items as zooplankton, crustaceans, and small fishes. They can easily dodge trawls. Waryfishes migrate from the bottom in the day to the mesopelagic zone at night. They are only likely to be seen by scientists. Young Waryfishes live higher in the water column than do adults, but even adults may be in the upper 200 m. However, some descend below 1,000 m. Waryfishes are hermaphrodites. Some species are known to migrate long distances over the ocean in search of food.

sources:  Marshall (1966); Bertelsen, Krefft, & Marshall (1976).

Distribution of Stomias boa

sources:  Gibbs (1969); Karrer (1973); Fink & Fink (1986); Gillis & Allard (1986); Treble et al. (2000); Jørgensen et al. (2005).



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Scopelosaurus lepidus (Krefft and Maul, 1955)

Blackfin Waryfish, guetteur à nageoire noire

biology:  Its food is crustaceans, such as euphausiids and

hyperiids, and small fishes fixated by the large eyes. Spawning occurs in mid-water far from shore, and there is known to be a concentration in the Sargasso Sea. This waryfish is eaten by Atlantic Cod and other large fishes.

common names: A local name is Nordlig Øglesmelt (Danish/

importance:  It is not economically important.

taxonomy:  The genus comes from the Greek skopelos (said to be

distribution:  The species is found in Davis Strait and on both sides of the Atlantic Ocean, including southwest and southeast Greenland, south to the Grand Banks, and off Nova Scotia.

Greenlandic).

an ancient name for a fish, used as a Lanternfish genus by Baron Georges Cuvier in 1816, now a synonym of Myctophum; note that the Greek word means “rock” and “peak,” among others) and sauros (lizard). The species name comes from the Greek lepidos (scale or scaly) or the Latin lepidus (pretty) (literature sources differ). This species may be a synonym of Scopelosaurus harryi (Mead, in Mead and Taylor, 1953).

description:  This species is distinguished by family characters

and the counts of fin-rays, scales, gill rakers, and pyloric caeca. There are 55–64 lateral-line scales. The teeth are small and pointed. The dorsal fin-rays number 10–12, anal fin-rays 16–19, long pectoral fins 10–15 rays, and pelvic fin-rays 9. The gill rakers on the lower arch are long, numbering 17–20. The gill openings are very wide and extend in front of the eye. The pyloric caeca are numerous, 16–39. The body is light to dark brown in colour with scale pockets outlined in black. The scales are usually lost. In larger specimens (>  12 cm) the pectoral fins have a large black patch at their base and a white band at the tip. The peritoneum is black. The species reaches 41.6 cm in length.

habitat:  Young Blackfin Waryfishes are found at 70–200 m, and

adults live deeper, at 500–995 m or more, where they are benthopelagic to mesopelagic. The species has been caught at 471.5–1,457.5 m and 2.2ºC–3.8ºC in Davis Strait. Other cruise data for Davis Strait indicate a depth range of 620–1,472 m. They migrate long distances to continental slopes to feed.

Distribution of Scopelosaurus lepidus

sources:  Karrer (1973); Munk (1977).

Scopelosaurus lepidus

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Anotopterus pharao

Family Paralepididae

Zugmayer, 1911

Daggertooth, pharaon

Barracudinas, Lussions common names: A local name is Dolktandfisk (Danish/Greenlandic). Other common names are Javelinfish and North Atlantic Daggertooth.

Brian W. Coad

taxonomy:  The genus comes from the Greek a (without), noton

(back), and pteron (fin). The species name comes from the Greek pharaoh, in allusion to the projecting lower-jaw tip that resembles the false beard seen on statues and paintings of Egyptian pharaohs. This species was formerly placed in its own family, Anotopteridae. Anotopterus arcticus Nybelin, 1946, was described from a head taken from the stomach of an Atlantic Halibut in Davis Strait and is a synonym. Barracudinas are found in all oceans including the Arctic and Antarctic Oceans. There are about 56 species, including 16 in Canada, of which 4 enter Arctic waters. These fishes are fairly rare. The maximum size is 146 cm although most species are a third of this or less. Barracudinas resemble the Barracuda Family members, hence their name, but they are not related. They have a fragile skeleton as an adaptation to mid-water life where food sources are sparse. The mouth is large and carries sharp fang-like teeth. On the lower jaw and the palatine bone in the upper mouth, fangs are alternately fixed and depressible. The eyes are rounded with round pupils. The dorsal fin is at the mid-body or absent, and there is an adipose fin. Some species have an adipose fin in front of the anal fin. The anus is usually near the pelvic fins, well separated from the anal fin. Dorsal fin-rays number 7–16, anal fin-rays 12–50, pectoral fin-rays 9–18, and pelvic fin-rays 8–13. The fins lack spines. Deciduous cycloid scales are present in some species, including those in Arctic Canada, numbering 51–92 or absent. The gill rakers are composed of multiple spines on bony plates. There is no gas bladder. There may be one to two luminous ducts in the belly muscles in some species. The overall colour is iridescent silver. These fishes are very numerous in mid-water, sometimes reaching the surface. They are thought to be very agile swimmers because few adults have been caught in nets. Observers in submersibles have seen some species swimming vertically with the head up at a 45° angle, using only the caudal region for propulsion. Others are reported to swim with the head down. This swimming is thought to aid in vision and lateral-line function by reducing body oscillations. Their food is fishes, crustaceans, and cephalopods. They are an important food for Salmons, tunas, Cods, sharks, albacores, whales, and seals. Some Barracudinas undergo a diurnal vertical migration, following their crustacean food. Some species have separate sexes, and others are simultaneous hermaphrodites. They are not commercially important.

sources:  Ege (1953); Rofen (1966a, 1966b).



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description:  The large jaws having about 11 strong, bladelike teeth on the palatine give this fish its English name and are distinctive. The absence of scales, of a dorsal fin, and of light organs, coupled with the presence of a large adipose fin, are also characteristic. There are two skin keels on each side of the caudal peduncle. The anal fin-rays number 12–17, usually 13–15; pectoral fin-rays 12–16, usually 14; and pelvic fin-rays 7–11. The bone skeleton of this fish is fragile despite its ferocious appearance, and the skin is easily ripped. The colour is dark grey to brownish yellow on the back, to yellowish white or silvery on the belly, with silvery sides and black tips of the jaws, the branchiostegal membranes, the tail fin, and the tips of the pectoral fins. The jaw and the gill and peritoneal cavities are black. The young are transparent. The species attains 146.0 cm in length and 1.65 kg in weight. habitat:  The Daggertooth can be found at the surface or at great

depths, to 5,100 m. The common depth range is 500–2,250 m in mesopelagic to bathypelagic waters. Younger fish are found in temperate waters such as those off the California coast, but the largest specimens come from cold waters in the Antarctic and the northern Atlantic and Pacific Oceans. It has been caught in Davis Strait at 180–350 m and 4°C–5°C. Elsewhere it has been recorded at 2°C–24°C.

biology:  Many specimens have been recovered from the stomachs of other fishes such as Greenland Halibut, Atlantic Cod, and even whales. Some have been caught by fishermen trolling for salmon. Daggertooths eat fishes, such as Sharpchin Barracudina in Davis Strait, Deepwater Redfish near Iceland, and commercial species like salmons. The Daggertooth may attack from a waiting position with the head up or down and causes incised or slash wounds on the salmon. The stomach is expandable, and the pectoral girdle is not attached to the skull, to allow large prey to be ingested. Smaller items like crustaceans, polychaetes, squids, coelenterates,

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Anotopterus pharao

and salps are also eaten. Daggertooths live at least six years, possibly 12–15 years. They are thought to be synchronous hermaphrodites with semelparous reproduction. When their gonads mature, the stomach and intestine atrophy and teeth are lost.

Arctozenus risso

importance:  It is not economically important.

common names:  A local name is Rissos Laksetobis (Danish/Greenlandic). Other common names are Ribbon Barracudina, Spotted Barracudina, barracudina longue quille, and barracudina pintade.

distribution:  This species is found in the North Atlantic Ocean

from Davis Strait and southwest and southeast Greenland to the Canary Islands. The single Canadian Arctic record is from Resolution Island in eastern Hudson Strait (Parsons, 1982).

(Bonaparte, 1840)

White Barracudina, lussion blanc

taxonomy:  The species is also spelt rissoi and is sometimes placed in the genus Notolepis. The genus comes from the Greek arktos (northern) and xenos (strange). The species is named after Guiseppe Antonio Risso, called Antoine Risso (1777–1845), author of Ichthyologie de Nice. Paralepis kroyeri Lütken, 1892, is regarded as a synonym or as a subspecies. The subspecies Arctozenus risso kroyeri has smaller teeth than the type subspecies (Arctozenus risso risso), and there are, on average, more vertebrae and anal fin-rays. description:  This species is distinguished by the pelvic fin origin being behind the dorsal fin origin and by having 28–35 anal fin-rays and 9–13 pectoral fin-rays. Dorsal fin-rays number 8–13, and pelvic fin-rays 9. Lateral-line scales number 59–70, are enlarged, embedded, and with pores, and are easily lost on capture. Gill rakers number 36–45 and may be obsolescent in large fish. Vertebrae number 75–85. The large eyes have a pupil angled forward to assist in detecting prey as the body hovers in the water column. The anus lies between the pelvic fins. The overall colour is iridescent silver with a dark upper-flank stripe or speckles. The jaw tips and nostrils are dark. There is a black spot or area at the anal and pelvic fin origins. The peritoneum is black. The species reaches 31.0 cm in standard length.

Distribution of Anotopterus pharao

sources:  Nybelin (1946); Hubbs, Mead, & Wilimovsky (1953);

Postolakii (1962); Rofen (1966b); Templeman (1970c); Parsons (1982); Hudon (1990a); Kukuev (1998).

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habitat:  This species is mesopelagic to bathypelagic and is found

mostly between 200 and 1,000 m, but as shallow as 64 m and as deep as 2,200 m. It occurs in small schools or individually, with the young being in shallower water than adults. In Hudson Strait it has been caught at 260–360 m and in Davis Strait at 433–1,247 m at 0.7°C–5.6°C. It is one of the better known Barracudinas because it enters shallow waters in the north, sometimes dying from cold and being cast up on shore.

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Arctozenus risso

biology:  Its food is small fishes, squids, and crustaceans such as

shrimps, caught pelagically. It is an important food species for a variety of fishes including Atlantic Cod, recorded from cod in Forbes Sound and at Killiniq (Port Burwell), Ungava Bay; Greenland Halibut in Davis Strait; and Ogac, Pollock, Swordfish, redfishes, and seals generally. Peak spawning is thought to occur in May south of the Arctic, but larval stages indicate that January–September is the season. Adults are found much further north than are post-larvae. Larvae are planktonic.

importance:  It has no importance in Arctic Canada, but it is very common in Greenland and locally important.

distribution:  The subspecies Arctozenus risso kroyeri is found only in the North Atlantic Ocean and adjacent Arctic waters. The type subspecies, Arctozenus risso risso, is found worldwide. In Canada the species has been reported from Davis Strait and eastern Hudson Strait, and southwest and southeast Greenland, southward through the Maritimes to Georgia.

Distribution of Arctozenus risso

sources:  Hildebrand (1948); Dunbar & Hildebrand (1952);

Rofen (1966a); Chumakov & Podrazhanskaya (1986); Janssen, Pankhurst, & Harbison (1992); Il’inskiy, Balanov, & Ivanov (1995); Jørgensen et al. (2005).



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Magnisudis atlantica (Krøyer, 1868)

Duckbill Barracudina, lussion à bec de canard.

common names:  A local name is Kort Laksetobis (Danish/ Greenlandic). Another common name is barracudine.

importance:  It is not economically important. distribution:  The species is found in tropical to Arctic waters

of the Atlantic, Pacific, and Indian Oceans and off both the Atlantic and the Pacific coasts of Canada, off southwest and southeast Greenland, and into Arctic waters in the Davis Strait without an exact locality (Berth, Schultz, & Vaske, 1979).

taxonomy:  The genus comes from the Latin magni (large) and sudis (stake, an old name for barracuda in reference to the shape). The species name comes from the Latin atlanticus (of the Atlantic). It is sometimes placed in the genus Paralepis. description:  This species is distinguished by having the pelvic fin origin below the dorsal fin; pectoral fin-rays numbering 15–18; anal fin-rays numbering 20–26; the rear of the premaxilla being clearly behind the nostril; and the longest gill raker being much less than twice as long as the next adjacent raker. The lower jaw has smooth-edged teeth. Dorsal fin-rays number 9–11, and pelvic fin-rays 9. The pectoral fins are shorter than the anal fin base. Lateral-line scales number 55–64, but the scales are easily lost. The lateral-line scales lack pores. The gill rakers number 31–40. There are three or more rakers on each base. Vertebrae number 63–66 in the North Atlantic. The anus is level with the pelvic fin tips. The colour is brownish above with silvery flanks and dark spots. The fins and fin bases may be darkened, but there is no black area at the base of the anterior anal fin. The gill cavity and peritoneum are black. The species reaches 50.0 cm in length.

?

Distribution of Magnisudis atlantica

habitat:  This species is pelagic-oceanic and oceanodromous. It is

usually found in deep mid-waters but may be found in surface waters too. It can be cast ashore in temperate to polar areas. It has been recorded down to 4,750 m.

sources:  Berth et al. (1979); Balanov, Il’inskiy, & Ivanov (1995); Il’inskiy et al. (1995).

biology:  The food items of this species include shrimps, squids, and mesopelagic fishes. It is eaten by sharks, tunas, and whales. Spawning occurs year round in tropical waters.

Magnisudis atlantica

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biology:  It is apparently quite common and an important food

Paralepis coregonoides

and Nordisk Laksetobis (Danish/Greenlandic). Another common name is Lancetfish.

for Atlantic Salmon, as well as cod, tuna, lancetfish, and seals. Atlantic Salmon caught in surface waters (about 3 m) of Davis Strait were found to have this species in their stomachs. Cod and Greenland Halibut feed on this species in Ungava Bay. Barracudina found in the stomach of cod, Greenland Halibut, and Atlantic Salmon point head forward, suggesting that they were captured tail first by pursuit. Their food is shrimps, fishes, and some plankton. Reproduction occurs from March to September in temperate waters.

taxonomy:  The genus comes from the Greek para (near) and

importance:  It is not economically important.

Risso, 1820

Sharpchin Barracudina, lussion à menton

common names: Local names are Putorutorsôak (Greenlandic)

lepis (scale). The species name comes from the Greek core (pupil of the eye) and gonia (angle) – the scientific name for whitefishes (Coregonus spp.) – and from eidos (likeness, form). The subspecies Paralepis coregonoides borealis Reinhardt, 1837, is recorded in Canada and was originally described from Greenland. Paralepis coregonoides coregonoides Risso, 1820, was described from France. Subspecies are distinguished by the vertebrae and myomere counts and by post-larvae characters, but the adults are externally the same. Greenland records may also include another subspecies, Paralepis coregonoides barracudina Fowler and Phillips, 1910, originally described from New Jersey, but whether this is valid and found in Canadian waters requires further study.

distribution:  The species is found in Davis Strait, northwest, southwest, and southeast Greenland, and the Atlantic Ocean south to the Grand Banks and off Georges Bank.

description:  This species is distinguished by the pelvic fin origin being below the dorsal fin origin; by having 22–24 anal fin-rays; by the rear of the premaxilla being under the nostril; and by the longest gill raker being more than twice as long as the next adjacent raker. The dorsal fin-rays number 9–11, pectoral fin-rays 14–17, and pelvic fin-rays 9. The lateral-line scales number 57–62. The gill rakers number 3–6 on each base but are lost in older fish. The vertebrae number 67–74. The overall colour is iridescent silver, overlain by black to brown in preserved fish, becoming lighter ventrally. The peritoneum is black. The species reaches over 50.0 cm in standard length. habitat:  The larvae hatch at 70–140 m, young fish are found from

the surface to 200 m (epipelagic to mesopelagic), and older fish are deeper (mesopelagic to bathypelagic) and seldom caught. It has been caught at 290–640 m and 1.5°C–3.0°C in Davis Strait, and elsewhere down to 1,032 m. Some specimens have been found stranded on beaches, apparently being adults that have drifted too far north and frozen to death.

Distribution of Paralepis coregonoides

sources:  Templeman & Squires (1960); Lear & May (1971); Kar-

rer (1973).

Paralepis coregonoides



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Family Myctophidae Lanternfishes, Poissons-lanternes

Brian W. Coad

Lanternfishes are found in all seas, from the Arctic through the tropics to the Antarctic, with 240 species worldwide. There are 83 species in Canadian waters, of which 8 are reported from the Arctic. These fishes are abundant. The maximum size is about 30 cm although most species are much smaller, about half this size. Lanternfishes have large eyes and a blunt snout; possess between 50 and 80 light organs (hence “Lanternfish”); have an adipose fin, a dorsal fin on the middle of the back, fins usually lacking strong spines except for a rudimentary spine at the anterior base of the dorsal, anal, pectoral, and pelvic fins; and have abdominal pelvic fins. The scales are easily detached and are usually lost when fish are brought up from deep water. Instead of lateral-line scales, the horseshoe-like organs left on the flank by detached scales are counted. The scales are usually cycloid. These fish have dark brown, black, or bluish black backs and silvery scales along the flanks.

Photophores are used to identify the species. They are individually and collectively named with abbreviations, and an explanation of these abbreviations may be found in the “Keys” section and the glossary. These deep-sea fishes are often damaged on capture and the photophores lost; therefore, specimens may be easily misidentified. Photophores produce a bluish light by a reaction in which the chemical luciferin is oxidized by the enzyme luciferase in the presence of oxygen. Each photophore has its own blood and nerve supply and consists of a scale embedded in the skin and modified into a shallow cup to support the tissues. The tissues include a reflector layer backed by dark pigment and, within the cup, the luminous tissue. The scale cup has a modified scale over it, forming a lens. The photophores, in their species-specific patterns, are believed to be important to the Lanternfishes in recognizing members of their own species. They may also block out the Lanternfish’s silhouette to predators from below by emitting a continuous, low-intensity light that matches the light coming from the surface of the sea. Some authors maintain that photophores can be switched on or off so that an illuminated fish can suddenly “vanish” in the dark ocean depths and so that a rhythmical pattern can be emitted to help in species recognition. Certainly the intensity of the light emitted can be varied, and in Benthosema glaciale the photophores emit repetitive flashes for a few seconds at 10–15 second intervals. A second common type of light organ in Lanternfishes is the supracaudal and infracaudal luminous glands found on the upper and lower caudal peduncle at the base of the tail. These glands have been called “stern chasers” and are believed to function to deflect predators. The blue flash of these organs has been likened to a spark of electricity. The flash attracts the predator to the tail of the fish, but by the time the predator strikes in that region the light has gone out and the Lanternfish has departed into the dark ocean with a flick of its tail. The flash of light may also serve to blind the predator. A predator attacking a school of Lanternfishes would be even more confused.

Abbreviations for photophores

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These fishes are very common offshore at depths of about 300– 1,200 m, although many reach the surface at night in a vertical migration of several hundred meters to feed on zooplankton. They can be attracted to lights and are then easy to catch. Young Lanternfish are found higher in the water column than adults, and larvae of most species are in the upper 125 m. Larval Lanternfish differ from adults in pigmentation and light-organ arrangement. Larvae may be stylophthalmoid, in which the eyes are on long stalks. Lanternfishes feed on such plankton as copepods, ostracods, krill, arrow worms, and fish eggs and larvae. Lanternfishes are not caught by anglers, and only a few are commercially important. Their potential importance is high because of their numbers and their possible use in fish oil and fish-meal production. They are, however, a numerous and important food for many other species of fish such as cods, Swordfish, tunas, and salmons. As well, these fishes are eaten by seals, whales, and, rarely, Thick-billed Murre chicks (0.07% at Akapatok Island, for example). Lanternfishes have been studied by biologists for this reason.

sources:  Paxton (1972); Nafpaktitis et al. (1977); Bekker (1983).

Benthosema glaciale (Reinhardt, 1837)

Glacier Lanternfish, lanterne glaciaire

common names: Local names are Mikiapic Kapisilik; Kapisalik, Kapisalingoak, and Keblernak (Greenlandic); and Isprikfisk (Danish/Greenlandic). Other common names are Glacial Lanternfish and Large-eyed Frogfish. taxonomy:  The genus comes from the Greek benthos (the ocean depths) and sema (a constellation of stars, a flag, or a signal). The species comes from the Latin glacialis (icy) and is often spelled glacialis in the literature. description:  This species is distinguished by a combination of characters including photophore patterns and fin-ray counts. The AO row is subdivided into an anterior (AOa) and a posterior group; the PLO is more than its diameter above the level of the upper base of the pectoral fin; the second PVO is at or below the upper base of the pectoral fin; there are two Prc, the second being much higher than the first; the Dn is minute; the PVO row is horizontal or almost horizontal; the PVO1 is not more than its diameter below the PVO2; the Vn is small and obscure; the VO2 is elevated; anal fin-rays number 17–19; and dorsal fin-rays number 12–15.

Benthosema glaciale



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Pectoral fin-rays number 10–13. Total gill rakers are 15–20, visible in the open mouth. The AO row numbers 11–14 photophores in total. Lateral-line organs number 36–37. There is one Pol photophore. The supracaudal luminous gland is developed best in males, and the infracaudal in females. The glands begin to develop at a length of 2.5 cm. The species reaches 10.3 cm in standard length.

habitat:  This is probably the most abundant Lanternfish off west

Greenland and in the North Atlantic Ocean. It is generally found near surface waters at night and down to 1,407 m during the day. The temperature range is −0.08°C to 21°C, but this species is most abundant at 4°C–16°C. In Davis Strait and southern Baffin Bay, temperatures for 258 fish were −0.08°C to 5.6°C at 336–1,456 m. In the Canadian Davis Strait off Baffin Island in 29 collections, this fish has been caught at 420–1,407 m and at 0.8°C–3.8°C. Its absence from northern Baffin Bay is due to the cold temperatures and long periods of daylight. In Davis Strait about 46% of the population feeds at night in the upper 60 m, having migrated from depths of 300– 900 m. The remaining fish are found at 150–550 m. At night all fish caught are in water of at least 2°C. Older fish are found deeper than young fish at night. In NAFO Subarea 0B, catches decreased in proportion to the increasing depth. Distribution of Benthosema glaciale

biology:  The prey items are selected by the Lanternfish and are

oil rich. Its food is principally calanoid copepods and euphausiid crustaceans. Other foods are ostracods, mysids, amphipods, arrow worms, and the urochordate Oikopleura. The Glacier Lanternfish is eaten by many other fishes including Atlantic Cod and Greenland Halibut in Ungava Bay, and by Ivory Gulls when concentrated at night in the southwest Davis Strait. Northern Fulmars and Black-legged Kittiwakes eat this species in southern Davis Strait. Thick-billed Murres eat this species at Akpatok Island in Ungava Bay and Hantzsch Island off the southern tip of Baffin Island. The species is mature at age two years (about 3 cm long), and lives up to six years off Nova Scotia and Newfoundland – to eight years elsewhere. Spawning occurs intermittently in early autumn and winter, such that larvae appear in April and May when zooplankton is abundant. Up to 2,000 eggs are produced. Full photophore development is achieved at 14–15 mm in length.

sources:  Hildebrand (1948); Dunbar & Hildebrand (1952); Bolin (1959); Halliday (1970); Gjösæter (1973a, 1973b); MacLaren Marex (1979a); Orr & Parson (1982); Gillis & Allard (1984); Gaston (1985); Albikovskaya (1988); Sameoto (1989); Treble et al. (2000); Chambers & Dick (2007).

importance:  It is not economically important.

Greenlandic). Another common name is lanterne crocodile.

distribution:  This species is found in Baffin Bay, Davis Strait,

Hudson Strait, and Ungava Bay. It is also found in the Mediterranean Sea and in the North Atlantic Ocean including northwest, southwest, and southeast Greenland, and southward in Atlantic Canada to off Cape Hatteras. It is possibly throughout the Arctic Ocean where the waters are deep enough. A single specimen is known from off Point Barrow, Alaska, and it has been reported recently from the Canadian Beaufort Sea.

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Lampanyctus crocodilus (Risso, 1810)

Jewel Lanternfish, lanterne-joyau

common names: A local name is Krokodille-prikfisk (Danish/ taxonomy:  The genus comes from the Greek lampe or lampas

(shine, lamp, or torch) and nyx or nykte (night). The species name comes from the Latin crocodilus (crocodile). The correct name for Atlantic Ocean specimens may be L. gemmifer (Goode and Bean, 1879), but this requires more study to verify.

description:  This species is distinguished by a combination of characters including photophore patterns and fin-ray and gill-raker counts. The AO row is subdivided into an anterior (5–8 photophores) and a posterior group; the PLO is more than its diameter above the level of the upper base of the pectoral fin; the second PVO is at or below the upper base of the pectoral fin; there are 4 Prc; the

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Dn is absent; there are 5–8, usually 5–7, AOa photophores; the eyes are not semi-telescopic; the dorsal fin-rays number 13–15; the pectoral fins are short, not reaching the anal fin origin; and gill rakers number 14–18. The anal fin-rays number 16–19, and the pectoral fin-rays 13–16. The first PO photophore and two PVO photophores are not on a straight line. The fourth PO photophore is highly elevated; there are no luminous, scale-like structures on the belly between the pelvic fin bases or these bases and the anus; there are four VO photophores; and the SAO series is strongly angled. There are two to three cheek photophores, luminous tissue at the adipose fin base, and 38–39 lateral-line organs. The AO photophores total 13–16. Both males and females have caudal luminous glands composed of overlapping, scale-like structures without a black pigment border. The supracaudal luminous gland consists of 3–4 poorly defined segments, and the infracaudal gland is about three times as long and has 8–10 scale-like segments. A very large Lanternfish, it reaches 30.0 cm in standard length.

importance:  It is not economically important. distribution:  This species was found in Ungava Bay at Killiniq

(Port Burwell) in Atlantic Cod stomachs (Dunbar & Hildebrand, 1952). The material was damaged, and the identification may be incorrect. However, it is also recorded from southwest and southeast Greenland, southward in Atlantic Canada, and generally in the North Atlantic Ocean and the Mediterranean Sea.

habitat:  This species is epipelagic to bathypelagic. Its depth

distribution by day is usually 275–1,000 m, with a maximum abundance at 600–800 m. At night it is found near the surface to 1,000 m, and maximum abundance varies with locality. Near Bermuda the night maxima were at 100–250 m and at 400–1,000 m. The maximum depth attained exceeds 1,200 m. Only juveniles migrate. In Ungava Bay it has been reported at 46 m.

biology:  Distribution is temperate and partially subtropical, and

it is the commonest Lanternfish in the temperate Atlantic Ocean. Some specimens enter sub-Arctic waters. It has been reported from Atlantic Cod stomachs in Ungava Bay. Its food is a wide variety of pelagic and benthopelagic invertebrates and even small fishes. Spawning occurs in the autumn in the Atlantic Ocean.

Distribution of Lampanyctus crocodilus

source:  Dunbar & Hildebrand (1952).

Lampanyctus crocodilus



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Lampanyctus intricarius Tåning, 1928

Diamondcheek Lanternfish, lanterne à joue pailletée

common names: A local name is Bredhalet Prikfisk (Danish/ Greenlandic). Another common name is Intricate Lanternfish.

taxonomy:  The species name comes from the Latin intricatus

(intricate) and -arius (forming an adjective to the noun).

description:  This species is distinguished by a combination of characters including photophore patterns and fin-ray and gill-raker counts. The AO row is subdivided into an anterior (8–9 photophores) and a posterior group; the PLO is more than its diameter above the level of the upper base of the pectoral fin; the second PVO is at or below the upper base of the pectoral fin; there are four Prc; the Dn is absent; there are 8–10 AOa photophores; the eyes are not semi-telescopic; the dorsal fin-rays number 14–16; the pectoral fins are long, extending beyond the anal fin origin; and the gill rakers number 15–18. The anal fin-rays number 17–20, and the pectoral fin-rays 13–15. The first PO photophore and the two PVO photophores are not on a straight line; the fourth PO photophore is highly elevated; there are no luminous, scale-like structures on the belly between the pelvic fin bases or these bases and the anus; there are four VO photophores; and the SAO series is strongly angled. There are one to two (rarely three) cheek photophores, luminous tissue at the adipose fin base, and 39–40 lateral-line organs. The AO photophores total 16–18. Both males and females have caudal luminous glands composed of overlapping, scale-like structures without a black pigment border. The supracaudal luminous gland has 2–3 scale-like segments, and the infracaudal 8–10. The species reaches 20.0 cm in standard length. habitat: This is a temperate or subpolar-temperate species in

both hemispheres, and it is epipelagic, mesopelagic, and bathypelagic. Depth distribution by day is generally 550–750 m, and by night 40–550 m, with a maximum abundance mostly of young at 75  m.

A single Canadian Arctic record was taken in Davis Strait over a bottom depth of 1,092 m and a bottom temperature of 4°C. The specimen was damaged, and identification was tentative.

biology:  Similarly to related species, its food is zooplankton. importance:  It is not economically important. distribution:  The species has been found in southern Davis Strait as a single record at 61.6167° N, 62.0167° W (ARC 8704979), in southwest and southeast Greenland, off the Atlantic coast of Canada, and in the Atlantic, Indian, and Pacific Oceans.

Distribution of Lampanyctus intricarius

sources:  See the family sources and the bibliography.

Lampanyctus intricarius

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from Davis and Hudson Straits is 245–968 m. In NAFO Subarea 0B, catches decreased in proportion to increasing depth.

Lampanyctus macdonaldi (Goode and Bean, 1896)

Rakery Lanternfish, lanterne-bouée râtelière

biology:  This species is epipelagic, mesopelagic, and bathypelagic. Its food is probably crustaceans.

common names: A local name is Macdonalds Prikfisk (Danish/ Greenlandic). Another common name is Rakery Beaconlamp.

importance:  It is not economically important.

taxonomy:  The species is named after Marshall McDonald

(1835–95), a U.S. Fish Commissioner, known for the development of hatchery apparatus and fish ladders.

description:  This species is distinguished by a combination of characters including photophore patterns and fin-ray and gill-raker counts. The AO row is subdivided into an anterior (6–8 AOa photophores) and a posterior group; the PLO is more than its diameter above the level of the upper base of the pectoral fin; the second PVO is at or below the upper base of the pectoral fin; there are four Prc; the Dn is absent; the eyes are not semi-telescopic; the dorsal finrays number 13–16; the pectoral fins are delicate and short, hardly extending to the pelvic fin origin; and the gill rakers number 21–27. The anal fin-rays number 15–19, and the pectoral fin-rays 12–14. The first PO photophore and two PVO photophores are not on a straight line; the fourth PO photophore is highly elevated; there are no luminous, scale-like structures on the belly between the pelvic fin bases or these bases and the anus; there are four VO photophores; and the SAO series is strongly angled. The AO photophores total 13–15, and the lateral-line organs 36–37. Both males and females have caudal luminous glands composed of overlapping, scale-like structures without a black pigment border. The supracaudal luminous gland has 3–4 poorly defined scale-like structures, and the infracaudal gland has 7–8 poorly defined scale-like structures, in both males and females. The species reaches about 20.0 cm in total length. habitat:  This species is subpolar-temperate in both hemispheres.

Its general depth distribution is 550 m to 1,000 m or more by day. At night the young are at 60–175 m, and adults are deeper than 850 m. It has been caught in Davis Strait at 423–1,187 m at 0.5°C–3.6°C and at 429.5–1,457.5 m at 0.5°C–4.3°C. Its depth range for cruise data

distribution:  The species is found in Davis Strait, Hudson Strait, Ungava Bay, southwest and southeast Greenland, in the Atlantic Ocean, and circumglobally in the southern hemisphere.

Distribution of Lampanyctus macdonaldi

sources:  Hildebrand (1948); D.B. Stewart et al. (1991); Nielsen et al. (1992); Treble et al. (2000); Jørgensen et al. (2005); Chambers & Dick (2007).

Lampanyctus macdonaldi



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Myctophum punctatum Rafinesque, 1810

Spotted Lanternfish, lanterne ponctuée

common names: A local name is Slankhalet Prikfisk (Danish/

Greenlandic).

taxonomy:  The genus supposedly comes from nyx (night) and phos (light) although “Myctophum” was never explained by the author, who was unaware that the fish had photophores. An alternative derivation is the Greek mykter (nose) and ophis (serpent). The species name comes from the Latin punctatus (spotted). description:  This species is distinguished by the AO row being subdivided into an anterior (AOa) and a posterior group; the PLO photophore is more than its diameter above the level of the upper base of the pectoral fin; the second PVO is at or below the upper base of the pectoral fin; the PVO row is inclined; there are two Prc

photophores; the Dn is minute; PVO1 is more than its diameter below PVO2; Vn is large and well defined; VO2 is level; SAO is in a straight or slightly angled line; the first SAO photophore is behind the third VO photophore; the eyes are not semi-telescopic; the dorsal fin-rays number 13–14; and the anal fin-rays number 20–22; There are 14–18 AO photophores and 7–10 AOp photophores with 3–4 over the anal fin base. The pectoral fin-rays number 13–15. The lateral-line organs number 43–44. The total gill rakers are 22–27. Adult males have one to four overlapping scale-like structures edged narrowly with black in the supracaudal luminous gland. The gland begins to develop at 3.5 cm. Adult females have two to five heartshaped, overlapping patches in the infracaudal gland. This gland appears at 3.8–4.0 cm. Females may also have one to two supracaudal patches. The species attains 10.7 cm in standard length.

habitat:  This species is found in boreal and temperate wat-

ers. Canadian fish may be expatriates from the south and east, not breeding but continuing to feed and grow to a large size at their range limit. These expatriates show irregular and regressive growth in scales, otoliths, and subopercles, and the caudal luminous glands

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are less developed than in other populations. These features suggest that the fish are under stress and that this is not their optimal habitat. However, some juveniles have been caught in northern expatriate areas, suggesting some reproduction may occur. They are mesopleagic, at 100–1,000 m by day, most abundantly at 700–800 m, and from the surface to 800 m by night, and down as deep as 1,350 m. Arctic Canadian records are from 398–721 m in Davis Strait.

Notoscopelus kroyeri (Malm, 1861)

Northern Saillamp, lampe-voilière du nord

biology:  This Lanternfish feeds on copepods, euphausiids, zoea

stages of Brachyura, and fish fry. It is an important food for Atlantic Cod and Swordfish and is eaten by Fin Whales. Sexual maturity is attained at about 5.0 cm in length. Females produce up to 12,968 eggs. Notoscopelus kroyeri

importance:  It is not economically important. distribution:  The species is found in Davis Strait, based on

three records from cruise data, and in southwest and southeast Greenland. It is found in the Atlantic Ocean from Brazil to Davis Strait, at Jan Mayen Island, and in the Kara and Mediterranean Seas.

Greenlandic). Other common names are Krøyer’s Lanternfish, Lancet Fish, and lanterne de Kroyer.

taxonomy:  The genus comes from the Greek noton (back) and skopelos (said to be an ancient name for a fish, used as a Lanternfish genus by Baron Georges Cuvier in 1816, now a synonym of Myctophum; note that the Greek word means “rock” and “peak,” among others). The species is named after the Danish zoologist Henrik Nikolaj Krøyer (1799–1870), who studied Greenland fishes and authored Danmarks Fiske. The species name has been spelled kroeyerii previously. The species is considered by some authors to be a subspecies of Notoscopelus elongatus (Costa, 1844), described from the Mediterranean Sea. description:  This species is distinguished by a combination of characters including photophore patterns and fin-ray counts. The AO row is subdivided into an anterior (AOa) and a posterior group; the PLO is more than its diameter above the level of the upper base of the pectoral fin; the second PVO is well above the upper base of the pectoral fin; the eyes are not semi-telescopic; and dorsal fin-rays number 20–23. The anal fin-rays number 18–20, usually 19, and the pectoral fin-rays 12–13. Stiff spine-like rays are found at the upper and lower caudal fin bases. The dorsal fin base is longer than the anal fin base. There are 24–31 gill rakers visible in the open mouth. There are no large glands on the upper and lower parts of the caudal peduncle outlined by heavy black pigment. A large luminous gland on top of the caudal peduncle is present in males, but there is no luminous tissue on the cheek or above the eye. In adult males the caudal peduncle gland occupies most of the upper caudal peduncle and is divided into eight to nine segments. Lateral-line organs number 40–43. Photophores are AOa 8–10, usually 9; AO posterior group 6–8, usually 7; total AO photophores 15–18, usually 16; the Dn is well developed; and there are three Prc. The species attains 18.9 cm in total length.

Distribution of Myctophum punctatum

sources:  See the bibliography.



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common names: A local name is Krøyers Prikfisk (Danish/

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Notoscopelus kroyeri

habitat:  This species is pelagic-oceanic and oceanodromous. It is

a common North Atlantic Lanternfish found at 325 m to more than 1,000 m by day, concentrated at 400–500 m, and from the surface to 125 m at night. Migration speed is 1.0–1.5 m/minute. In the Davis Strait it has been caught at 290–980 m, and collections in Hudson Strait were down to 414 m. Off Baffin Island in the Davis Strait it has been caught at 545–1,451 m and 1.1°C–4.5°C. In the Canadian Arctic it is found in less than 10% of sets. Off the Atlantic coast of Canada it is found at temperatures of 3.0°C–23.4°C.

biology:  Euphausiid crustaceans are its most important food

item. In Canadian waters it is food for Atlantic Cod in Ungava Bay, and for Atlantic Salmon at Killiniq (Port Burwell), Ungava Bay. It is also eaten by seals. Spawning occurs mainly in February–March in Canadian waters of the Grand Banks–Flemish Cap when fish are about three years old. Its maximum age is nine years. Growth is slower until age three in the northwest Atlantic, as opposed to the growth in the northeast. One-year-old fish, 50–70 mm long, in the Labrador Basin have been carried by currents out into the North Atlantic from the Grand Banks and back again.

importance:  It is not economically important. Distribution of Notoscopelus kroyeri

distribution:  The species is found in Davis Strait, eastern Hud-

son Strait, and the northern Labrador Sea. It is found in the North Atlantic Ocean only in temperate to subpolar waters. It has been reported from southwest and southeast Greenland south to the Gulf of St Lawrence, off the Scotian Shelf, and south of the Flemish Cap.

330

sources:  Karrer (1973); Nafpaktitis (1975); Neilson & Gillis (1979); Mazhirina & Filin (1987); Filin (1990, 1997, 1998); Hudon (1990a); D.B. Stewart et al. (1991); Jørgensen et al. (2005).

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biology:  Its food is probably crustaceans. These Lanternfishes

Protomyctophum arcticum (Lütken, 1892)

Arctic Telescope, télescope arctique

common names:  A local name is Arktisk Prikfisk (Danish/

are food for Ivory Gulls when they are concentrated at night in the southwest Davis Strait, which indicates a shallower occurrence at this latitude. Northern Fulmars, Black-legged Kittiwakes, and Ringed Seals also eat this species in southern Davis Strait. Sexual maturity is attained at 3.0 cm in length.

Greenlandic).

importance:  It is not economically important.

taxonomy:  The genus comes from the Greek proto (first or original) and supposedly from nyx (night) and phos (light) although “Myctophum” was never explained by the author, who was unaware that the fish had photophores. An alternative derivation is the Greek mykter (nose) and ophis (serpent). The species name comes from the Greek arktikos (Arctic), having been described from Davis Strait.

distribution:  The species is found in the Davis Strait off southern Baffin Island, and from 70°29' N on the west coast of Greenland, south to below 40° N in the western Atlantic Ocean.

description:  This species is uniquely distinguished by the semi-­ telescopic eyes, that is, elongate and directed upwards; the AO row is continuous; and the PLO is below the upper base of the pectoral fin. The PLO is usually slightly higher than the first PVO photophore. The PLO and the first and second PVO photophores do not form a triangle shape. The lateral line is poorly developed. AO photophores number 14–16, the Dn is minute, and there are two Prc. The dorsal fin-rays number 10–13, usually 12; anal fin-rays 21–24; and pectoral fin-rays 14–17. Total gill rakers number 18–21. Adult males have a single supracaudal luminous gland, edged in black and first appearing at 2.1 cm. Females have a single infracaudal patch at 2.5–2.6 cm, a second patch develops later, and, when the fish attain 3.2 cm, the two patches coalesce. The species reaches 6.0 cm in standard length. habitat:  This is an epipelagic, bathypelagic, and oceanodromous

species. It is the second most common Lanternfish in the Newfoundland Basin after the Glacier Lanternfish. Depth distribution generally is 250–850 m by day and 90–325 m by night. The largest catches are taken at 350 m by day and 250 m by night. Elsewhere it occurs at much greater depths, below 1,600 m.

Distribution of Protomyctophum arcticum

sources:  Bekker (1967); MacLaren Marex (1979a, 1979b); Orr & Parsons (1982); Nielsen et al. (1992).

Protomyctophum arcticum



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Symbolophorus veranyi

biology:  It is eaten by Swordfish in southern Canadian waters.

(Moreau, 1888)

importance:  It is not economically important.

North Atlantic Cornerlantern, lanterne-de-coin nord-atlantique

distribution:  In Arctic Canada Symbolophorus sp. has been

common names: Other common names are Largescale Lanternfish and lanterne à grandes écailles.

recorded from northern Ungava Bay at ca. 60°55' N, 66°44' W (Allard, 1980), and S. veranyi from Davis Strait without an accurate locality (D.B. Stewart, Dunbar, & Bernier, 1993). The species is found also in the North Atlantic Ocean, the Mediterranean Sea, and off Brazil.

taxonomy:  The genus comes from the Greek symbolon (mark,

signal) and pherein (to carry). The species is named after Chevalier Jean Baptiste Vérany (1800–65), a French naturalist and co-founder of the Muséum d’histoire naturelle de Nice.

description:  This species is distinguished by a combination of characters including photophore patterns and fin-ray counts. The AO row is subdivided into an anterior and a posterior group; the PLO photophore is above the level of the upper pectoral fin base; the second PVO photophore is at or below the level of the upper pectoral fin base, the PVO photophores are not horizontal but inclined, and the first PVO is more than its diameter below the second PVO; there are two Prc photophores; the Dn is minute; the Vn is large; the SAO is strongly angled, the first SAO photophore is in advance of the third VO photophore; the eyes are not semi-telescopic; and the anal fin-rays number 21–23. Lateral-line organs number 41–42. AO photophores number 14–17. VO photophores are level, and there is one Pol photophore. The dorsal fin-rays number 12–14, and the pectoral fin-rays 12–14. Total gill rakers are 18–21. Adult males have a supracaudal luminous gland made up of one to three small coalescing patches, which appears in fish of 5.2 cm in length. Adult females have two to four patches infracaudally, which appear at 5.8 cm in length. The species reaches at least 13.0 cm in length. habitat:  This Lanternfish favours temperate waters and is not

? ?

Distribution of Symbolophorus veranyi

sources:  Allard (1980); Hudon (1990a); D.B. Stewart, Dunbar, & Bernier (1993).

uncommon off the Atlantic Canadian coast. It is found at 100 m and at 550–800 m during the day and most commonly at the surface at night, but also more rarely down to 800 m.

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Family Macrouridae Grenadiers, Grenadiers

Claude B. Renaud and Brian W. Coad

The family, also commonly called the Rattails, is distributed in all oceans and comprises four subfamilies (Bathygadinae, Macrourinae, Macrouroidinae, Trachyrincinae), 34 genera, and over 350 species, including 26 in Canada, 9 of which occur in Canadian Arctic waters. Members of this family vary in adult total length from 25 to 150 cm. The four subfamilies have each been treated as distinct families in alternative classifications. Grenadiers are characterized by their tadpole-like appearance, a largish head, and a long tapering tail, which is subject to breakage and regeneration, and the absence of a caudal fin, except in one genus of Trachyrincinae (Trachyrincus), which possesses a minute one. A chin barbel is usually present. Teeth are present on the premaxillae and dentaries but not on the roof of the mouth. Branchiostegal rays number 6–8. Both the second dorsal fin and the anal fin are very elongate and about the same length. Counts of rays in these fins are seldom given because of the high number (sometimes over 150 rays). The first dorsal fin is high, and the second very low. The Bathygadinae and Macrourinae do not possess true spines in their first dorsal fin but have two pseudospines, which are unsegmented, unpaired, modified soft rays; the Trachyrincinae possess only one pseudospine, and the Macrouroidinae none. Most possess cycloid scales with spinules, except for the Bathygadinae, which have no spinules on their scales. Although these spinules superficially resemble the ctenii on the ctenoid scales of higher fishes, they are not developed from the same structures. The subfamily Bathygadinae contains two genera and 25 species, which have a large terminal mouth, a rounded snout, rays of the second dorsal fin that are longer than those of the anal fin, and long, slender gill rakers. The Macrourinae contains 28 genera and over 320 species, although well over half of the species are found in only three genera – Coelorinchus (occasionally misspelled Caelorinchus), Coryphaenoides, and Nezumia – and the subfamily is characterized by a subterminal or an inferior mouth, a blunt or pointed snout, rays



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of the second dorsal fin that are shorter than those of the anal fin, and tubercular gill rakers. The Macrouroidinae contains two genera with a single species each, which possess an inferior mouth, a large bulbous head, one low dorsal fin only, and long, slender gill rakers. The Trachyrincinae has two genera and six species, which possess a small inferior mouth; a long, pointed, and flattened snout; two dorsal fins; relatively short rays in the first dorsal fin; rays of the second dorsal fin that are somewhat longer than those of the anal fin; large, rough scutes along the base of the dorsal and anal fins; and relatively short, slender gill rakers. Grenadiers are deep-water benthopelagic or bathypelagic fishes. The great majority of macrourids occupy the continental shelf and slope at depths between 200 m and 2,000 m, but some are found as deep as 6,000 m or more. The total world catch of macrourids in 1987 amounted to 51,226 t, 40% of which was attributable to one species, Coryphaenoides rupestris. None of the nine species that occur in the Canadian Arctic is utilized locally.

sources:  Cohen, Inada, Iwamoto, & Scialabba (1990); Endo (2002); Orlov & Iwamoto (2008); Devine et al. (2012).

Coryphaenoides armatus (Hector, 1875)

Russet Grenadier, grenadier roux

common names:  A local name is Pansret Skolæst (Danish/

Greenlandic). Other common names are Abyssal Grenadier, Armed Grenadier, Cosmopolitan Rattail, and Smooth Abyssal Grenadier.

taxonomy:  The genus comes from the Greek koryphaina (dolphin fish) and oides (similar to). The species name comes from the Anglo-Saxon arm (arm) and the Latin atus (pertaining to), and thus, “armed.” It was originally described under the genus Macrurus. This species belongs to the subfamily Macrourinae. Some authors place it in the genus Nematonurus or the subgenus Nematonurus of Coryphaenoides. Synonyms include Coryphaenoides variabilis Günther, 1878, described from the central North Pacific Ocean; Macrurus asper Goode and Bean, 1883, and Macrurus goodii Günther, 1887, both described from the western North Atlantic Ocean; Coryphaenoides gigas Vaillant, 1888, described from the eastern North Atlantic Ocean; Nematonurus cyclolepis Gilbert, 1896, described from the eastern North Pacific Ocean; Macrurus suborbitalis Gill and Townsend, 1897, described from the Bering Sea; and Nematonurus abyssorum Gilbert, 1915, described from off California. According to authors, there are two subspecies: C. armatus variabilis in the North Pacific Ocean and C. a. armatus everywhere else.

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Coryphaenoides armatus

description:  This species is distinguished from its congeners by the upper jaw extending back to the posterior third of the orbit or more; the total inner gill rakers on the first arch number 11–14; the mandibular teeth are in one row; the premaxillary teeth are in one to two rows, one in large adults; and the interorbital width is 21%–27% of the head length. The mouth is of moderate size and subterminal to inferior. The upper jaw projects beyond the lower jaw. The snout is pointed, without any enlarged scutes, and is partially naked on the underside. The premaxillary teeth are large. Of head length, the chin barbel is 9%–23%, the snout length is 20%–31%, the eye diameter is 14%– 27% (usually 23% or less), and the interorbital width is 21%–27%. There are two well-separated dorsal fins: the first fin is short, with two pseudospines, the second pseudospine being serrated, and has 8–10 rays; the second fin is long and has similarly sized short rays. The anal fin is long and has similarly sized longer rays, and its origin is anterior to the origin of the second dorsal fin. The pelvic fins have the first ray elongate and a total of usually 10 rays in the Atlantic Ocean and 11–12 rays in the Pacific Ocean. The pectoral fins have 17–22 rays. The branchiostegal rays number 6. The total outer gill rakers number 7–9. The outer gill slit is 12%–18% of head length. The cycloid scales are spinulated (in 3–10 divergent rows, but the spinules diminish with age), thin, deciduous, and cover the body but are mostly absent from the ventral aspects of the head. The pyloric caeca number 12. In colour the body is blackish blue to brownish, and the tail is brownish grey. The fins, mouth, gill cavity, and peritoneum are blackish. The species attains 102.0 cm in total length. habitat:  The species is benthopelagic to bathypelagic, at depths

of 257–5,180 m, and is usually found below 2,500 m. In the Canadian Arctic small individuals (16–18 cm in total length) have been collected at 1,950–1,960 m depth. Elsewhere larger, older individuals are found at greater depths.

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biology:  As young it feeds on benthic invertebrates (amphipods,

isopods, cumaceans, bivalve molluscs, and sea cucumbers), and as adults mostly on mesopelagic and bathypelagic fishes, sea urchins, and cephalopods. While young adults rely on vision to detect food, older adults use smell and are known to be opportunistic scavengers. The switch in the sensory mechanism used for food detection occurs at around 40–50 cm in total length. The larger Russet Grenadier is a solitary forager that swims constantly in a cross-current fashion in order to pick up odours. Temporary aggregations of individuals only occur at food falls. After feeding on a carcass, individuals disperse in different directions. This species reaches at least 29 years of age. Fecundity is several million eggs per female. It is believed that the Russet Grenadier is a semelparous (dying after a single spawning event) broadcast spawner. However, the time and the location of a spawning event have yet to be documented. The Russet Grenadier is one of the most abundant grenadier species in the world’s oceans at depths greater than 2,000 m; its abundance has been estimated at 38 billion fish. Individuals are estimated to travel up to 3,000 km/year. The mean swimming speed is 0.17 body lengths per second. The mean respiration rate is 3.2 mL oxygen per hour per kg, a metabolic rate that is 3.7–7.1 times lower than measured in Atlantic Cod at a similar temperature. Internal parasites include cestodes and trematodes.

importance:  It is not economically important. distribution:  It has been found in Davis Strait as three records – ZMB 23761, ZMB 23762, and Jørgensen (1996). It is also found in southwest (up to 63° N) and southeast Greenland and in all oceans, except the central Arctic, at latitudes between 65° N and 61° S.

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taxonomy:  The species name comes from the Latin brevis (short) and barba (beard). The species belongs to the subfamily Macrourinae. Some authors place it in the subgenus Chalinura either under Coryphaenoides or under Macrurus. Others consider Chalinura to be a distinct genus, and this species was originally described under that genus.

Distribution of Coryphaenoides armatus

sources:  Haedrich & Polloni (1976); Karrer (1976); Wilson & Waples (1984); Jørgensen (1996); FAO (2011); Hunter, Bergstad, Høines, & Gjelsvik (2012); Orlov (2012); Bergstad (2013).

description:  This species is distinguished from its congeners by the upper jaw extending back to the posterior third of the orbit or more; the total inner gill rakers on the first arch number less than 17; the mandibular teeth are in one row; the premaxillary teeth are in three or more rows; and the interorbital width is 26%–41% of the head length. The mouth is subterminal, the upper jaw projecting beyond the lower jaw. The snout is blunt, overhangs slightly beyond the upper jaw, and is partially or fully naked on the underside. The chin barbel is 8%–19% of head length, the snout length 15%–44%, the eye diameter 16%–26% (usually 23% or less), and the interorbital width 26%–41%. The body scales have a few slightly divergent rows of depressed spinules. There are two dorsal fins and one anal fin. The fin-rays of the second dorsal fin are shorter than those of the anal fin. The first dorsal fin has two pseudospines, the second pseudospine being serrated, and 9–10 rays. The pectoral fins have 18–20 rays, and the pelvic fins 8–9 rays, the first one elongate. The branchiostegal rays number 6. The outer gill slit is 14%–30% of head length. The pyloric caeca number 10. The colour of the body is pale brown, brownish black, pale grey, or whitish silvery. The lips, the lining of the gill cavities, and the peritoneum are blackish. Juveniles are almost black on the head and abdomen and brownish black elsewhere. The species attains 45.0 cm in total length. habitat:  This species is bathypelagic, occurring at depths

Coryphaenoides brevibarbis (Goode and Bean, 1896)

Shortbeard Grenadier, grenadier à barbe courte

common names: A local name is Butsnudet Skolæst (Danish/

Greenlandic).

between 430 m and 4,700 m. It is found at 1,056–1,461 m and 3.0°C–3.4°C in the Canadian Davis Strait, and also at 429.5–1,441.5 m and 2.9°C–3.5°C. In the Canadian Arctic, individuals of 13–45 cm in total length have been collected from 1,950–1,960 m depth.

biology:  Its food consists of chaetognaths, small crustaceans (gammarid amphipods, copepods, euphausiids, mysids),

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polychaetes, fishes, and cephalopods. The sex ratio is about one male to one female. Its lifespan is about 14–15 years. It spawns in Greenland waters.

importance:  It is not economically important. distribution:  The species is found in Davis Strait, in southwest (at least 64° N) and southeast Greenland, and south to Georges Bank, southwest of Nova Scotia. It occurs east of southern Greenland, south of Iceland, west of Ireland, and as far south as the Bay of Biscay off northern Spain.

Coryphaenoides carapinus Goode and Bean, 1883

Carapine Grenadier, grenadier à barbillon court

common names: A local name is Tipsnudet Skolæst (Danish/

Greenlandic).

taxonomy:  The species name comes from the Latin –inus (pertaining to) and from Carapus, an unrelated group of fishes with a similar body shape. The species belongs to the subfamily Macrourinae. It has been placed in the genera Chalinura, Lionurus, and Macrurus, and some authors place it under the subgenus Lionurus. Nematonurus farrani Fraser-Brunner, 1935, described from the Irish Atlantic slope, is a synonym. description:  This species is distinguished from its congeners by

Distribution of Coryphaenoides brevibarbis

sources:  Karrer (1976); Jørgensen (1996); Jørgensen et al. (2005); R. Hunter et al. (2012); Orlov (2012); Bergstad (2013).

the upper jaw extending back to the posterior third of the orbit or more; total inner gill rakers on the first arch number less than 17; and the mandibular teeth are in two or more rows. The mouth is small and inferior, with the upper jaw projecting beyond the lower jaw. The snout overhangs the upper jaw and is partially naked on the underside. Rows of pointed teeth are present in the upper and lower jaws. The chin barbel is 8%–25% of head length, the snout length 24%–44%, the upper jaw 33%–39%, the eye diameter 11%–35% (usually 23% or less), and the interorbital width 22%–40%. There are two dorsal fins. The first dorsal fin has two pseudospines, the second pseudospine being serrated, and 8–9 finrays. The anal fin-rays are longer than those of the second dorsal fin. The pelvic fins have 9–11 rays, the first of each being elongate. The pectoral fins have 17–20 rays. The branchiostegal rays number 6. The cycloid scales have a few slightly divergent rows of depressed spinules. Total outer gill rakers on the first arch number 8–11 and are stubby. The pyloric caeca number 9. In colour the body is blackish blue to pale grey, and the tail is brown grey. The peritoneum is dark

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brown. The species attains 45.0 cm in total length. An individual of 39.0 cm total length weighs 185 g.

Coryphaenoides guentheri

habitat:  This species is benthopelagic at depths between 384 m

Günther’s Grenadier, grenadier de Günther

and 5,610 m but is usually found above 2,400 m. The larger individuals are mostly female and inhabit greater depths. In the Canadian Arctic it has been collected at 1,950–1,955 m depth.

biology:  It feeds almost exclusively on benthic organisms such

as annelids (polychaetes), crustaceans (amphipods, copepods, isopods, mysids, and decapods), and brittle stars (ophiuroids). It often hovers, head down, to feed. It reaches its greatest abundance (2.4 individuals per 1,000 sq m) in the transition area between the continental slope and the continental rise (1,800–2,400 m). Females reach maturity at about 7 cm total length. The lifespan is about 14–15 years. Fecundity varies from 50,000 to 220,000 eggs per female. The eggs are 0.5 mm in diameter. It is believed to spawn in winter. Internal parasites include cestodes, nematodes, and trematodes.

importance:  It is not economically important. distribution:  The species has been found in Davis Strait as two records, ZMB 23763 and Jørgensen (1996). It is also found in southwest Greenland and occurs throughout the North Atlantic Ocean between 63° N and the equator, in the southeast Atlantic Ocean, and in the Indian Ocean.

(Vaillant, 1888)

common names: A local name is Günthers Skolæst (Danish/ Greenlandic).

taxonomy:  The species name honours Albert Karl Ludwig Gotthilf Günther (1830–1914), a German-born British zoologist, ichthyologist, and herpetologist who at the time was Keeper of Zoology at the Natural History Museum, London. Originally described under the genus Macrurus, the species belongs to the subfamily Macrourinae and to the subgenus Coryphaenoides. At various times it has been placed in the genera Chalinura, Lionurus, and Macruroplus. Macrurus ingolfi Lütken, 1898, described from the southwest of Iceland, is a synonym. description:  This species is separated from its congeners by the upper jaw not extending back to the posterior third of the orbit. The mouth is small and inferior, and the upper jaw projects beyond the lower jaw. The pointed snout has a scute at its tip and one on each side. The snout overhangs the upper jaw. The teeth are small and numerous on both jaws. The chin barbel is 6%–14% of head length, the snout length 29%–36%, the eye diameter 21%–37%, and the interorbital width 21%–26%. There are two separate dorsal fins, the space between them being 16%–48% of head length. The first dorsal fin is short, with two pseudospines, the leading edge of the second pseudospine being serrated, and has 9–11 rays. The second dorsal fin is long and has very short, similarly sized rays. The anal fin is long, with similarly sized but longer rays, its origin being well ahead of that of the second dorsal fin. The pelvic fins have 7–9 rays, usually 8, the first one being only slightly elongate. The pectoral fins have 19–22 rays.The branchiostegal rays number 6. Cycloid scales are distributed over most of the body, but much of the snout region is scaleless. The body scales have depressed spinules in 8–12 nearly parallel or convergent rows (note that scales may often be lost in this species and others). The gill rakers on the first arch are tubercular and spiny. The body colour is light greyish brown, and bluish on the abdomen and gill covers. The fins are pale to dark. The mouth and gill cavities are black. The pyloric caeca number 10–11. The species attains 58.0 cm in total length. habitat:  It is benthopelagic at depths of 420–2,830 m, usually

Distribution of Coryphaenoides carapinus

sources:  Haedrich & Polloni (1976); Karrer (1976); Jørgensen (1996); R. Hunter et al. (2012); Orlov (2012); Bergstad (2013).



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below 1,000 m and at temperatures from 3.0°C to 4.3°C, but it prefers temperatures between 3.2°C and 3.6°C. Larger adults are found at the greater depths. The species can occur in schools of about 200 individuals. In the Canadian Arctic it has been recorded at depths down to 1,960 m. It is found at 1,156–1,463 m and at 3.1°C in the Canadian Davis Strait, and generally in Davis Strait at 1,062.5–1,457.5 m and 3.3°C–3.8°C.

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Coryphaenoides guentheri

biology:  Its most important food items are crustaceans

(gammarid amphipods) and annelids (polychaetes), followed by holothuroids (sea cucumbers) and bivalve molluscs, indicating benthic feeding habits. It is also reported to feed on other benthic crustaceans (ostracods, isopods, and mysid shrimps), copepods, oligochaetes, cephalopods, and fishes. The sex ratio is about one male to one female. The maximum age is 31 years. The spawning period is in the spring.

importance:  It is not economically important. distribution:  This species is found in Davis Strait, in southwest (to 66° N) and southeast Greenland, and south to northern Labrador in the west. It occurs in the northeast Atlantic Ocean between 20° and 63° N and in the Mediterranean Sea.

sources:  Karrer (1976); Jørgensen (1996); Jørgensen et al. (2005); R. Hunter et al. (2012); Orlov (2012); Bergstad (2013).

Coryphaenoides rupestris Gunnerus, 1765

Rock Grenadier, grenadier de roche

common names: A local name is Almindelig Skolæst (Danish/ Greenlandic). Other common names are Black Grenadier, Bluntnose Grenadier, Blunt-snouted Grenadier, Roundnose Grenadier, and Round-nosed Grenadier. taxonomy:  The species name is the Latin rupestris (living among rocks). The species belongs to the subfamily Macrourinae and to the subgenus Coryphaenoides. Various authors have placed it in the genera Coryphaena and Macrurus. Synonyms include Macrourus stroemii Reinhardt, 1825, and Lepidoleprus norvegicus Nilsson, 1832, both described from Norway.

Distribution of Coryphaenoides guentheri

338

description:  This species is distinguished from its congeners by the upper jaw extending back to the posterior third of the orbit or more; total inner gill rakers on the first arch number 17–20; and the snout is rounded and tipped with a large, blunt scute. The mouth is inferior, with the upper jaw projecting beyond the lower jaw. The tip of the snout has a large, blunt scute. The snout overhangs the upper jaw and is rounded and fully scaled. The head is large with numerous scaleless ridges. Rows of small, sharp teeth are present on the premaxillae (upper jaw) and the dentaries (lower jaw). A small chin barbel is 2.5%–8.0% of head length, the snout length 19%–37%, the eye diameter 19%–39% (usually 24% or more), and the interorbital width 29%–40%. There are two separate dorsal fins, the space between them being 37%–54% of head length.

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The first dorsal fin is short with two pseudospines, the second pseudospine being serrated, and has 8–11 rays. The second dorsal fin is long and has 103–183 very short, similarly sized rays. The single anal fin is long, with 104–193 similarly sized but longer rays, and its origin is well ahead of the origin of the second dorsal fin. The pelvic fins have 7–8 rays, the first one being elongate and about equal to head length. The pectoral fins have 16–23 rays. The lateral line is straight and above the midline. The branchiostegal rays number 6. The gill rakers on the first arch are tubercular and spiny. Small, relatively adherent cycloid scales with highly dense spinules are present on the head, including its underside, and those of the trunk and tail are larger and highly deciduous. The body scales have many slender retrorse spinules arranged randomly. The vertebrae (including the urostyle) number 63–79, rarely 89. The pyloric caeca number 29–31 and are long and slender. The body colour is brownish to greyish violet. The orbits, fins, mouth, and gill cavities are blackish to brownish grey. The species attains a maximum total length of 150.0 cm and weight of 1.9 kg.

habitat:  Benthopelagic to bathypelagic from 155 m to 3,000 m

depth, the species usually occurs between 500 m and 1,700 m. It has been found associated with different substrate types, from soft to hard bottoms, and over deep-water coral reefs. On the Canadian side of Davis Strait the fishes are concentrated between 600 m and 800 m depths. Across its range it is found in waters from 0.6°C to 11.5°C, but in Baffin Bay and Davis Strait it prefers waters from 0.6°C to 5.6°C. Adults undergo vertical daily migrations of up to 1,000 m for feeding purposes. However, it is considered to be a poor swimmer, unlikely to undertake extended migrations. It may occur in schools of 250 or more.

biology:  Its diet consists of pelagic crustaceans (copepods, cumaceans, and shrimps, notably glass shrimps) and to a lesser extent mysids, euphausiids, hyperiid amphipods, ostracods, isopods, tunicates, arrow worms, and small fishes belonging to the families

Myctophidae (Lanternfishes), Gonostomatidae (Bristlemouths), Serrivomeridae (Sawtooth Eels), Stomiidae (Dragonfishes), Paralepididae (Barracudinas), Bathylagidae (Deepsea Smelts), Cottidae (Sculpins), Alepocephalidae (Slickheads), Platytroctidae (Tubeshoulders), redfishes (Sebastes spp.), and Macrouridae (Grenadiers), and their eggs, polychaetes, ctenophores (comb jellies), squid, and sea urchins. Most of the feeding occurs during the autumn and winter months. They are an important food item for Greenland Halibut in Davis Strait. Whales and redfishes also feed on them. It is a slow swimmer, caught by trawls at speeds below three knots. Its first year of life is spent as a pelagic larva and juvenile, and this is followed by a demersal phase. The fish size generally increases with depth and latitude. There is an exception in Rockall Trough, off the west coasts of Scotland and Ireland, where large adults are most abundant in the shallowest depths (500–750 m), a mixture of juveniles and large adults occurs at intermediate depths (about 1,000 m), and fish of intermediate length are found at the greater depths. Females grow at a faster rate than do males in both length and weight. Fish measuring 60–80 cm in total length are 10–15 years old, and those of 95 cm in total length have been determined to be 25 years old based on scales; another study using otoliths claims that fish of 90–100 cm in total length can attain 40 years or more. The maximum reported age is 72 years. The generation time is 17 years. Spawning takes place in batches and is believed to occur off the bottom in water deeper than 1,400 m. Across its range the spawning is thought to occur throughout the year, the peaks of intensity varying with location. In the northeast Atlantic the peak spawning takes place around Iceland from January to April, and June to September, and in the Skagerrak in late autumn. Spawning has so far not been confirmed in the northwest Atlantic although mature specimens are encountered in Davis Strait and on the Scotian Shelf. Some authors have suggested that spawning may occur at deeper waters in the northwest Atlantic than those usually being fished (i.e., > 1,000 m), and this would explain why spawning has so far gone unreported in this area. The alternative is that the fishes in Davis Strait represent

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a dead-end population, with drifting eggs and larvae continuously being recruited from off Iceland, with no evidence of a return migration to Iceland. The sex ratio in the northwest Atlantic Ocean is about 65% males and 35% females. On average, females become sexually mature at 79–83 cm in total length, and their fecundity ranges from 6,000 to 63,500 eggs, with a mean of about 30,000. The egg diameter is 1.8–2.6 mm. The eggs are spherical and free floating and possess a single oil globule. On average, males become sexually mature at 70–76 cm in total length. Some individuals are mature at age 6, but others remain immature until age 20, the average age being 11 years. Females may not spawn every year. Drumming muscles are present on the gas bladder and may be important in producing sound for communication. At least 20 species of parasites have been recorded in Rock Grenadier from Davis Strait to the Grand Banks. The parasites include two nematodes, Anisakis simplex and Hysterothylacium aduncum, which are potentially dangerous to humans.

importance:  The flesh is of excellent texture and taste and rich

in vitamins A, B2, and B12. A directed fishery began on both the Canadian and the Greenland sides of Davis Strait in 1967, involving factory stern trawlers that used primarily bottom, but also midwater, trawls at depths usually between 500 m and 1,500 m. Russia and Germany were the two main countries engaged in this fishery. Annual catches between 1967 and 1989 varied from a high of 12,318 t (1974) to a low of 50 t (1984). In the northwest Atlantic Ocean as a whole, the greatest catch (83,700 t) was taken in 1971, but continuous declines in the annual catch in subsequent years resulted in the closure of the directed fishery in the early 1990s. It was sold frozen and as fish-meal. The estimated biomass and abundance in NAFO Division 0B in 2000 were 1,660 t and 9.2 million fish, decreasing in 2001 to 1,256 t and 7.9 million fish. This species is currently showing signs of over-exploitation. Rock Grenadier was assessed as “Endangered” by the Committee on the Status of Endangered Wildlife in Canada. It is a by-catch of the Greenland Halibut, Atlantic Cod, and Sebastes spp., as well as the deep-water shrimp fisheries.

distribution:  The species is found at Mittimatalik (Pond Inlet) on northern Baffin Island, in Davis and eastern Hudson Straits, and in northwest, southwest, and southeast Greenland. It occurs on both sides of the North Atlantic Ocean in generally temperate waters from about 37° N (off the United States) to 70°44' N (Baffin Bay, Greenland), off Iceland and Norway (70° N), with isolated records at lower latitudes in the Bahamas (24° N) and off North Africa (20° N).

Distribution of Coryphaenoides rupestris

sources:  Richardson (1836a); Kotthaus & Krefft (1957); Anonymous (1960a); Konstantinov & Podrazhanskaya (1972); Parsons (1975); Haedrich & Polloni (1976); Sandeman & Buchanan (1979); Atkinson, Bowering, Parsons, Horsted, & Minet (1982); Atkinson (1983b, 1995); Chumakov & Podrazhanskaya (1986); Crawford (1992); Jørgensen (1996); Treble et al. (2000); Jørgensen et al. (2005); Rodger (2006); COSEWIC (2008); M.R. Simpson, Miri, et al. (2011); Devine et al. (2012); R. Hunter et al. (2012); Orlov (2012); Bergstad (2013).

Gadomus longifilis (Goode and Bean, 1885)

Threadfin Grenadier, grenadier à filaments

common names: A local name is Langfinnet Skolæst (Danish/

Greenlandic). The common name was coined by Claude B. Renaud.

taxonomy:  The genus name was not explained, possibly derived from Gadus and the Latin domus (a family or race). The species name comes from the Latin longus (long) and filum (thread), in reference to the long barbel or possibly to the elongate fin-rays. description:  This species is distinguished by the second dorsal

fin-rays being equal to or longer than the anal fin-rays; the gill rakers are elongate; the outer gill slit is without a fold over the upper and lower parts; the gap between the first and second dorsal fins is small;

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Gadomus longifilis

the mouth is wide and terminal; the snout is rounded; the scales are without spinules; a post-temporal fossa is absent; and there are very elongate rays in the first dorsal, pectoral, and pelvic fins. The barbel is long, longer than eye diameter, and 40%–50% of head length. The interobital space is wide, 21%–25% of head length. The teeth in the jaws are in a narrow band of four to six teeth, depressible and incurved. The first dorsal fin has 2 unbranched rays, the second ray is elongate (1.5–2.0 times head length), and branched rays number 9–11. The pectoral fin-rays number 13–18, and the pelvic fin-rays 8. The first pectoral and pelvic fin-rays are prolonged into a filament. The lateral line is not strongly arched over the pectoral fin. The lower-arch gill rakers number 27–31. The gill filaments are only 20% of gill-raker length at the cerato-branchial centre. The pyloric caeca are 5–13, increasing in number with size. The species is brownish in colour overall. The mouth and the gill chamber are dark brown to black. There is a dark median stripe on the gill lamellae. The species reaches 30.0 cm in total length.

of its distinctive appearance and its known occurrence in adjacent waters in Greenland.

habitat:  The species is benthic to bathypelagic with a depth range

of 630–2,165 m. The Canadian Davis Strait record was caught at 1,271  m.

biology:  It eats small crustaceans such as copepods and decapods.

Distribution of Gadomus longifilis

importance:  It is not economically important. distribution:  The species is found in Davis Strait, based on a

source:  Howes & Crimmen (1990).

single record from cruise data at 63.0692° N, 58.5297° W, in southwest Greenland, and the Atlantic Ocean and the Mediterranean Sea generally. Voucher specimens deposited in a museum are needed to verify its occurrence in Canadian waters. It is included here because



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Macrourus berglax Lacepède, 1801

Roughhead Grenadier, grenadier berglax

common names:  Local names are Imingoak, Ingmingoak, Ingminniset, and Tupissut (Greenlandic); and Nordlig Skolæst (Danish/Greenlandic). Ingminniset means “it bellows when dying,” in reference to the drumming muscles. Other common names are Rough-headed Grenadier and Onion-eye Grenadier. taxonomy:  The genus comes from the Greek makros (long) and oura (tail). The species name comes from the Norwegian berg (cliff) and lax (salmon). This species belongs to the subfamily Macrourinae. The genus was misspelled Macrurus in the early literature. Early authors used the species name rupestris for this species, which caused confusion with Coryphaenoides rupestris Gunnerus, 1765 (see that species account), until the name berglax was proposed by Lacepède. It has been placed in the genus Coryphaenoides. Macrourus fabricii Sundevall, 1842, described from Norway, is a synonym. description:  This species is distinguished by the origin of the second dorsal fin being in front of the anus level; there are no gill rakers on the outer side of the first gill arch; there are six branchiostegal rays; and the anus is at the anal fin origin. (See under Trachyrincus murrayi for other characters.)

The mouth is small and inferior. The upper jaw projects beyond the lower jaw. The snout is pointed, with a scute at its tip, and overhangs the upper jaw. A prominent suborbital ridge runs from the tip of the snout to the operculum. The head is hexagonal in cross-section. Small pointed teeth are arranged in three to five rows on the premaxillae (upper jaw) and two to three rows on the lower jaw. The chin barbel is 11%–19% of head length, the snout length 30%–34%, and the eye diameter 31%–40%. There are two separate dorsal fins: the first is short with two pseudospines, the second pseudospine being finely serrated, and 9–11 rays; the second dorsal fin is very long with 105–124 short rays becoming progressively shorter posteriorly. There is one anal fin with 113–148 slightly longer rays, its position being well behind the origin of the second dorsal fin. The pelvic fins have 7–9 rays, usually 8, the prolonged second ray measuring 34%–40% of head length. The pectoral fins have 16–20 rays. There are four to six ridges on top of the head with scute-like scales bearing strong spinules. The underside of the snout is scaleless or has only a few spinulated cycloid scales. The rest of the body has small spinulated cycloid scales; those on the top of the head do not overlap, while those elsewhere do. The pyloric caeca number 15–26. The body colour of freshly caught specimens is grey, darker on the ventral aspect of the trunk. The anal fin has a dark edge, and the first dorsal and pectoral fins are darkly pigmented. The maximum total length is 156.0 cm, and weight is 2.3 kg. Fish measuring 50–60 cm in total length weigh 1.0–1.2 kg.

habitat:  The species is benthopelagic between 100 m and 2,740 m

depth, with greatest numbers occurring at 600–1,500 m. It is found at water temperatures from −2°C to 8°C but prefers waters between

Macrourus berglax

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−2°C and 4°C. It has been reported from 230–530 m in Hudson Strait, 223–1,463 m and 0.24°C to 5.6ºC in Davis Strait and southern Baffin Bay, 240–649 m in Ungava Bay, and off Baffin Island at 897 m and 3.6°C. In the Barents Sea it is found at salinities over 35‰. It may occur in schools of 150 individuals. Generally, larger, older individuals are found at greater depths.

biology:  Younger individuals feed preferentially on crustaceans (amphipods and mysid shrimps) and polychaetes, while older individuals prefer crustaceans (notably the shrimp Pandalus borealis), fish (Capelin, Goitre Blacksmelt, Northern Cutthroat Eel, Sea Tadpole, Eelpouts, redfishes, Barracudinas, Sculpins, and Grenadiers), and squid. Ophiura (an echinoderm called serpent star), sea urchins, sea cucumbers, anomuran crabs, isopods, benthic bivalve molluscs, marine gastropods (Buccinum, Fusus), and ctenophores (comb jellies) are also common food items. Cannibalism is known to occur. About 70% of the diet is benthic, and the rest pelagic. The Atlantic Cod is a known predator. The overall sex ratio is about 1:1, but females predominate among the larger individuals, growing faster than males, beginning at the age of seven years. The species may attain an age of 40 years. The Roughhead Grenadier in the Barents Sea undergoes a spawning migration to an area near the Lofoten Islands, just off the Norwegian coast. In the western North Atlantic Ocean a spawning area occurs in the northern Labrador Sea. The spawning period is thought to be prolonged and occurs between late winter and late spring. The age at first maturity is 13–16 years. The generation time is 19 years. Females are sexually mature at 65 cm in total length and produce 2,000–80,000 eggs. The egg diameter is 3–4 mm. It is a batch (serial) spawner because three sizes of eggs are found in the ovaries. Males are sexually mature at 54.0 cm in total length. The males are believed to produce sound through the use of well-developed drumming muscles associated with their gas bladder. The species is known to harbour a wide variety of parasites (hematozoans, myxosporidians, cestodes, nematodes, trematodes, crustaceans, a monogenean, and an acanthocephalan). importance:  There is no large-scale directed fishery for this species. It is taken principally as by-catch, although in the northeast Atlantic Ocean it is occasionally the target of some small fisheries. It was only in the late 1980s that Germany and Portugal began commercial exploitation of this species in the northeast Atlantic Ocean. By 2000 there were already 10 countries participating in the fishery. In 2001 the catch was 8,795 t, 92% of which was taken by Spain. On the Canadian side of Baffin Bay and southeast of Newfoundland it is taken mostly by trawl as a by-catch of the Greenland Halibut fishery. In the northwest Atlantic it is also taken as a by-catch of the deepwater shrimp fishery, and of the Rock Grenadier fishery when it was active. The dramatic declines in the Canadian population of Roughhead Grenadier appear to be due to environmental factors operating on long time scales (North Atlantic Oscillation winter index with a 14-year lag time) rather than over-exploitation. Roughhead Grenadier was assessed as of “Special Concern” by the Committee on the Status of Endangered Wildlife in Canada. In the Barents Sea it is



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taken as by-catch by the Russian fleet in the bottom trawl and longline fisheries for Greenland Halibut, Atlantic Cod, and Haddock. The flesh is white and flaky, and medicinal oil is extracted from the liver.

distribution:  The species is found in Baffin Bay, Davis Strait, Hudson Strait, and Ungava Bay. It is also found in northwest (at least up to 72° N), southwest, and southeast Greenland and on both sides of the North Atlantic Ocean and into the Arctic Ocean between 37° N (off the United States) and 82° N (Spitsbergen).

Distribution of Macrourus berglax

sources:  Kotthaus & Krefft (1957); Anonymous (1960a); Kon-

stantinov & Podrazhanskaya (1972); Parsons (1975); Sandeman & Buchanan (1979); Allard (1980); Allard & Gillis (1986); Gillis, Allard, & Axelsen (1987); Hudon (1990a); D.B. Stewart et al. (1991); Khan, Threlfall, & Whitty (1992); Jørgensen (1996); Treble et al. (2000); Jørgensen et al. (2005); COSEWIC (2007); Devine et al. (2012); Orlov (2012); Bergstad (2013); Neat & Campbell (2013).

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Nezumia bairdii

longer than those of the second dorsal fin. The origin of the anal fin is distinctly anterior to that of the second dorsal fin. The pelvic fins each have 6–7 rays, usually 7, the first of which is elongate. There is a small light organ between the pelvic fin bases, visible as a “window.” The pectoral fins have 16–20 rays. The lateral line has a slight arch anteriorly and straight thereafter. The scales are cycloid with spinules. The body is fully scaled except for a median strip on the underside of the snout. The colour of fresh specimens is grey or brownish grey with a violet tinge on the lower jaw and the trunk, and a silvery shine on the gill covers, the trunk, and the lower half of the tail. The first dorsal fin is pale except for a black membrane between the second pseudospine and the first segmented ray. The pelvic fins are black, and the remaining fins paler. The mouth and the gill cavities are black. The species attains at least 43.0 cm in total length.

common names:  A local name is Bronze Skolæst (Danish/

habitat:  The species is benthopelagic, reported from depths

Nezumia bairdii

(Goode and Bean, 1877) Marlin-spike, grenadier du Grand Banc

Greenlandic). Other common names are Marlin-spike Grenadier, Common Grenadier, and grenadier de Baird.

taxonomy:  The genus comes from the Japanese nezumia (mouse), and the species is named after the American ichthyologist Spencer Fullerton Baird (1823–87), a head of the U.S. Commission of Fish and Fisheries and the first curator and the second Secretary at the Smithsonian Institution. The species belongs to the subfamily Macrourinae. Originally described under the genus Macrurus, it has also been placed in the genera Lionurus and Coryphaenoides. description:  This species is distinguished by having seven branchiostegal rays and the anus well anterior to the anal fin. (See under Trachyrincus murrayi for other characters.) The mouth is inferior (the upper jaw slightly projects beyond the lower jaw). The pointed snout overhangs the upper jaw. The small teeth are in bands on the premaxillae (upper jaw) and on the lower jaw. The chin barbel is 12%–15% of head length, the snout length 30%– 33%, the eye diameter 30%–34%, and the interorbital width 26%–29%. The two dorsal fins are widely separated. The first is short, with two pseudospines, the second pseudospine being serrated anteriorly, and has 9–11 rays. The second is long, with 137–159 very short rays. The single, long anal fin has 120–138 similarly sized rays, which are much

between the surface and 2,295 m, usually 90–780 m. It occurs at temperatures of 2.6°C–13.0°C, but usually 3.0°C–5.0°C, and salinity of 31‰–34‰. Occurrences at depths greatly below 1,000 m should be viewed with suspicion, but it occurs at 529–1,156 m and 3.4°C–3.6°C off Baffin Island, and at 524.5–1,254.5 m and 2.6°C–3.9°C generally in southern Baffin Bay and Davis Strait. It can occur at very shallow depths at lower latitudes in areas where the surface water temperature is cold. It is associated with mud bottoms.

biology:  Its food consists primarily of small benthic

crustaceans (euphausiids and amphipods) and polychaetes, and secondarily of molluscs and fishes. It is preyed upon by Swordfish. The maximum reported age is 11 years. The sex ratio is highly skewed towards females, which account for 70% or more of individuals. At lower latitudes (Gulf of Maine) it spawns in summer and fall. The eggs have not been described. It is known to harbour various parasites (cestodes, nematodes, trematodes, myxosporidians, hematozoans, acanthocephalans). Isopods are often attached behind the first dorsal fin.

importance:  Owing to its small size and limited numbers,

there is no directed fishery for this species, but it is caught by bottom trawlers as by-catch in the fisheries for Atlantic Cod, redfishes (Sebastes spp.), and flounders off Labrador and Newfoundland.

Nezumia bairdii

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distribution:  It is found in Baffin Bay and Davis Strait, as well as in southwest Greenland to 65° N. It is also found in the western North Atlantic Ocean from the Canadian side of Baffin Bay (73°13' N) to as far south as the southern tip of Florida. One questionable record from 3,520 m depth was reported from the eastern North Atlantic Ocean.

Trachyrincus murrayi Günther, 1887

Roughnose Grenadier, grenadier-scie

common names:  A local name is Murrays Skolæst (Danish/Greenlandic). Another common name is Murray’s Longsnout Grenadier. taxonomy:  The genus comes from the Greek trachys (rough) and

rhynchos (snout). The species is named after the pioneering Scottish oceanographer, marine biologist, and limnologist Sir John Murray (1841–1914), born in Cobourg, Ontario, who was a naturalist on the Challenger Expedition (1872–6) to survey the deep oceans and was an editor of the ensuing reports. The species belongs to the subfamily Trachyrincinae. The genus name is often misspelled Trachyrhynchus or Trachyrinchus.

Distribution of Nezumia bairdii

sources:  Anonymous (1960a); Konstantinov & Podrazhanskaya (1972); Karrer (1973); Parsons (1975); Sandeman & Buchanan (1979); Khan et al. (1992); Jørgensen (1996); Jørgensen et al. (2005).

description:  This species is distinguished by having the second dorsal fin-rays equal to or longer than the anal fin-rays (versus shorter in all other Canadian Arctic species); the gill rakers are flattened and elongate (versus short tubercles or absent); the outer gill slit is without a fold over the upper and lower parts (versus present); and a post-temporal fossa, appearing as a dent, is present (versus absent). The mouth is inferior, with the upper jaw projecting beyond the lower jaw. The snout is long and pointed, overhanging the upper jaw, and 39%–42% of head length. Bands of small teeth are present on the premaxillae (upper jaw) and the lower jaw. There is a small chin barbel. The eyes are large, 28%–30% of head length. The first dorsal fin is short with one pseudospine and 9–10 rays; the second dorsal fin is very long. There is one long anal fin, with its origin posterior to that of the second dorsal fin. A minute caudal fin is present but may be

Trachyrincus murrayi



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lost. The pelvic fins have 6–7 rays, the leading one being elongate. The pectoral fins have 21 rays. The branchiostegal rays number 7. Bony ridges are present on the head, with spinulated cycloid scales. Two rows of keeled scutes are present on each side of the body, one along the base of the dorsal fins and the other along the base of the anal fin. On the sides of the body, between these two rows of scutes, there are small, easily lost, spinulated cycloid scales. Apparently there are no scales present on the belly. The body colour is grey. The species attains a maximum of 62.0 cm in total length.

Family Moridae Codlings, Moros

Claude B. Renaud

habitat:  It is benthopelagic and generally occurs between 530 m and 1,630 m depth at temperatures between 3.1°C and 3.8°C. It is found at 507–1,247 m in Davis Strait.

biology:  This species is believed to eat crustaceans. Its maximum

reported age is 40 years. The sex ratio is about 1:1. The spawning period is March–May.

importance:  It is not economically important. distribution:  It is found in Davis Strait and in southwest (to

64°49' N) and southeast Greenland. It is distributed on both sides of the North Atlantic Ocean, being in the western North Atlantic south to Hamilton Inlet, Labrador.

Distribution of Trachyrincus murrayi

sources:  Karrer (1973); Tyler (1988); Jørgensen (1996); Jørgensen et al. (2005); Neat & Campbell (2013).

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Members of this family, also known as Moras, Deepsea Cods, Flatnose Cods, or Hakelings, and in French as Mores, are found in all oceans except the central Arctic. Eretmophoridae is a junior synonym. There are 11 species in Canada, of which 3 are in Arctic waters. There are over 100 species in this family, grouped in about 18 genera, although half of the species belong to only two genera, Physiculus and Laemonema (not in Arctic Canada). The maximum size is about 90 cm in length. The Codlings are relatively elongate, usually 50–70 cm in length, with large gill openings, and have a distinct externally symmetrical caudal fin. The caudal fin is separated from the dorsal and anal fins by a narrow caudal peduncle. The fins are without spines, but one pseudospine (Antimora) or two pseudospines (Halargyreus) may be present in the first dorsal fin. The pseudospines are unpaired and unsegmented modified soft rays. Two dorsal fins and one anal fin are present. However, some species appear to have three dorsal fins and two anal fins because of very deep indentations in the second dorsal fin and the single anal fin, respectively. The pelvic fins are jugular in position (in advance of the pectoral fins) and separated. The cycloid scales are small and overlap. The gas bladder is physoclistous (not connected to the oesophagus) and in firm contact with the auditory capsules (otophysic connection) through a pair of anterior projections. The neural spine of the first vertebra is firmly attached to the rear of the skull. Some species have a light organ on the belly between the pelvic fins. Palatine teeth are always absent, but vomerine teeth may be present on the roof of the mouth. These fishes are pelagic to benthopelagic, ranging in depth from shallow coastal waters to over 2,500 m. The world catch in 1986 was 37,000 t. The three species that occur in the Canadian Arctic are not utilized locally.

sources:  Cohen et al. (1990); Endo (2002).

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Antimora rostrata (Günther, 1878)

Blue Antimora, antimore bleu

common names: A local name is Blå Antimora (Danish/Green-

landic). Other common names are Blue Hake, Flatnose Codling, Long-finned Cod, and Violet Cod.

taxonomy:  The genus comes from the Greek anti (opposite) and

Mora, a related genus. The species name comes from the Latin rostratus (long snouted). The species was originally described under the genus Haloporphyrus. Haloporphyrus viola Goode and Bean, 1879, described from the outer edge of LaHave Bank, Nova Scotia, is a synonym.

description:  This species is distinguished by the snout having shelf-like projections on each side, and the upper jaw projects beyond the lower jaw. The mouth is inferior, and the snout is flattened and pointed, overhanging the upper jaw. There is a prominent bony shelf from the tip of the snout to under the eye on either side of the head. Teeth are present on the premaxillae (upper jaw) and dentaries (lower jaw), and there are sharp teeth on the roof of the mouth (head of the vomer). A chin barbel is present. There are two dorsal fins with their bases touching: the first dorsal fin is short and has 4–7 rays, the first of which (a pseudospine) is elongate; the second dorsal fin is long, is slightly indented, and has 48–58 rays. There is one long, very deeply indented anal fin (appearing as two fins) with 36–49 rays, and its origin is posterior to that of the second dorsal fin. The pelvic fins are jugular with 5–7 rays, the second of these being greatly elongate. The pectoral fin-rays number 17–25. The caudal fin is truncate but with rounded upper and lower lobes.

The scales are cycloid and small, numbering 115 along the straight but faint lateral line. Total outer gill rakers on the first arch number 10. It is distinguished from a closely related species, Antimora microlepis Bean, 1890, in the North Pacific Ocean based on the presence of fewer gill filaments on the first gill arch, 76–90 versus 90–103 according to one author; however, one study found a range of 62–100 gill filaments in A. rostrata from the northwest Atlantic that virtually encompasses both counts, so perhaps A. microlepis is another synonym. The branchiostegal rays number 7. The vertebrae (including the urostyle) number 57–60. The body colour is deep violet or blue grey, but at depths of 2,200 m or more it is black, including the ventral aspect. The species attains 75.0 cm, or more, in total length and about 2.7 kg in weight.

habitat:  The species is benthopelagic and known to occur at

depths between 200 m and 3,277 m and temperatures of 0°C–9°C, but it prefers depths of more than 1,400 m and temperatures of 3°C– 5°C, living close to mud bottoms. Juveniles (< 20 cm total length) predominate at depths less than 1,000 m, and adults predominate at depths greater than 1,000 m. The species is found at depths between 557 m and 1,463 m in Davis Strait, based on 34 collections, but perhaps this is simply an artefact of sampling.

biology:  It feeds predominantly on fishes (Sebastes spp.) and

squids but also on decapods, amphipods, chaetognaths, and polychaetes. It is believed to be an opportunistic scavenger of fishes. Its stomach contents are often lost because the stomach is everted due to the expansion of the gas bladder when the fish is brought to the surface. The growth rates are similar between sexes (one study, however, states that males have a slower rate than females), but males are generally smaller than females, do not appear to live as long, and tend to be found at shallower depths. Longevity based on presumed annuli (not validated) of sagittal otoliths is 25 years.

Antimora rostrata



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No mature individuals, eggs, or larvae have been found in Canadian waters, and there is no evidence of spawning in U.S. waters. A few spent individuals have been collected off Iceland between March and July, but none in spawning condition. A maturing female had an estimated 1,351,300 eggs. The mean swimming speed is 0.39 body lengths per second. The species is known to harbour blood parasites (hematozoans).

importance:  A dominant species at depths between 500 m and

2,800 m, it is taken as a by-catch by trawlers and deep-set bottom long lines of the Greenland Halibut fishery off southeast Newfoundland at the 800–1,500 m depth range, but apparently it is not utilized commercially, probably because its flesh is soft and watery. The gas bladder contains a thick, spongy, white or creamy mass or “foam,” which may serve as a diffusion barrier, and maintains high gas and oxygen pressures. Scientists have studied this species for its “foam” composition, lipid production, and haemoglobin function.

distribution:  The species is found in Baffin Bay and Davis Strait

on both the Canadian side (up to 74° N) and the Greenland side (up to 65° N), and in southeast Greenland. It is found in all oceans except the North Pacific north of 10° N and the central Arctic.

Halargyreus johnsonii Günther, 1862

Dainty Mora, more délicat

common names: A local name is Slank Ridder (Danish/Greenlandic). Other common names are Johnson’s Deepsea Cod, Slender Cod, and Slender Codling. taxonomy:  The genus comes from the Greek als (salt) and

argyros (silver). The species name likely honours the British ichthyologist and botanist James Yate Johnson (1820–1900), who gave the type material from Madeira to the Natural History Museum, London. Halargyreus brevipes Vaillant, 1888, described from off Morocco, and H. affinis Collett, 1904, described from Faroe Bank between Iceland and Scotland, are synonyms.

description:  This species is distinguished by the snout lacking

a shelf-like projection on each side; the upper jaw is not markedly projecting beyond the lower jaw; there is no chin barbel (a tubercle is present); the second dorsal fin-ray is not elongated; and the vomer is without teeth. The mouth is terminal. The lower jaw projects slightly beyond the upper jaw. The teeth on the jaws are very small. The eye diameter is about equal to the snout length. There are two dorsal fins, with bases nearly touching. The first one is short, with 6–9 rays (the first two being pseudospines), none of which are greatly elongate, and the second dorsal fin is long, with 47–59 similarly sized rays. The anal fin is long and deeply indented, with 41–53 rays; its origin is posterior to that of the second dorsal fin. The pectoral fins have 14–19 rays and do not reach the anal fin origin. The pelvic fins are jugular with 5–6 rays, the first two being elongate. The caudal fin is forked. The branchiostegal rays number 7. Total outer gill rakers on the first arch number 20–28 and are longer than the gill filaments. There are about 122 scales along the lateral line. The pyloric caeca number 8–11. Total vertebrae (including the urostyle) number 50–57. The body colour is pale bluish or reddish brown on the dorsal and lateral aspects, and silvery on the ventral aspect. The mouth and the gill cavities are black. The peritoneum is black. The species attains 56.0 cm in total length.

habitat:  This species is benthopelagic to pelagic over the contin-

Distribution of Antimora rostrata

sources:  Templeman (1970a); Konstantinov & Podrazhanskaya (1972); Karrer (1973); Small (1981); Mauchline & Gordon (1984); Gordon & Duncan (1985); D.B. Stewart et al. (1991); Khan et al. (1992); Magnússon (2001); Kulka, Simpson, & Inkpen (2003); Jørgensen et al. (2005); Fossen & Bergstad (2006).

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ental slope. It is found at 700–720 m depth in Davis Strait in the Canadian Arctic and at 653–1,500 m in southwest Greenland. Elsewhere it occurs at depths of 350–3,000 m. It has been captured at temperatures between 3.8°C and 10.2°C.

biology:  It feeds almost exclusively on crustaceans (copepods,

mysids, and shrimps) and incidentally on chaetognaths and fishes. The stomach is often everted and its contents lost when the fish is brought to the surface. The Taillight Gulper (Saccopharynx ampullaceus) and the Black Scabbardfish (Aphanopus carbo) are predators. Its lifespan may be at least 7 years. It is known to occur in large numbers of 100 to 350 or more.

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Halargyreus johnsonii

importance:  It is not economically important.

Lepidion eques

distribution:  It has been found in Davis Strait as a single record at 63°09' N, 60°30' W (ZMB 22566). It has an antitropical distribution, being found also in southwest and southeast Greenland and widely scattered in the Pacific Ocean, throughout the Atlantic Ocean, and along a band around Antarctica at 60° S. In the North Atlantic Ocean it is found off Morocco and Madeira, off the west coast of Ireland, the Faroe Islands, south and west of Iceland, and off the Grand Banks.

Largeye Lepidion, lépidion à grands yeux

Distribution of Halargyreus johnsonii

sources:  Templeman (1968a); Cohen (1973); Karrer (1973); Mauchline & Gordon (1984).



UTP Fishes Book 5pp04.indb 349

(Günther, 1887)

common names: A local name is Blå Ridder (Danish/Greenlandic). Another common name is North Atlantic Codling. taxonomy:  The genus comes from the Greek lepidion (a small scale). The species name is the Latin eques (horseman), in reference to the long dorsal fin-ray that suggests a rider. It was originally described under the genus Haloporphyrus. description:  This species is distinguished by the snout lacking shelf-like projections on each side; the upper jaw is not markedly projecting beyond the lower jaw; a chin barbel is present; the second dorsal fin-ray is very elongated (the first is small and hidden under the skin); and the vomer has a few teeth on its head. The mouth is terminal (jaws of equal length) or subterminal (upper jaw protruding slightly). A chin barbel is present. The large eye is 32.2%–38.5% of head length. The posterior nostril is immediately anterior to the eye. There are two dorsal fins, their bases touching. The first dorsal fin is short, with five rays, the first of these being rudimentary and hidden under the skin, while the second ray is greatly elongate, measuring about 26%–57% of standard length. The second dorsal fin is long and indented, with 52–60 rays. A single indented anal fin has 49–54 rays, and its origin is posterior to that of the second dorsal fin. The pectoral fins have 21–25 rays, and the pelvic fins 7–8 rays, the second and third being elongate. There is a small dark and scaleless patch over a light organ on the belly in front of the anus. The caudal peduncle is slender, and the caudal fin spatulate. The small scales number about 180 along the complete and arched lateral line. There are lateral-line pit organs on the head instead of pores. The branchiostegal rays number 7. Total outer gill rakers on the first arch number 14–22, and there are 8–13 pyloric caeca. Total vertebrae (including the urostyle) number 57–62. Fresh individuals are yellowish brown. The species attains 49.0 cm in total

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Lepidion eques

length. Individuals that were 29–31 cm in total length and partially dehydrated from being in 50% ethanol weighed 150–180 g.

habitat:  The species is benthopelagic at depths of 127–1,880 m,

the larger individuals occurring at the greater depths. It has been caught at 700–899 m in Davis Strait. The fry are pelagic. The species is taken at temperatures between 2.0°C and 9.5°C, but it prefers temperatures between 5°C and 6°C. It occurs singly or in schools numbering over 100 individuals. The maximum abundance occurs at 750 m depth, and the greatest biomass at 1,000 m depth.

biology:  Its preferred food is crustaceans (amphipods, copepods,

decapods) and cephalopods, but also polychaetes. It feeds on the sea bottom (epibenthic) and off the sea bottom (hyperbenthic). The mean length of adults increases with depth. Males and females are very similar in size. Longevity based on otoliths (non-validated) is 13 years for an individual of 32 cm in total length. Spawning occurs in April–May in the Rockall Trough, west of Scotland, and in February–June in the waters south of Iceland. Fecundity ranges from 3,424 to 110,023 eggs for females of 25–33 cm in standard length. Intestinal parasites include nematodes, trematodes, a cestode, and an acanthocephalan. Distribution of Lepidion eques

importance:  It is not economically important. distribution:  It is found in Davis Strait both on the Canadian

side (up to 64°06' N), and on the Greenland side (up to 63°05' N), in southeast Greenland, and south along the coast of Labrador to the southern coast of Newfoundland. It is found in the northwest and the northeast Atlantic Ocean but is rare in the former. It also occurs from the Bay of Biscay, along the western shores of the British Isles, and south and west of Iceland.

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sources:  Kotthaus & Krefft (1957); Templeman (1970a, 1970b); Karrer (1973); Mauchline & Gordon (1980, 1984); Gordon & Duncan (1985); Magnússon (2001).

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sources:  Svetovidov (1948, 1986); Cohen & Russo (1979); Cohen et al. (1990); Endo (2002); Francisco, Robalao, Stefanni, Levy, & Almada (2014).

Family Phycidae Phycid Hakes, Phycidés

Claude B. Renaud

Enchelyopus cimbrius (Linnaeus, 1766)

Fourbeard Rockling, motelle à quatre barbillons

Phycid Hakes are mostly found in the temperate continental-shelf waters of the northern hemisphere. The family comprises 5 genera and 25 species, and only 5–6 of these species are found in the temperate waters of the southern hemisphere. There are seven species in Canada, of which four are found in Arctic waters in two subfamilies (Gaidropsarinae and Phycinae). Both subfamilies are treated as families in an alternative classification. The Phycid Hakes have also been treated as subfamilies of the Family Gadidae and even as part of the subfamily Lotinae under the Gadidae. The AFS list placed Gaidropsarus in Gadidae without providing justification. Gaidropsarus is here placed in Gaidropsarinae within Phycidae. Members of this marine family vary in adult total length from 25 to 120 cm. The main characteristics of the family are gill openings extending above the level of the pectoral fins; the gas bladder is not connected to either the rear of the skull or the oesophagus (a gas bladder is absent in Gaidropsarus argentatus and G. ensis); the neural spine of the first vertebra is firmly attached to the rear of the skull; the scales are cycloid; the fins have soft rays only (except in the genera Enchelyopus and Gaidropsarus, where the first ray of the first dorsal fin is both unpaired and unsegmented and termed a pseudospine); and the eggs are small (< 1 mm in diameter) with oil globules. The species possess one anal fin and a chin barbel. The subfamily Gaidropsarinae contains 3 genera and 15 species, which possess three dorsal fins and two to four barbels on the snout. The Phycinae contains 2 genera and 10 species, which possess two dorsal fins and no barbels on the snout. Some authors consider that Gaidropsarus possesses two dorsal fins, but others present convincing X-ray evidence that it possesses three. The species of this family are demersal or benthopelagic. In 1987 the world landings of the four most economically important species of this family (Urophycis tenuis, U. chuss, Phycis blennoides, and U. brasiliensis in decreasing order of importance) amounted to 35,933 t. None of the four species occurring in Canadian Arctic waters is utilized locally.



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Enchelyopus cimbrius

common names: A local name is Firtrådet Havkvabbe (Danish/

Greenlandic).

taxonomy:  The genus comes from the Greek enchelys (eel), and

the species name from the Latin cimbrius (Welsh). This species belongs to the subfamily Gaidropsarinae. It was placed in the Gadidae but now belongs to the Phycidae. Originally described under the genus Gadus, it has also been placed at various times in the genera Rhinonemus (a junior synonym of Enchelyopus), Onos, and Gaidropsarus. Motella caudacuta Storer, 1848, described from off Massachusetts, and Couchia edwardii Couch, 1866, described from off England, are synonyms. Motella pacifica Temminck and Schlegel, 1846, has been suggested as a synonym, but the only specimen known came from Japan.

description:  This species is distinguished by the first dorsal fin being an elongate ray; the second dorsal fin has very short rays, each similar in size, hair-like, and inconspicuous; the first and second pelvic rays are not markedly elongate; snout barbels are present; and the end of the dorsal and anal fins and the lower caudal fin lobe have black pigment blotches. The upper jaw projects beyond the lower jaw. The snout is blunt. Small teeth are present in the upper (premaxillae) and lower (dentaries) jaws, and there are six to eight larger teeth in the anterior part of the upper jaw. There are teeth on the head of the vomer (roof of the mouth). A chin barbel is present, and there is a small barbel on the tip of the snout, and a larger one at each of the two anterior nostrils. The first dorsal fin consists of an elongate ray that is unsegmented and unpaired (a pseudospine). The second dorsal fin consists of a series of about 50 small, fleshy, unsegmented but bilaterally symmetrical, free-standing filaments that lie in a groove. The

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Enchelyopus cimbrius

third dorsal fin is long and unindented, with 37–55 regular rays. The single anal fin is long and unindented, with 36–49 rays. The caudal fin is rounded or truncate, with rounded lobes. The pelvic fins are anterior to the pectoral fins and have 5–7 rays, the second of which is somewhat elongate. The pectoral fins have 15–19 rays. The lateral line is interrupted along its entire length and has 51–57 scales. Small cycloid scales cover the body. Total outer gill rakers on the first arch number 5–13. There are 7, rarely 6, branchiostegal rays. Total vertebrae (including the urostyle) number 50–56. The body colour varies from yellowish olive to dusky brown on the dorsal aspect and is paler on the lateral and ventral aspects, with small brown spots restricted to the lateral aspects. There is a long dark blotch along the ventral edge of the caudal fin and the posterior portion of the anal fin. The third dorsal fin has either one dark blotch at the posterior end or four or five dark blotches – one each at the anterior and posterior ends and two or three in the middle area. The interior of the mouth is dark purple or blue. The species attains 42.0 cm in total length, usually 20–35 cm.

years, 25 cm at seven years, and 29–30.5 cm at nine years. The maximum age is nine years based on otoliths. The total length at first maturity is about 15 cm and is attained at three years of age. Along the east coast of North America spawning extends from April to November, with the peak being in June at water temperatures of 9°C–10°C. Is also known to spawn in the Baltic Sea and around the southwest coast of Iceland. Spawning occurs at depths of less than 140 m. Fecundity is 5,000–45,000 eggs per female. The eggs are spherical, possess an oil globule, and are 0.65– 0.86 mm in diameter, buoyant, and transparent. Larvae are 1.6–2.4 mm at hatching. It is parasitized by protozoans, nematodes, trematodes, and a cestode.

habitat:  This species is demersal as an adult and pelagic as a larva

Hudson Strait as three records from cruise data. It is also found in southwest and southeast Greenland and on both the North American and European sides of the North Atlantic Ocean. In the western North Atlantic it occurs south to the northern Gulf of Mexico. In the eastern North Atlantic there is a lone record off Mauritania (21° N), and then the distribution occurs without interruption from the northern Bay of Biscay up to the Baltic Sea into the Gulf of Finland and to the Barents Sea up to about 71° N. It also occurs around Iceland.

at depths ranging from 0 m to 650 m (exceptionally down to 1,325 m), but it is usually found between 20 m and 55 m. Arctic Canadian fish have been found at 143–672 m. It is associated with soft muddy-sand or gravel substrates. In soft muddy-sand substrates it stays in burrows during the day and forages at night. Generally it is sedentary but is thought to undertake short migrations to shallow waters in autumn and winter and return to deeper waters in spring and summer.

biology:  It feeds on young flatfishes, small crustaceans (amph-

importance:  The species is not very important because it is not

very abundant. It is taken in trawls. In Russia it is smoked and used in soups.

distribution:  It has been found in Davis Strait and western

ipods, copepods, decapods, isopods, mysids), polychaetes, and bivalve molluscs. The food may be taken by vigorously disturbing bottom mud and gulping in any organisms that appear or can be swallowed with the mud. Sea Ravens and White Hakes are the dominant predators. Larvae that are 17–20 mm long are silvery and pelagic. Those that are 35–48 mm long begin to descend to the bottom. Growth is slow: 15 cm in total length is attained at three years, 20 cm at five

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taxonomy:  The genus comes from the Greek gaia (earth), hydor (water), and psaros (speckled like a starling). The species name comes from the Latin argentum (silver) and atus (provided with). The species belongs to the subfamily Gaidropsarinae. It was placed in the Gadidae but now belongs to the Phycidae. Originally described under the genus Motella, it has also been placed in the genera Onos and Onogadus. Motella reinhardi Collett, 1879, described from the Barents Sea off Bear Island, is a synonym.

Distribution of Enchelyopus cimbrius

sources:  See the bibliography.

Gaidropsarus argentatus (Reinhardt, 1837)

Silver Rockling, mustèle argentée

common names:  Local names are Saviliursak (Greenlandic)

and Korstrålet Arktisk Havkvabbe (Danish/Greenlandic). Other common names are Arctic Rockling and motelle arctique.

description:  This species is distinguished by the first dorsal fin being an elongate but relatively short ray (not reaching the third dorsal fin); the second dorsal fin has very short rays, each similar in size, hair-like, and inconspicuous; the first and second pelvic rays are not markedly elongate; snout barbels are present; there are no black blotches at the end of the dorsal and anal fins and the lower caudal fin lobe; and the interorbital width is less than the eye diameter. The upper jaw projects beyond the lower jaw. Numerous sharp teeth are present on the upper (premaxillae) and lower (dentaries) jaws and on the head of the vomer (roof of the mouth). Two barbels are present on the snout, and one barbel on the chin. The head length measures 20%–25% of total length. The interorbital width is smaller than the eye diameter. The first dorsal fin lies in a groove and consists of one elongate and thickened unsegmented ray (a pseudospine). The second dorsal fin lies in the same groove and consists of a series of short, fine, unsegmented rays that are not connected by a membrane. The third dorsal fin consists of 52–65 longer, similarly sized, segmented rays that are connected by a membrane. The elongate ray of the first dorsal fin measures 31%–43% of head length and 3.5%–8.8% of total length. There is one anal fin, with 43–51 similarly sized rays. The pectoral fins have 22–24 rays. The pelvic fins have 7–9 rays, the second of which is moderately elongate. Total outer gill rakers on the first arch number 8–11 and are short and stubby in adults. Total vertebrae (including the urostyle) number 51–53. Lateral-line pores are present on the head. The lateral line is interrupted along its entire length and has a sharp bend downward at the origin of the anal fin. The body colour is red, reddish brown, or dark grey, and the head and belly are bluish grey. Immature individuals are silvery in colour, hence the origin of the scientific and common names. The edges of the dorsal, anal, and pectoral fin membranes

Gaidropsarus argentatus



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Gaidropsarus argentatus

are orange to red. The buccal cavity is light. The species attains 45.0 cm in total length.

habitat:  The species is epibenthic, occurring offshore at depths

from 100 m to 2,260 m (usually > 400 m) at water temperatures of 0°C–4.1°C. It has been caught at 682–1,389 m and 0.05°C–1.23°C in northern Davis Strait and at 437–948 m in southern Davis Strait, at 545–650 m and 1.6°C–2.9°C in Davis Strait generally, and at 442.5– 1,412.5 m and 0.1°C–4.1°C in southern Baffin Bay and Davis Strait generally. This species is closely associated to bottoms of gravel, sand, mud, shells, and stones.

biology:  It feeds on crustaceans (amphipods, decapods, and euphausiids) and sometimes fishes. Spawning occurs in Greenland waters.

importance:  It is not economically important. distribution:  The species is found in Baffin Bay, Davis and Hudson Straits, and northern Ungava Bay as well as all the coasts of Greenland. Generally it is found on both sides of the North Atlantic Ocean. In the eastern North Atlantic it occurs off Iceland, the Faroe Islands, and Norway. In the western North Atlantic it occurs on the Greenland side of Baffin Bay up to Upernavik (72°55' N), on the Canadian side (up to 74°23' N), and southwards along Labrador to the Grand Banks.

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Distribution of Gaidropsarus argentatus

sources:  Treble et al. (2000); Jørgensen et al. (2005).

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Gaidropsarus ensis (Reinhardt, 1837)

Threebeard Rockling, mustèle arctique à trois barbillons

common names:  A local name is Langstrålet Arktisk Havkvabbe (Danish/Greenlandic). Other common names are Arctic Threebeard Rockling and Threadfin Rockling. taxonomy:  The species name is the Latin ensis (sword).

The species belongs to the subfamily Gaidropsarinae. It was placed in the Gadidae but now belongs to the Phycidae. Originally described under the genus Motella based on material from off western Greenland, it has also been placed in the genera Onos and Onogadus. Onos rufus Gill, 1883, described from off New Jersey, is a synonym.

description:  This species is distinguished by the first dorsal fin-

ray being relatively long, reaching the third dorsal fin; the second dorsal fin has very short rays, similar in size, hair-like, and inconspicuous; snout barbels are present; there are no black blotches at the end of the dorsal and anal fins and lower caudal fin lobe; and the interorbital width is equal to the eye diameter. The upper jaw projects beyond the lower jaw. Numerous sharp teeth are present on the upper (premaxillae) and lower (dentaries) jaws and on the head of the vomer (roof of the mouth). Two barbels are present on the snout, and one on the chin. The head length measures 18%–20% of the total length. The interorbital width is about equal to the eye diameter. The first dorsal fin lies in a groove and consists of one elongate and thickened unsegmented ray (a pseudospine). The second dorsal fin lies in the same groove and consists of a series of short, fine, unsegmented rays that are not connected by a membrane. The third dorsal fin consists of 52–64 longer, similarly sized, segmented rays that are connected by a membrane.

The elongate ray of the first dorsal fin measures 86%–114% of head length and 11.3%–25.9% of total length. The anal fin has 40–48 similarly sized rays. The pectoral fin-rays number 20–27. The pelvic fin-rays number 6–9, the second of which is elongate. Total outer gill rakers on the first arch number 11–13 and are elongate and slender in adults. The vertebrae (including the urostyle) number 52–53. Lateral-line pores are present on the head. The lateral line is interrupted along its entire length and has a sharp bend downward at the origin of the anal fin. The body colour is light brown or red on the dorsal and lateral aspects, and the belly is reddish with a bluish-grey tinge. The species attains 42.0 cm in total length.

habitat:  This species is epibenthic, occurring at depths of 229–

2,020 m at water temperatures of just below 0°C to 4.4°C. It is found at 235–649 m in Ungava Bay for 44 collections, 398–1,600 m at −0.08°C to 2.57°C for 59 collections from Davis Strait and Baffin Bay, and at 266–530 m in Hudson Strait for 32 collections. It has been caught at up to 4.4°C generally in southern Baffin Bay and Davis Strait.

biology:  It is believed to feed on epibenthic crustaceans. Hooded

Seals are known to eat them (the original description of the species was based on two specimens removed from the stomach of a Hooded Seal taken in western Greenland at Uummannaq). It spawns in Greenland waters.

importance:  On the Canadian side of Baffin Bay it is collected as a by-catch of the Greenland Halibut fishery.

distribution:  The species is found in Baffin Bay (to 74°18' N), Davis and Hudson Straits, the entrance to Cumberland Sound, and northern Ungava Bay. It is found also on all the coasts of Greenland and generally in the northwest Atlantic Ocean, south through Labrador, the Flemish Cap, the Grand Banks, and along the Scotian Shelf to Cape Hatteras (35° N).

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taxonomy:  The genus comes from the Greek phykos (seaweed), in reference to the habitat in which these fishes hide. The species name honours Captain Hubbard C. Chester from Noank, Connecticut. The species belongs to the subfamily Phycinae. It was originally described under the genus Phycis, then placed in the genus Urophycis, but now returned to Phycis.

Distribution of Gaidropsarus ensis

sources:  Kotthaus & Krefft (1957); Allard (1980); Hudon (1990a); D.B. Stewart et al. (1991); Treble et al. (2000); Jørgensen et al. (2005).

Phycis chesteri

Goode and Bean, 1878 Longfin Hake, merluche à longues nageoires

description:  This species is distinguished by having two dorsal fins, the rays of the first dorsal fin, after the very elongate third ray, being progressively shorter but clearly visible; the first and second pelvic fin-rays are greatly elongate (the second almost to the caudal fin); and barbels are absent from the snout. The upper jaw projects beyond the lower jaw. The mouth is large, with fine teeth on the premaxillaries (upper jaw), the dentaries (lower jaw), and the head of the vomer (roof of the mouth). A chin barbel is present. The two dorsal fins touch at the base. The first dorsal fin is short, with 8–12 rays, the third of these being extremely elongate, much longer than the head length. The second dorsal fin is long and slightly indented, with 50–63 rays. A single, long, and slightly indented anal fin has 43–56 rays. The caudal fin is rounded. The second dorsal fin and the anal fin are not connected to the caudal fin. The pectoral fin-rays number 14–18. The pelvic fins are jugular (anterior to the pectoral fins), each consisting of two extremely elongate rays and a third rudimentary ray. The second elongate pelvic fin-ray is the longest and reaches the end (insertion) of the anal fin or beyond. The branchiostegal rays number 7. Total outer gill rakers on the first arch number 17–24, and the epibranchial gill rakers 4–5. The pyloric caeca number 11–14. The vertebrae (including the urostyle) number 45–52. Lateral-line pores are present on the head. The scales are cycloid, and the lateral-line scales number 90–91. The first half of the lateral line is slightly arched above the midline and is continuous, and the other half is straight, is at the level of the midline, and becomes interrupted at about the insertion of the anal fin. The body colour is olive brown to olive on the back, lighter on the sides, and silvery white on the underside. The margins of the dorsal, caudal, and anal fins are darker. The species reaches 45.0 cm in total length. habitat:  The species is benthopelagic at depths from 60 m to

1,400 m but usually occurs between 300 m and 900 m over soft, silty substrates. It occurs at temperatures of 1.6°C–13.0°C, but mostly at 4°C–9°C. The larvae and juveniles are pelagic, and when the latter reach a standard length of 14 cm, they become demersal. In Davis Strait and Baffin Bay they have been found at depths of 642–1,130 m.

biology:  Its preferred food is crustaceans (amphipods, euphausPhycis chesteri

common names:  A local name is Langstrålet Skælbrosme

(Danish/Greenlandic).

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iids, and shrimps), but it also feeds on polychaetes, molluscs (bivalves and cephalopods), and vertically migrating fishes (Lanternfishes, Marine Hatchetfishes, and Bristlemouths). They are preyed on by Spiny Dogfish, White Hake, and Goosefish. Aging has been attempted using otoliths, but this has proved unreliable. The females outnumber the males by almost three to

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Phycis chesteri

one. The females are larger than the males, and maturity is attained at 22.4 cm for males and 29.2 cm for females in U.S. waters. Females of 24–41 cm in total length produce 450,000–1,305,700 eggs. The transparent, buoyant, and spherical fertilized eggs measure 0.8 mm in diameter and possess an oil globule. Across the species’ range, spawning occurs from the end of September to April, with peak activity in December to January. Swimming Longfin Hake extend their pelvic fins forward, dragging the tips of the elongated rays along the substrate. Parasites include cestodes, nematodes, trematodes, and an acanthocephalan.

importance:  It has no commercial importance at present. How-

ever, its flesh is white and flaky, and despite its being rather soft, the abundance of the fish at depths between 500 m and 700 m make it a potential candidate for a trawling fishery.

distribution:  The species has been found in southern Baffin Bay as a single record at 70º40' N and in Davis and Hudson Straits, as well as in southwest and southeast Greenland. It is found generally in the northwest Atlantic Ocean from Baffin Bay south to the Straits of Florida but is not abundant south of Cape Hatteras. Distribution of Phycis chesteri

sources:  Karrer (1973); Allard (1980); Wenner (1983); Methven (1985); Hudon (1990a).



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Family Gadidae Cods, Morues

Claude B. Renaud

This family, also known as the Codfishes, comprises two subfamilies (Gadinae and Lotinae; both are treated as families in an alternative classification), 15 genera, and 26 species. There are 17 species in Canada, of which 8 are in the Arctic. Some authors also include the subfamily Phycinae within the Gadidae, but it is treated as a family here. Members of the family vary in adult total length between 15 cm and 200 cm, are typically marine – except for the freshwater Burbot, Lota lota; and for some freshwater populations of Atlantic Tomcod, Microgadus tomcod (Walbaum, 1792) – and are mostly found in the polar and temperate continental-shelf waters of the northern hemisphere. Only one species is found in the tropical and temperate waters of the southern hemisphere, Micromesistius australis Norman, 1937. The main characteristics of the family are gill openings that extend above the level of the pectoral fins; the gas bladder has anterior horn-like processes that are not connected to the auditory capsules at the rear of the skull (no otophysic connection), and it is not connected to the oesophagus (physoclistous); the neural spine of the first vertebra is firmly attached to the rear of the skull; small cycloid scales are present; the fins have soft rays only; the head of the vomer (roof of the mouth) has teeth; and a chin barbel is usually present. The subfamily Gadinae contains 12 genera and 21 species, which possess three dorsal fins, two anal fins, a truncate or slightly forked caudal fin, and eggs without an oil globule. The subfamily Lotinae contains 3 genera and 5 species, which possess one or two dorsal fins, one anal fin, a rounded caudal fin, and eggs with an oil globule. Most are benthopelagic, except for the predominantly pelagic Arctogadus, Boreogadus, Gadiculus, and Micromesistius species. They are often found in large schools. The eggs and larvae are usually pelagic. A long pelagic life and spawning and feeding migrations result in the wide distribution of many species. Adults feed on fishes and various invertebrates. At least four of the species found in the Canadian Arctic, Boreogadus saida, Eleginus gracilis, Gadus

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morhua, and G. ogac, are specially adapted against freezing through the possession of glycoproteins in their bloodstream. Additionally, Arctogadus glacialis, B. saida, and G. morhua possess multiple forms of haemoglobin, enabling them to bind oxygen over a wide range of temperatures. The family is second only to the Herrings (Clupeidae) for world catch of marine fishes, with landings of 11,581,872 t in 1987. Five of the eight species that occur in Canadian Arctic marine waters (A. glacialis, B. saida, E. gracilis, G. morhua, and G. ogac) are exploited locally to some level. A sixth species, Lota lota, is also exploited but only in fresh waters because its occurrence in marine waters is very sporadic. Overfishing continues to be a problem, and the collapse of the Atlantic Cod stocks is infamous. Canada was practically founded on the Atlantic Cod fisheries, the Grand Banks stocks being fished by Europeans before the voyage of John Cabot in 1497 to these waters. In Arctic waters Boreogadus saida has an extensive scientific literature because it is a keystone species. This species transfers up to 75% of the energy from lower trophic levels to top predators.

sources: Vladykov (1933a); Svetovidov (1948); Milne & Smiley

(1976, 1978); Van Voorhies, Raymond, & DeVries (1978); Markle (1982); Dewar, Johnson, Layton, & Marshall (1983); Burcham, Osuga, Rao, Bush, & Feeney (1986); Burcham, Osuga, Yeh, & Feeney (1986); Cohen (1989); Crawford (1989); Cohen et al. (1990); Fabijan (1991a, 1991c); D.B. Stewart et al. (1991); Marshall & Layton (1997); Marshall (1999a); Endo (2002); Møller, Jordan, Gravlund, & Steffensen (2002); Geoffroy, Majewski, Reist, & Fortier (2012); Verde, Giordano, di Prisco, & Andersen (2012).

Arctogadus glacialis (Peters, 1872)

Polar Cod, saïda imberbe

common names: Local names are Uugavik (Inuvialuktun) and

Istorsk (Danish/Greenlandic). Other common names are Arctic Cod (Europe), Arctic Greenland Cod, East Siberian Cod, Ice Cod, Toothed Cod, morue arctique, and saïda barbu. The use of different common names in the European and North American literature can lead to some confusion. Europeans refer to this species as Arctic Cod, whereas North Americans officially call it Polar Cod. The problem lies in that, for Europeans, Polar Cod refers to Boreogadus saida (see that species account), while North Americans officially refer to B. saida as Arctic Cod. Additionally, when Arctogadus borisovi was synonymized with A. glacialis (see below under “Taxonomy”), their French common names referring to chin barbel development were no longer appropriate and could cause confusion because the length of the chin barbel is due to intraspecific variation. Therefore, a more appropriate name suggested is saïda polaire.

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Arctogadus glacialis

taxonomy: The genus comes from the Greek arktos (north) and gados (codfish), and the species name is the Latin glacialis (icy). The species belongs to the subfamily Gadinae and was originally described under the genus Gadus. The type locality is off Sabine Island, off the east coast of Greenland. Synonyms include Arctogadus borisovi Dryagin, 1932, described from the delta of the Kolyma River, Siberia, Russia; Boreogadus pearyi Nichols and Maxwell, 1933, described from Lincoln Bay, Nunavut (82°07' N, 61°50' W, not northern Greenland as often stated), and Phocaegadus megalops Jensen, 1948, described from Baffin Bay, off the west coast of Greenland. It would appear that Gadus (Boreogadus) Bottemannei Bleeker, 1873, described from the Iceland Sea, is also a synonym. description: This species is distinguished by having three dorsal

fins and two anal fins (about equal in size); its jaws are about equal in length or the lower jaw protrudes slightly; the lateral line is not continuous but is interrupted along its entire length; and the palatine bones in the roof of the mouth have teeth. Teeth are present on the premaxillaries (upper jaw), dentaries (lower jaw), and the head of the vomer (the roof of the mouth). Palatine teeth are almost invariably present (97%) with 3–17 teeth per bone. The chin barbel varies in development, from being absent to being relatively small to being well developed, and its length from 0.0 mm to 15.7 mm, representing 0.0%–15.5% of head length. The eye is relatively large, its diameter being 4.9%–11.1% of standard length. The interorbital width is 4.4%–7.0% of standard length. The individual dorsal fins and the anal fins are all widely separated from each other. The first dorsal fin has 9–16 rays, the second dorsal fin 14–24 rays, and the third dorsal fin 16–25 rays. The first anal fin has 15–25 rays, and the second anal fin 15–26 rays. The caudal fin is truncate, having rounded lobes or being forked with rounded lobes. The pelvic fins are jugular (anterior to the pectoral fins) with 6 rays, the second one being elongate. The pectoral fins have 16–23 rays. The branchiostegal rays number 7. Total outer gill rakers on the first arch



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number 26–38. The pyloric caeca number 30–47. Total vertebrae (including the urostyle) number 54–61. The lateral line is interrupted along its full length and has a sharp curve downwards under the second dorsal fin, after which it continues straight either at or under the midline to the base of the caudal fin. The cycloid scales on the body overlap like shingles on a roof. Pit organs instead of pores are present on the head. The body coloration (live) is uniformly dark olive, brown, bluish, or yellowish on the dorsal and upper lateral aspects, grey on the lower lateral aspect, and light grey with dark speckling on the ventral aspect. The young are a light grey, with the head and fins being chocolate brown. The ventral aspect of young individuals is often silvery. The fins are black or dark grey. The peritoneum is dark brown to black. The species reaches 60.0 cm in total length and 1.5 kg in weight.

habitat: This is a pelagic, relatively sedentary, schooling spe-

cies, associated with offshore pack ice or drifting ice, at or beyond the edge of the continental shelf, but usually close to mud and shale bottoms in coastal waters. It is often found near or in beds of macroalgae such as Laminaria spp. and Fucus spp., and it has been found in a medusa at 253 m. It may sometimes be collected in river estuaries or in fiords. It is found at depths of 0–930 m, temperatures of −1.6°C to 3.4°C, usually below 0°C, and salinities of up to 33.5‰, but it can also tolerate the low salinities of estuaries. It has been collected at 191–1,439 m in Baffin Bay and Davis Strait. It dominates in the otter trawl catches of Bridport Inlet, Melville Island.

biology: The diet includes fishes such as Arctic Cod and mysids,

but it comprises predominantly pelagic amphipods associated with the underside of sea ice. It is eaten by Narwhal and Bearded, Harp, Hooded, and Ringed Seals. Fishes of 52–68 mm in fork length are one year old, based on otoliths. The species reaches 13 years of age, based on otoliths. Females mature at three to four years. Spawning is assumed to take place during summer and in sea water.

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importance: It is fished in Norway using trawls at depths

between 230 m and 930 m in the northeastern part of the East Siberian Sea and off both coasts of Greenland. It has little importance as food but is used as fish-meal and oil. In the western and north-central coastal Canadian Arctic mainland and archipelago this species is caught in the subsistence fishery by jigging in ice cracks.

Boreogadus saida

distribution: It is found throughout Arctic Canada including

Equaluaq, Itok, Ogac, Ogaq, Ôgark, Ogâ tsuk, Ovac, and Uugavik (Inuktitut); and Uugaq (Inuvialuktun); in Greenland they are Ekalluak, Ekalugak, Eqalugaq, and Ûvak. Other common names are Polar Cod (Europe), Rock Cod (see the “Common Names” section under Arctogadus glacialis), and morue arctique. Some older literature uses the English common name Tomcod, but this name is now restricted to members of the genus Microgadus, which are found outside the Arctic.

the Arctic islands, but not in western Hudson Strait, Hudson and James Bays, and the Foxe Basin. A record from the Daly Bay area of northwestern Hudson Bay in Stewart and Bernier (1984) requires confirmation by voucher specimens. It is found also in northwest, southeast, and northeast Greenland and off Iceland. It occurs across the northern Barents Sea; along the coast of Russia in the Kara, Laptev, and East Siberian Seas and offshore up to about 81° N; and along the northern coast of Alaska. It is usually found above the Arctic Circle except in Davis Strait, eastern Hudson Strait, Ungava Bay, and the southern tip of Greenland.

(Lepechin, 1774)

Arctic Cod, saïda franc

common names: Local names in Canada and Alaska are

taxonomy: The genus comes from the Greek boreas (north) and gados (codfish), and the species name is the Russian for this fish. The species belongs to the subfamily Gadinae and was originally described under the genus Gadus. Synonyms include Merlangus polaris Sabine, 1824, described from Baffin Bay; Gadus fabricii Richardson, 1836, described from the northern bays of Greenland; and Gadus agilis Reinhardt, 1837, also described from Greenland. description: This species is distinguished by having three dorsal

Distribution of Arctogadus glacialis

sources: Walters (1955, 1961); Nielsen & Jensen (1967); Buchanan,

Cross, & Thomson (1977); Hunter (1981); Finley & Gibb (1982b); Finley & Evans (1983); Strong (1988); Andersen, Nielsen, & Smidt (1994); Graham, Johnson, Stenton, Welch, & Welch (1996); Treble et al. (2000); Jordan, Møller, & Nielsen (2003); Jørgensen et al. (2005); Aschan et al. (2009); Michel et al. (2012); Naughton (2012).

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fins, two anal fins (about equal in size), 35–47 gill rakers, no palatine teeth, and a very wavy lateral line. The jaws are equal or the lower jaw projects slightly beyond the upper jaw (the mouth is terminal or slightly upturned). Teeth are present on the premaxillaries (upper jaw), dentaries (lower jaw), and the head of the vomer (roof of the mouth), but not on the palatines (also roof of the mouth). A minute chin barbel is present and is much shorter than the pupil diameter. The head length is about 22% of total length. Individual dorsal fins and the anal fins are widely separated from each other. The first dorsal fin has 10–17 rays, the second dorsal fin 11–18 rays, and the third dorsal fin 16–24 rays. The first anal fin has 13–22 rays, and the second anal fin 17–24 rays. The caudal peduncle is slender. The caudal fin is forked, with rounded upper and lower lobes. The pelvic fins are jugular (anterior to the pectoral fins), with 6 rays each, the second ones being elongate. The pectoral finrays number 18–21. The branchiostegal rays number 6–7. The pyloric caeca number 20–37. The vertebrae (including the urostyle) number 49–58. Pit organs instead of pores are present on the head. The lateral line is interrupted along its full length and becomes wavy under the second dorsal fin and posteriorly. The scales are cycloid, small, and not overlapping, but scattered. Individuals of more than 25 cm in total length have bony roughening structures (contact organs), one per scale, which erupt from under their scales. The body colour is brownish to black on the upper surface and silvery on the lateral and lower surfaces, with dark speckling scattered over the body. A violet or yellowish sheen may be present. The dorsal, anal, and caudal fins are dark with pale edges. The peritoneum is dark. The species attains a total length of 40.0 cm but is usually about 25.0 cm.

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Boreogadus saida

habitat: The species is pelagic, associated with a wide variety of

habitats from ice-free, nearshore marine waters to brackish lagoons to almost freshwater river mouths (the estuaries of Rupert Bay, of Eastmain and La Grande Rivers flowing into eastern James Bay, and of Great Whale, Little Whale, and Innuksuac Rivers flowing into eastern Hudson Bay), to 175 km offshore, to under smooth sea ice or in ice cracks, in narrow wedges along the edges of melting ice floes, in surface channels on top of ice in meltwater, and under jellyfish in Resolute Bay. It is often associated with ice, either pack or drifting, that acts as an “inverted benthos.” Pack ice also provides protection from seabird and marine mammal predators, and the fish are more dispersed than when there is no ice cover. Wedges may harbour up to 28 resting fish, perhaps a strategy to reduce energetic requirements and avoid predators. Concentrations of the dense benthopelagic layer of fish reach 37 g/sq m in the southeast Beaufort Sea, and total biomass in the Amundsen Gulf reaches about 250,000  t. These large shoals are governed by local currents that concentrate and trap zooplankton prey. Smaller fish (less than a year old) are found in an epipelagic layer to about 60 m depths in the southeast Beaufort Sea. Larger individuals (age 1+) have been found at more than 200 m there during the ice-free season over the continental slope, similar to their winter distribution, and accounting for the “missing” biomass in nearshore waters. The biomass in nearshore waters has been found to be too low to support the energetic requirements of known predators. In winter in the Amundsen Gulf, aggregations migrate progressively deeper, from 220 m to 550 m bottom depths, in response to increasing light intensity, presumably a mechanism to avoid visual predators such as the Ringed Seal. Fish that became trapped in bays, such as Franklin Bay, during the winter provide a dense concentration accessible to the diving range of marine mammals. However, small cod (< 25 g) in Franklin Bay migrate into a cold layer (90–150 m, −1.4°C) at night from December to April to avoid being captured by shallow-diving and sight-feeding immature seals. Large cod (25–95 g) stayed below 180 m, presumably to avoid



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deep-diving mature seals. The diel vertical migration of small cod has been attributed to the lack of prey and to the foraging interference by larger cod in deeper water. A bioluminescent copepod (Metridia longa) might provide light for the cod to feed on copepods in the deep, dark water. During the day cod of all sizes form a dense aggregation at 160–230 m and −1.0°C to 0°C. The species occurs at depths from the surface down to 1,390 m, but the larvae are most abundant between 5 m and 20 m depths. In shallow areas (< 25 m) with a rocky substrate it is found in association with kelps (Laminaria, Agarum, and Alaria spp.). This species is tolerant of wide variations in temperature (−1.85°C to 12.5°C), salinity (1‰–30‰), and turbidity. Its preferred temperature is below 3.5°C. It has been recorded at 146–1,383 m and −0.08°C to 2.3°C for 34 collections in Davis Strait and Baffin Bay, at 0–530 m for 178 collections in Hudson Strait, and at 115–620 m for 120 collections in Ungava Bay. It undergoes distant migrations for spawning.

biology: Its food consists mostly of planktonic and benthic crustaceans (amphipods, cirripeds, copepods, cumaceans, euphausiids, isopods, mysids, and shrimps) and, less often, fish eggs and fry, including at times their own; it will also eat phytoplankton, free-living tunicates, chaetognaths, opisthobranch gastropods, polychaetes, and squid. Larval Arctic Cod are generalists and feed on a wide variety of organisms but predominantly on copepod eggs and nauplii larvae. Adults feed at times on the bottom, as evidenced by the presence of pebbles in the gut. The Arctic Cod is termed a keystone species because of the central role it plays in the Arctic marine food web, providing the main link between phytoplankton and small pelagic crustaceans (amphipods and copepods), on the one hand, and other fishes (Arctic Char, Dolly Varden, Atlantic Cod, Atlantic Salmon, Fourhorn Sculpin, Shorthorn Sculpin, Greenland Cod, Greenland Halibut), seabirds (Arctic Tern, Northern Fulmar, Black-legged Kittiwake, Common and Thick-billed Murres, Black Guillemot, Glaucous Gull, Ivory Gull, Sabine’s Gull, Ross’s Gull, Thayer’s Gull, Parasitic Jaeger),

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marine mammals (Bearded, Harp, Hooded, Harbor, Ringed, and Spotted Seals, Beluga, Bowhead Whale, Narwhal, Walrus, Polar Bear), and terrestrial mammals (Arctic Fox, the Inuit, and their dogs), on the other. It has been estimated that 1–10 million Arctic Cod are eaten monthly along land-fast ice edges by seabirds alone. There are numerous anecdotal reports of its utilization as food by other species in the Arctic. One Arctic Cod was conveniently dropped by an Eider Duck at Cape Fullerton, Hudson Bay, and added to the Canadian Museum of Nature’s fish collection (CMNFI 19580169). It is fed on by Bowhead Whales in the Alaskan Beaufort Sea and by Atlantic Salmon at Killiniq (Port Burwell), Ungava Bay. At Allen Bay, Cornwallis Island, on 31 August 2010 during a run on shore, the fishes were eaten by Northern Fulmars, kittiwakes (presumably Black-legged Kittiwakes), Glaucous and Ivory Gulls, and Arctic Terns. Northern Fulmars and Thick-billed Murre chicks eat this species at Prince Leopold Island, Lancaster Sound. Ivory Gulls eat this cod in the North Water Polynya between Canada and Greenland. It is also eaten by Thick-billed Murre chicks at Coats Island; by Sabine’s Gull, Bearded Seals, Arctic Char, and Fourhorn Sculpins at Bernard Harbour; by Shorthorn Sculpins at Blacklead Island, Cumberland Sound; and by Bearded Seals, Harp Seals, Ringed Seals, and Belugas in southeastern Baffin Island. In Allen Bay, Cornwallis Island, the presence of schools was a significant indicator for the distribution of Northern Fulmars. Demersal schools there separate into satellite schools to feed at the surface, reducing competition but increasing the risk of predation. It is the main food of Narwhals in Lancaster Sound and a major food of

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Belugas at Resolute Bay, Creswell Bay, and Aujuittuq (Grise Fiord) in Jones Sound. Even Common Ravens eat Arctic Cod in the Canadian High Arctic, after the fish has been discarded by seabirds. It comprises about 40% of the diet of Ringed Seals in Amundsen Gulf, and about 29,000 t are eaten annually by Ringed Seals, Belugas, and marine birds on the Alaskan continental shelf. The movements of marine mammals and birds on the Canadian Beaufort shelves are influenced by this cod. The species occurs in large aggregations that may involve hundreds of millions of individuals, along the coast during summer, fall, and winter. The larvae of this cod are the most wide-spread and abundant fish larvae from the Chukchi Sea to Amundsen Gulf. On the southern coast of Cornwallis Island, Lancaster Sound, two such aggregations contained at least 900 million individuals, the schools had a surface area of 59 hectares, and total biomass was 30,715 t (520 t/ha). Large schools are a defensive measure against predators. Massive shore strandings numbering thousands of individuals sometimes occur after a storm event. It is known to leap onto ice to escape predators, notably Beluga. It is the most abundant and has the highest biomass of any fish in Arctic marine waters. For example, the number of Arctic Cod in Lancaster Sound has been estimated at 400,000 t. However, its abundance throughout the Arctic waters is not uniform, and the number estimated to occur in Mackenzie Bay is one order of magnitude less than that found in Lancaster Sound, and in Frobisher Bay it is intermediate between these two. Coastal fish grow at a faster rate than do offshore fish, reaching 6.0 cm in total length at the end of their first year, compared to 2.0

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cm in the latter case. The maximum age is 6 years in Labrador and up to 10 years in the Arctic. Females make up the majority of the older fish. Maturity is attained at 2–3 years for males and 3–4 years for females. The Arctic Cod is an r-selected species characterized by early maturity, rapid growth, larger numbers of offspring at a given parental size, small body size, most young being produced at an early age, high mortality, and short lifespan. Such fish are better adapted to an environment that is liable to catastrophic, unpredictable changes. It is of no selective advantage to invest in large, long-lived fish if adults are as liable as young to sudden disaster. Spawning under ice near the sea bed occurs from October to March, peaking in January and February. Females produce up to 21,000 buoyant eggs of 1.9 mm in diameter. Larval survival is limited at sea ice cover greater than 50% and sea surface temperature of less than 0°C. Therefore, summer cohorts have better survival than do spring cohorts, but the latter persist, presumably because they reach a larger size before the onset of the Arctic winter, which is a survival advantage. Hatching may start as early as January and last into July in seas that have a large river discharge, such as Hudson Bay and the Beaufort Sea, but is restricted to April–July where there is little freshwater input, such as the Canadian Arctic Archipelago and north Baffin Bay. Increased river discharges linked to climate change, more frequent winter polynyas, and a warmer surface layer all help young cod to grow to larger sizes and survive the oncoming winter. The species possesses a glycoprotein in its blood that prevents it from freezing. Parasites include nematodes of the genus Thynnascaris, digenetic trematodes of the genera Genolinea and Derogenes, and copepods belonging to the genera Clavella and Haemobaphes.

distribution: The species is found throughout Arctic Canada as far north as the northern tip of Ellesmere Island, and almost to the North Pole at 88º26' N, 126º26' E, off Russia; and on all coasts of Greenland. It occurs throughout the entire Arctic Ocean basin but does not usually occur south of 60° N except in the Bering Sea, Hudson and James Bays, Ungava Bay, the southern tip of Greenland, and along the coast of Labrador into the Gulf of St Lawrence (Miramichi Bay, New Brunswick, 47°08' N) and into Saguenay Fjord, Québec (48°25' N).

importance: In Canadian waters Arctic Cod is exploited in a

Distribution of Boreogadus saida

minor way by the Norwegians. As its flesh is of low quality, it is used mainly for fish-meal and liver oil. In 1984 the world catch totalled 23,709 t, and in 1987 that number had declined to 11,713 t. The types of gears used are bottom and mid-water trawls. The major fishing grounds are in the White and Barents Seas and the northwest Atlantic Ocean and involve Russia, Norway, Denmark, and Germany. In the western and north-central coastal Canadian Arctic and archipelago and along Baffin Island this species is caught in the subsistence fishery by jigging in ice cracks. For sport, children jig for it in spring at Igloolik through ice cracks. At Aujuittuq (Grise Fiord) beach-stranded Arctic Cod are also harvested for domestic use. The Inuit use it as food for themselves and their dogs. Around the Kent Peninsula it has been used for winter stores and for dog food. The cod is reputed by the Inuit to have an immortal soul that returns to the sea when the body has been eaten. Inuit used to arrange a fish catch on the ice in a circle, with their heads facing inwards to the fishing hole, and the fisherman standing next to the fishing hole, so that the cod would return to be caught again. The Inuvialuit harvest this species (about 300 fishes yearly) for domestic purposes out of Paulatuk, Northwest Territories. In Greenland they are caught as a by-catch of the shrimp fishery, and on the Canadian side of Baffin Bay as a by-catch of Greenland Halibut fisheries.



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sources: Ross (1826); Scofield (1899); Johansen (1926); Rasmussen (1927); Grainger (1953); Scholander, Flagg, Walters, & Irving (1953); Walters (1953c, 1955); McLaren (1958); Anders (1968); Drainville (1970); Mansfield, Smith, & Beck (1975); McAllister (1977); Craig & Griffiths (1978); den Beste & McCart (1978); Sergeant & Hay (1978, 1979); MacLaren Marex Inc. (1979b); Sandeman & Buchanan (1979); T.G. Smith, Hammill, Doidge, Cartier, & Sleno (1979); Lowry & Burns (1980); Morin (1980); Hunter (1981); Finley & Gibb (1982b); Finley & Evans (1983); Ratynski (1983); Sameoto (1984); Bradstreet et al. (1986); Chumakov & Podrazhanskaya (1986); Allen & Smith (1988); Strong (1988); Hudon (1990a); Fabijan (1991a); Gaston (1991); Gilbert (1991); D.B. Stewart et al. (1991); Renaud & Morrison (1992); Crawford & Jørgensen (1993a, 1996); Andersen et al. (1994); Graham et al. (1996); Treble et al. (2000); Gradinger & Bluhm (2004); Fortier, Sirois, Michaud, & Barber (2006); Karnovsky, Hobson, Brown, & Hunt (2009); Benoit, Simard, Gagné, Geoffroy, & Fortier (2010); Bouchard & Fortier (2011); Geoffroy, Robert, Darnis, & Fortier (2011); Walkusz, Paulic, Williams, Kwasniewski, & Papst (2011); Geoffroy, Majewski, Reist, & Fortier (2012); Matley, Crawford, & Dick (2012a, 2012b); Michel et al. (2012); Naughton (2012); Mathias (2013); Benoit, Simard, & Fortier (2014).

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Brosme brosme (Ascanius, 1772) Cusk, brosme

common names: Local names are Iloruleqqortooq and Tinguttooq (Greenlandic). Other common names are Scrod, Torske, and Tusk. taxonomy: The genus and species name either comes from the

Greek brosis (eat), presumably in reference to its voracious appetite, or refers to the Danish name brosme for the fish. It belongs to the subfamily Lotinae. Originally described under the genus Gadus, it has also been placed in the genus Trisopterus. The type locality is Denmark. Synonyms include Gadus torsk Bonnaterre, 1788, described from Norway; Gadus lubb Euphrasen, 1794, described from off Bohuslän, Sweden; Brosmius flavescens Lesueur, 1819, described from the Newfoundland Banks; and Brosmus vulgaris Fleming, 1828, described from the British Isles.

description: This species is distinguished by having one dorsal

fin and one anal fin, both joined or nearly joined to the caudal fin, and each separated from it by a notch. The upper jaw projects slightly beyond the lower jaw, or the jaws are equal. The mouth is large with rows of sharp teeth on the upper jaw (premaxillae), the lower jaw (dentaries), and the roof of the mouth (the head of the vomer and the palatines). The chin barbel is equal to or longer than the eye diameter. The caudal fin is rounded. The dorsal fin-rays number 85–108, and the anal finrays 62–77. The pelvic fins are small, with 5–6 equal rays, and their tips are free. The pectoral fin-rays number 22–25. The branchiostegal rays number 7. Total outer gill rakers on the first arch number 11–13. The pyloric caeca number about 15. Total vertebrae (including the urostyle) number 63–67. The scales are small and cycloid on the head and the rest of the body and are deeply embedded. Lateral-line pores are present on the head. The lateral line is arched over the pectoral fin and is continuous anteriorly but

becomes interrupted in the posterior 15% of its length. Drumming muscles are present on the gas bladder. The body colour is dark reddish or greenish brown on the upper aspect and cream or white on the lower aspect. Young individuals may have six yellow bars on the side. The unpaired fins are the same colour as the body at their base, have a black line near the margin, and are white at the margin. The species attains 120.0 cm in total length, but usually 40–60 cm, and 30.0 kg in weight.

habitat: The species is benthopelagic and usually found off-

shore on hard, rough bottoms of boulders, rocks, gravel, or pebbles. It occurs at depths between 18 m and 1,185 m, but usually above 550 m, at temperatures of 0°C–14°C, usually 6°C–10°C, and salinities of 32‰–34‰. It is solitary or found in small groups. It appears to be fairly sedentary, being sluggish, and is a weak swimmer.

biology: In the northeast Atlantic Ocean it feeds on crustaceans

(especially crabs and pandalid shrimps), molluscs, benthic fishes (such as flatfishes, gurnards, and Sculpins), and occasionally on starfish. Its stomach contents are often lost when the fish is brought to the surface because the stomach is everted due to the expansion of the gas bladder. It is prey to Spiny Dogfish, Atlantic Cod, Goosefish, Greenland Halibut, and other fishes and to Hooded Seal and Gray Seal. Individuals reach 34–45 cm in total length at five years of age, 53–59 cm in total length at seven years, 62–63 cm in total length at nine years, and 69–72 cm in total length at eleven years. Longevity is up to 21 years, based on otoliths, but some authors believe this is an under-estimate. Sexual maturity is reached at 43–50 cm in total length or at about five years in males and six years in females, but the ages have been questioned, and maturity may be reached at about age ten years. In the Gulf of Maine and on the Scotian Shelf spawning takes place between March and November, with the peak in late June, at depths of less than 183 m. Fecundity ranges from 100,000 to 3,927,000 eggs for females between 56 cm and 90 cm in total length, respectively. The fertilized eggs are 1.1–1.5 mm in diameter, spherical,

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and buoyant and have a coppery tint and a pinkish oil globule. Larvae hatch at 4 mm in total length and remain pelagic until 50 mm in total length, when they go to the bottom. It is parasitized by the nematode Pseudoterranova decipiens and the trematode Prosorhynchus squamatus.

importance: In the North Atlantic Ocean the 1987 landings

amounted to 46,254 t. The Cusk is taken as by-catch in the Greenland Halibut and Atlantic Cod fisheries. It is caught more often on long lines than in otter trawls. Its flesh is white, flaky, and of good flavour, supposedly like lobster. It is sold fresh, frozen as fillets, dried salted, in brine, or smoked. Canadian live-weight landings, mostly from the western portion of the Scotian Shelf, were 6,290 t in 1982, 3,150 t in 1984, 1,148 t in 2003, and 469 t in 2011. The decreased landings are due to the change from long-lining to otter trawling. The species is of no importance in Arctic Canada. The Committee on the Status of Endangered Wildlife in Canada has assessed it as “Endangered.”

distribution: The species has been found in Davis Strait with

four records from cruise data, and in southwest and southeast Greenland. It occurs on both sides of the North Atlantic Ocean between 37° and 83° N. In the northeast Atlantic Ocean it occurs off Iceland and the northern British Isles, and in the North, Norwegian, and Barents Seas. In the northwest Atlantic Ocean it occurs south from Davis Strait to the Gulf of St Lawrence, in the Strait of Belle Isle, on the Grand Banks of Newfoundland, and off New Jersey.

Eleginus gracilis (Tilesius, 1810)

Saffron Cod, navaga jaune

common names: Local names are Ogak, Ogavik, Siuryuktuuq,

and Uuqaq (Inuktitut). Other common names are Far Eastern Navaga and morue boréale. Some popular accounts use the common name Tomcod, but this is unfortunate because that name applies exclusively to members of the genus Microgadus, which are found outside the Arctic.

taxonomy: The genus comes from the Greek eleginos (a schooling fish mentioned by Aristotle), and the species name is the Latin gracilis (slender). The species belongs to the subfamily Gadinae. It was originally described under the genus Gadus and has been placed in the genus Pleurogadus. The species was treated as a subspecies of Eleginus navaga (Pallas, 1811), now known as Eleginus nawaga (Walbaum, 1792). The fishes in the American Arctic were also considered to be closer to the European subspecies navaga than to the Pacific subspecies gracilis. A taxonomic revision of the genus is required. Gadus wachna Pallas, 1814, described from Kamchatka, Russia, is a synonym. On the basis of molecular evidence, the genus Microgadus Gill, 1865, was considered by authors as a synonym of Eleginus Fischer, 1813, but they incorrectly attributed priority to Microgadus in violation of the International Code of Zoological Nomenclature. description: This species is distinguished by having three dorsal

Distribution of Brosme brosme

sources: Kotthaus & Krefft (1957); Karrer (1973); Rodger (2006); COSEWIC (2012).



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fins, two anal fins (about equal in size), and 14–26 total gill rakers; the lateral line is not wavy; pit organs are present on the head canals; and the last half of the lateral line is interrupted. The upper jaw projects beyond the lower jaw. Teeth are present on the upper jaw (premaxillaries), lower jaw (dentaries), and roof of mouth (the head of the vomer, but not on the palatines). A chin barbel is present, no longer than half the eye diameter. Individual dorsal fins and the anal fins are distinctly separated from each other. The first dorsal fin has 11–16 rays, the second dorsal fin 15–24 rays, and the third dorsal fin 17–24 rays. The first anal fin has 20–27 rays, and the second anal fin 18–23 rays. The pelvic fins are jugular (anterior to the pectoral fins), with 6 rays, the second of which is elongate. The pectoral fin-rays number 18–21. The caudal fin is square or slightly indented. Total vertebrae (including the urostyle) number 57–64. The lateral extensions (parapophyses) of the precaudal vertebrae, beginning with the seventh, eighth, ninth, or tenth vertebra, are expanded and hollow. The gas bladder horns are short and straight. Pit organs instead of pores are present on the head. The lateral line is continuous until slightly past the origin of the second dorsal fin; then it continues in an interrupted fashion to the end of the fleshy portion of the tail. The dorsal and lateral body aspects have dark grey-green to brown vermiculations on a yellowish background. The flanks can be silvery tinged with violet. The ventral aspect is yellowish to silvery white. The dorsal, anal, and caudal fins are dark

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Eleginus gracilis

with white edges. The peritoneum is silvery with black specks. The species attains 63.0 cm in total length, but usually is between 25 cm and 35 cm, and 1.3 kg in weight.

habitat: The species is benthopelagic in coastal waters, from the

surface down to 300 m, but is usually found above 60 m depth. It may occur at temperatures up to 10.5°C. It enters the brackish and fresh waters of rivers under tidal influence and is even found in a freshwater lake on Bering Island east of the Kamchatka Peninsula. In the southeastern Beaufort Sea it is found in salinities of 1.1‰– 8.0‰ in June–July, 10.0‰–27.0‰ in August, and 6.4‰–33.0‰ in September. The species is associated with silty bottoms for feeding and with sandy, pebbly bottoms for spawning. Young fish remain in shallow water throughout the year, and adults undergo limited migrations. In early winter the adults leave the silty bottom inshore waters of 30–60 m depth, where they were feeding, for brackish inshore sand-and-pebble areas, 2–10 m deep, where they spawn (December–March). They spend winter under ice and move offshore in early spring to the colder and more saline waters of the open sea, where they spend the summer feeding.

biology: The young and adults feed on fishes, notably Four-

horn Sculpins and coregonines, as well as calanoid copepods, epibenthic crustaceans (amphipods, decapods, isopods, and mysids), polychaetes, and bivalve molluscs. Plant material can also be present in up to 69% of stomachs. Larval and post-larval fish feed on copepods such as Microcalanus pygmaeus, Pseudocalanus minutus, Eurytemora raboti, and Oithona similis in the Beaufort Sea. It is preyed upon by Bearded, Harp, Ringed, and Spotted Seals and eaten by Beluga in Kugmallit Bay and east Mackenzie Bay. Longevity is up to 19 years. Both males and females become sexually mature at two to three years of age at a size greater than 17 cm total length. Fecundity is highly variable, from 4,900 to 680,000

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eggs. The eggs are 1.0–1.1 mm in diameter, demersal, and possibly adhesive. Spawning occurs once a year (December–March) at temperatures between −1.8°C and 0°C, and individuals can spawn up to 10 times during their lifetime. Larvae hatch out at 3.5 mm in total length in April–May and lead a planktonic life at 2.5–7.5 m depth for two to three months, reaching a total length of 7.5 mm in August. Young Saffron Cod can be found associated with jellyfishes. It possesses a glycoprotein in the blood that prevents it from freezing.

importance: A fishery has existed in the northwest Pacific Ocean

for the last 100 years. In 1987 the catch totalled 27,929 t, all taken by Russia. In Russia it is sold either fresh or frozen for human consumption. The flesh is said to be very tasty in fish caught during fall and winter. Some fishing is also carried out during late autumn and winter by Alaskan fishermen on the Alaskan side. The gear type is extremely varied and includes hook and line, beach seines, gillnets, hoop nets, and trawls. In the western coastal Canadian Arctic it is used domestically as food for humans and dogs. It occurs in the by-catch of the anadromous salmonid fisheries of the Mackenzie Delta. It is jigged under ice in large numbers in Tom Cod Bay, Northwest Territories. The Inuvialuit harvest this species for subsistence purposes (about 2,500 fishes yearly) out of Tuktoyaktuk, Paulatuk, and Sachs Harbour, Northwest Territories. At the offshore islands in Coronation Gulf 50 have been caught in a night’s fishing, but generally little use is made of this species.

distribution: The species is found in Simpson Strait, Queen Maud Gulf, Dease Strait, Melville Sound, Bathurst Inlet, Coronation Gulf, Dolphin and Union Strait, Amundsen Gulf, and Beaufort Sea. It is found in the North Pacific Ocean along the Asian coast from the Yellow Sea to the Chukchi Sea, and in North America along virtually the entire coast of Alaska as far south as Sitka.

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common names: Local names are Ogac, Ovak, and, along the

Labrador coast, Ugak and Uugak (Inuktitut); and Aalisangar, Saarullik, Saraudlik, Saraudlirksoak, Sârugdligaraq, Sârugdlik, and Sâugdlik (Greenlandic). Other common names are Codfish, Northern Cod, Scrod, morue commune, morue de l’Atlantique, and cabillaud. In Newfoundland and Labrador fishermen simply call this species “fish,” while all other fish species have a particular name.

Distribution of Eleginus gracilis

taxonomy: The genus comes from the Greek gados (codfish), and the species name is New Latin for codfish. The species belongs to the subfamily Gadinae. The species name often appears misspelled as morrhua in the older literature. Six subspecies have been recognized; however, only four of these (callarias, kildinensis, marisalbi, and morhua) refer to this species, and the other two, macrocephalus and ogac, are recognized as distinct species. Synonyms include Gadus callarias Linnaeus, 1758, described from the Baltic Sea; Gadus heteroglossus Walbaum, 1792, and Gadus vertagus Walbaum, 1792, both described from off Finland; Gadus arenosus Mitchill, 1815, and Gadus rupestris Mitchill, 1815, both described from off New York; Morhua vulgaris Fleming, 1828, described from off Scotland; Gadus nanus Faber, 1829, described from off Iceland; and Morrhua americana Storer, 1839, described from off Massachusetts.

sources: Scofield (1899); Walters (1955); Abrahamson et al.

description: This species is distinguished by having three dorsal

(1964); Slaney & Company (1975); Fraker et al. (1979); Hunter (1981); Burcham, Osuga, Yeh, & Feeney (1983); Ratynski (1983); Dunn & Vinter (1984); Lawrence et al. (1984); Burcham, Osuga, Rao, et al. (1986); Allen & Smith (1988); Lacho (1991); Carr et al. (1999); Naughton (2012).

Gadus morhua Linnaeus, 1758

Atlantic Cod, morue franche

Gadus morhua



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fins and two anal fins (about equal in size); total gill rakers number 18–28; the lateral line is not wavy and is white; there are no palatine teeth; the body has many greenish, brownish, or reddish spots; and the barbel length is less than eye diameter. The upper jaw projects slightly beyond the lower jaw, and the snout is blunt. The head and mouth are large, head length being about 25% of total length. There are numerous small, sharp teeth on the upper jaw (premaxillaries) and lower jaw (dentaries) and on the roof of the mouth (the head of the vomer). A chin barbel is present. The dorsal and anal fins are all separated from each other. The first dorsal fin has 12–16 rays, the second dorsal fin 14–24 rays, and the third dorsal fin 15–23 rays. The first anal fin has 17–27 rays, and the second anal fin 15–22 rays. The length of the fin-rays in each of these fins generally forms a gentle gradation, the longest ones being anterior and the shortest posterior. The pelvic fins are jugular (anterior to the pectoral fins) with 6 rays, the second one being elongate. The pectoral fins have 18–21 rays. The branchiostegal rays number 7. The caudal fin is square or slightly indented. Lateral-line pores are present on the head. The lateral line is continuous through most of its length and becomes interrupted only under the third dorsal fin and beyond. The scales are small, cycloid, and overlapping like shingles on a roof. Small horn-shaped or spade-shaped roughening structures (contact organs), one per scale, rarely two, are present at the posterior end of the scales of individuals of more than 40 cm total length. In individuals of 30 cm total length and greater, the anterior horns of the gas bladder are long (each horn being 26–65 mm when straightened) and either bent or coiled into a ball. There are 250–280 pyloric caeca. The peritoneum is leaden silvery in colour with black dots. The outer wall of the ovaries is

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Gadus morhua

unpigmented. The testicular lobes have either smooth or scalloped edges. Total vertebrae (including the urostyle) number 49–59. The dorsal and lateral parts of the body are covered with dark spots (reddish, brownish, or greenish) on a lighter background, but the lateral line is unpigmented and stands out as such. The overall colour is very variable depending on the background – usually brown to dark red or grey to green. Some fish may be almost black or even brick red. The ventral part of the body is pale. The fins are coloured like the neighbouring body. The species reaches 200.0 cm in total length and 95.9 kg in weight but is usually under 100.0 cm and 10.0 kg.

habitat: The species is benthopelagic and associated with a wide

variety of bottoms (rocky and pebbly, gravelly, sandy, or muddy) of the continental shelf and slightly beyond, from surface waters down to 700 m and possibly more, but usually above 150–200 m. However, it occurs pelagically early in its development, during spawning, and sometimes when feeding. It is tolerant of a wide range of salinity (almost fresh to full-strength sea water) and temperature (−2.0°C to 20.0°C, but preferred temperatures are 0°C to 8°C). Its distribution is partly influenced by temperature as it has been found at higher latitudes along the western coast of Greenland during warmer periods. The species is known to be landlocked in at least three coastal meromictic lakes (i.e., with a relatively fresh upper layer, a middle saline layer where the cod live, and an anoxic high-salinity bottom layer) on Baffin Island (Ogac Lake in Frobisher Bay, and Qasigialiminiq and Tariujarusiq Lakes in Cumberland Sound) and one island lake in Russia (Lake Mogil’noe on Kil’din Island in the Barents Sea). The three Baffin Island lakes receive influxes of tidal water during the open-water period but were apparently colonized between

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8,000 and 5,000 years ago, with little or no gene flow since. The Lake Mogil’noe population is recognized as a distinct subspecies (kildinensis). Atlantic Cod form compact schools during the day and disperse at night. Younger, smaller fish are usually found in shallow, warm waters, and older, larger fish in deeper, colder waters. Some populations are relatively sedentary (e.g., in the Baltic Sea), while others undergo migrations (feeding and spawning) that can attain over 1,000 km. There is a record of a 57 cm total length individual that was tagged on the North Sea and then captured 3.5 years later measuring 75 cm on the Grand Banks off southeast Newfoundland, a minimum distance of over 3,200 km. They can move at 25.7 km/day over a period of nearly a month. Those that are migratory move offshore in winter and nearshore in spring. The size of schools can reach tens of kilometers and hundreds of millions of fish. One school was about one-third the size of Prince Edward Island.

biology: The yolk-sac larvae feed on phytoplankton; after yolk

absorption they feed on zooplankton (copepods). Young fish up to 25 cm in total length feed mainly on small crustaceans (amphipods, euphausiids, isopods, mysids), and older fish mainly on decapods (crabs, lobster, shrimps), polychaetes, molluscs (mostly squids, but also cockles, sea clams, sea mussels), brittle stars, and fishes including Alewife, American Plaice, young Atlantic Cod, Atlantic Herring, Atlantic Mackerel, Arctic Cod, Blue Whiting, Capelin, Eelpouts, young Greenland Halibut, Grenadiers, Haddock, Lancetfishes, Lanternfishes, Pricklebacks, Rainbow Smelt, redfishes, Sand Lances, Sculpins, Silver Hake, Snailfishes, and Sticklebacks. Atlantic Cod from one of the Baffin Island lakes (Ogac Lake) eat sea urchins and sea stars. Small food items are located on or in the sea bed using the

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Gadus morhua

chin barbel and the pelvic fin-rays, which have touch and taste cells. Mid-water food items, as small as 2 mm, as well as larger bottom foods are located by sight. Small Atlantic Cod are eaten by squid and several predatory fishes, including Atlantic Cod, Atlantic Herring, Atlantic Mackerel, Fourspot Flounder, Pollock, Sea Raven, and Spiny Dogfish, and larger cod are eaten by marine mammals, including Gray, Harbor, Harp, and Hooded Seals, White-beaked Dolphins, and Pilot Whales. The growth rate is rather high but varies from region to region, with fish from lower latitudes generally growing faster, but not for as long as those from higher latitudes. The overall sex ratio (male to female) can vary considerably from 1:1 to 1:2.6. Males predominate at early ages (2–3 years old), and females predominate at older ages (6–11 years old). In the Canadian Arctic a 60 cm fish can vary from 5–6 to 12–15 years old depending on the locality and year collected. Its longevity is 29 years. Both male and female Atlantic Cod are sexually mature as early as two years old in the eastern North Atlantic Ocean and 2+ years old in the western North Atlantic Ocean. First maturity may be as late as 15 years old. Sexes are separate, but some hermaphroditic individuals have been reported. Fecundity is positively related to size, and females can produce between 23,000 and 12,000,000 eggs, the average being 1,000,000. Spawning occurs once a year, and



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some individuals may spawn only in alternate years, at temperatures between −1°C and 12°C (usually below 6°C) and salinities between 10‰ and 35‰. The eggs are released not all at once but in batches. During the spawning season males and females pair up and perform an elaborate courtship called a zigzag dance that starts at lower depths and culminates in spawning higher in the water column. Spawning takes place mostly at night. Males produce low-frequency grunting sounds during courtship by activating the drumming muscles attached to their gas bladder, which in turn serves as a resonator. Spawning may occur at various depths from near the surface down to 600 m. Throughout the range, spawning occurs virtually year round, but in any given region the spawning season lasts three to six months. In the Canadian Atlantic Ocean area, spawning begins in the north as early as February and ends in the south as late as December. The fertilized eggs are spherical, transparent, non-adhesive, and buoyant and are 1.1–1.9 mm in diameter. There is no oil globule. Depending on the incubation temperature, larvae 3–6 mm long hatch about one or two months post-fertilization. The eggs and early larvae are pelagic and subject to drifting caused by oceanic currents. After about three months, when the larvae reach 2.5–5.0 cm in total length, they settle to the bottom. This species is host to over 40 species of parasites. Prevalent parasites are the nematode Phocanema decipiens, commonly known as codworm or sealworm, found in the muscle; the nematodes

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Anisakis sp. and Contracaecum sp., found in the liver; and the copepod Lernaeocera branchialis, attached to the gills. Cod in Newfoundland and Labrador contain glycoproteins in their blood during winter, enabling them to be freeze tolerant.

importance: In the northwest Atlantic Ocean the Atlantic Cod

has been fished for 3,500 years and commercially for 500 years. It is the most important commercial fish in the northwest Atlantic Ocean (about 200 million tonnes were harvested over the 500-year period) and one of the most important food fishes worldwide. Its fishery represents about 30% of the world’s total catch of ground-fish (bottom-dwelling fishes). The 1987 world catch was 2,054,721 t. Of this total, about 70% was fished from the northeast Atlantic Ocean, mainly by Denmark (Faroe Islands), Iceland, Norway, Russia, and the United Kingdom, and about 30% was fished from the northwest Atlantic Ocean, mainly by Canada, France, Greenland, Portugal, Spain, and the United States. Between 2000 and 2009, landings in the North Atlantic varied from 769,000 t (2008) to 945,000 t (2001), with the northwest Atlantic representing about 6% and the northeast Atlantic 94% of these annual values. A moratorium on fishing the northern stock (off southern Labrador and eastern Newfoundland) was imposed by Canada in 1992 because of low catches over a number of years due to a multiplicity of factors including over-exploitation and environmental conditions. The northern stock cod fishery reopened in 1999 but at a much reduced total allowable catch of 9,000 t (8,500 t were reported caught), compared to that of 370,000 t in 1987. The fishing devices used are varied and include bottom (otter) and pelagic trawls, gill-nets, handlines (for jigging), cod traps, long lines baited with squid, and Danish seines. Since the early 2000s Atlantic Cod farming has been carried out in Norway and conducted on an experimental basis in New Brunswick, Newfoundland, and Labrador. The flesh of Atlantic Cod is white, and it is marketed fresh or frozen, whole or as fillets, in blocks or sticks, salted or sugar-salted, dried (either salted or unsalted), and in brine or smoked. Other market products include salted cod cheeks, fresh cod “tongues” (actually flesh taken from the throat area), liver, liver oil (an important source of omega-3 fatty acids and vitamins A and D), eggs (either smoked or frozen), fish-meal, and even glue. In Canadian Arctic marine waters there was an attempt in the early 1980s to revive a commercial inshore fishery for this species (one had existed intermittently from 1897 to 1975), operated by the Makivik Corporation off Killiniq Island on the Ungava Bay side. A local processing plant at Killiniq (Port Burwell) produced salted fish for export to southern markets. However, uncertainty surrounding the availability of Atlantic Cod every year made its continued exploitation for export problematic. Commercial exploitation of this species was contemplated in Cumberland Sound and Ogac Lake, on Baffin Island, in the late 1980s. In Hudson Strait and Ungava Bay and on the Canadian side of Davis Strait, Atlantic Cod has constituted a small by-catch of the shrimp (Pandalus borealis and P. montagui) fisheries since these began in 1977. There are also subsistence fisheries for the species along the coasts of Baffin Island and northern Québec.

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The Canadian populations of Atlantic Cod have been assessed as six designatable units (DUs) by the Committee on the Status of Endangered Wildlife in Canada (April 2010). Four DUs (Laurentian North, Laurentian South, Newfoundland and Labrador, and Southern) were assessed as “Endangered”; one DU (Arctic Lakes) was assessed as “Special Concern”; and one DU (Arctic Marine) was assessed as “Data Deficient.”

distribution: This species is found in southern Baffin Bay (at 70°27' N), Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, and Ungava Bay, as well as in northwest (to about 73° N), southwest, and southeast Greenland. It is found on both sides of the North Atlantic Ocean from Cape Hatteras (35° N), up into the Gulf of St Lawrence to Baffin Bay, around Iceland and the British Isles, south to the Bay of Biscay, into the Baltic Sea, up the coast of Norway, into the White Sea and along the western coast of Novaya Zemlya, and very occasionally just barely into the Kara Sea. The species is also found in the Barents Sea and the Greenland Sea up to the southern coast of Spitsbergen (as far as ca. 80° N).

Distribution of Gadus morhua

sources: McKenzie (1934); Anonymous (1951a, 1960a); Dunbar (1954b, 1956, 1958); Grainger (1954c, 1955); Kotthaus & Krefft (1957); Brawn (1961); Patriquin (1967); Konstantinov & Podrazhanskaya (1972); MacLaren Marex Inc. (1979b); Sandeman & Buchanan (1979); Lear (1984); Gillis & Allard (1984, 1986); Vladykov, Renaud, & Laframboise (1985); Allard & Gillis (1986); Renaud, Speers, Qadri, & McAllister (1986); Renaud (1989a, 1989b); Gillis et al. (1987); Hudon (1990a); Renaud & Morrison (1992); Rose (1993); Andersen et al. (1994); Goddard et al. (1994); Kulka (1995, 2011); Kulka & Tillman (2000); Treble et al. (2000); Evseenko & Pobalkova (2001); J.B.C. Jackson et al. (2001); Jørgensen et al. (2005); Evseenko, Laurel,

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Brown, & Malikova (2006); Hardie, Gillett, & Hutchings (2006); Rodger (2006); Helfman (2007); Carr & Marshall (2008); Hardie et al. (2008); Hardie & Hutchins (2011); Naughton (2012); Species at Risk Public Registry (2013).

Gadus ogac

Richardson, 1836 Greenland Cod, ogac

common names: Local names are Ôarsuk, Ogac, Ogak, Oggaksuk, Ovak, O-wuk, Ugak, Uugaq, Uugavik, and Uugayak, and along the Labrador coast, Uugaatsuk (Inuktitut); and Ûvak (Greenlandic). Other common names are Fjord Cod, Pilot, Ogac, Rock Cod, morue du Groenland, morue de roche, and morue ogac. It is the only species in the Canadian Arctic that shares the same common name, Ogac, in English, French, and Inuktitut. taxonomy: The species name is the Inuktitut ogac (codfish).

The species belongs to the subfamily Gadinae, and it has been treated as a subspecies of Gadus morhua. Gadus callarias marisalbi Derjugin, 1920, described from the White Sea, Russia, was considered by an author to be a probable synonym of Gadus ogac, but this is incorrect as the former is a synonym of Gadus morhua (see that species account). Based on the lack of genetic differentiation, numerous authors treated Gadus ogac as a junior synonym of the Pacific Cod (Gadus macrocephalus Tilesius, 1810), but this is also incorrect because it fails to take into account evidence of their specific distinctiveness based on differences in the larval pigmentation patterns and, in adults, the pelvic fin and chin barbel

pigmentation, the shape of the testicular lobes, and the shape and relative length of the contact organs described by other authors.

description: This species is distinguished by having three dorsal

fins and two anal fins (about equal in size); total gill rakers number 16–23; the lateral line is not wavy and is blotched; there are no palatine teeth; the body lacks many brown to reddish spots; and the barbel length equals or exceeds the eye diameter. The upper jaw projects slightly beyond the lower jaw. The head and the mouth are large. There are numerous small, sharp teeth on the upper jaw (premaxillaries), the lower jaw (dentaries), and the roof of the mouth (the head of the vomer). A chin barbel is present. The dorsal and anal fins are all separated from each other. The first dorsal fin has 12–17 rays, the second dorsal fin 14–22 rays, and the third dorsal fin 15–21 rays. The first anal fin has 17–25 rays, and the second anal fin 15–21 rays. The length of the fin-rays in each of these fins, except in the first dorsal fin that is more arch-like, generally forms a gentle gradation, the longest ones being anterior and the shortest posterior. The pelvic fins are jugular (anterior to the pectoral fins). The pelvic fins have 6 rays, the second rays being elongate. The pectoral fins have 17–19 rays. The branchiostegal rays number 7. The total outer gill raker count on the first arch is 16–23, based on specimens examined from across the range; an upper limit of 30 reported by one author appears too high, and of 43 by others is certainly erroneous. The caudal fin is square or slightly indented. Lateral-line pores are present on the head. The lateral line is continuous through most of its length and becomes interrupted only under the third dorsal fin or on the caudal peduncle and beyond. The scales are small, cycloid, and overlapping like shingles on a roof. Large club-shaped roughening structures (contact organs), one per scale, are present at the posterior end of the scales of individuals of more than 25 cm total length. In individuals of 30 cm total length and greater, the anterior horns of the gas bladder are short (13–26 mm when straightened) and bent. The body has dark-brownish

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vermiculations on a yellowish background, and this pattern is also present over the lateral line, making the latter inconspicuous. The dorsal and caudal fins have black margins. The chin barbel is black. The pectoral and pelvic fins are reddish brown to black. The belly is grey to white. The peritoneum is black, and the outer wall of the ovaries is darkly pigmented. The testicular lobes have scalloped edges. The total number of vertebrae (including the urostyle) is 51–57. The species attains 77.0 cm in total length and about 7.0 kg in weight, but usually less than 50.0 cm, and 3.0 kg.

habitat: The species is benthopelagic and found along the coast

from surface waters down to 462 m at temperatures of 0°C–10°C. It is associated with soft bottoms and is found near rocky outcrops. It is found in salinities of up to 23‰, but it can tolerate low salinities down to 4‰ and is sometimes found in estuaries (Eastmain and La Grande Rivers flowing into eastern James Bay, and Great Whale, Little Whale, and Innuksuac Rivers flowing into eastern Hudson Bay). It is non-schooling and non-migratory other than making local seasonal migrations between the estuary (winter) and the coastal waters (summer), although one tagged individual was recovered at 67 km from the tagging site (a migration from Cambridge Bay to Wellington Bay). Abundance varies; for example, it is numerous for one year in seven along the west coast of Parry Peninsula in Franklin Bay.

biology: Its diet is diverse and includes fishes (Arctic Cod, Arctic Shanny, Capelin, Cisco, Fourhorn Sculpin, small Greenland Cod, Greenland Halibut, Northern Sand Lance, Slender Eelblenny, Smelts, Threespine Stickleback), crustaceans (amphipods, crabs, cumaceans, euphausiids, isopods, mysids, shrimps), other invertebrates (bivalve molluscs, echinoderms, nemerteans, polychaetes, squids), and plant material. It is eaten by Atlantic Cod, Beluga, and seals. Fish of about 50 cm total length are five to six, or nine to ten, years of age depending on the locality and the year collected. In northwest Hudson Bay at 63°40' N, fish of 35 cm total length are seven years old. The maximum age is 21 years, but fish in James Bay, for example, live only nine years. The sex ratio is 1:1. The species attains sexual maturity at two to four years old and spawns annually thereafter in shallow brackish waters from February to June. Fecundity varies between 1,000,000 and 7,350,000 eggs/female. The eggs are spherical, 0.6–1.3 mm in diameter, adhesive, and without an oil globule. The eggs sink to the bottom after spawning. Hatching occurs at 38 days. The larvae are about 4 mm long at hatching and pelagic. Parasites include the nematode Phocanema decipiens, commonly known as codworm or sealworm, found in the muscle, and the copepod Lernaeocera branchialis, attached to the gills. Glycoproteins in its blood prevent it from freezing in sea water that is close to the freezing point of –1.8°C. importance: The world catch in 1987 was 4,017 t, mostly caught

in Greenland. It is also caught as a by-catch of the Atlantic Cod and Arctic Char fisheries. Its flesh is white, flaky, and firm and is said

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to be sweeter tasting than that of Atlantic Cod. In Canadian Arctic marine waters there are subsistence fisheries in the north-central coastal Arctic and Arctic Archipelago, Hudson Bay, Foxe Basin, along Baffin Island, and northern Québec. The method of capture is jigging in ice cracks. In northern Québec the catch from nine villages along the east shore of Hudson Bay up into Ungava Bay is about 8,000 kg yearly. In Nunavut, exploratory fisheries with quotas of 1,200–2,000 kg each were permitted in 1991 at Taloyoak (Spence Bay) and Soper Lake, Baffin Island, and in 1993 at Peterson Bay, King William Island, and at Ogle Point, Thunder Cove, and Richardson Point (all three localities on the Adelaide Peninsula). A small commercial fishery started in 1985 at Kimmirut (Lake Harbour), Baffin Island, but it has not expanded beyond the local community. In the Belcher Islands Greenland Cod is eaten raw with or without berries. It is also utilized as dog food.

distribution: The species is found in Davis Strait, northern Baffin Island at Ikpiarjuk (Arctic Bay) and Mittimatalik (Pond Inlet), Frobisher Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, Gulf of Boothia, Queen Maud Gulf, Dease Strait, Melville Sound, Bathurst Inlet, Coronation Gulf, Dolphin and Union Strait, Amundsen Gulf, western Victoria and Banks Islands, and the Beaufort Sea. General reports for Cumberland Sound have not been confirmed by specimens. It is found also in northwest (at least up to 73°30' N) and southwest Greenland; northern Alaska; and south along the coast of Labrador, into the Gulf of St Lawrence, in the Saguenay River, and in Bras d’Or Lake on Cape Breton Island.

Distribution of Gadus ogac

sources: Richardson (1836a); Walters (1955); Kotthaus & Krefft

(1957); Legendre (1961); James Bay Development Corporation (1976); McAllister (1977); Fraker et al. (1979); Sandeman & Buchanan (1979);

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R. Morin (1980); Hunter (1981); Gillis & Allard (1984, 1986); Vladykov et al. (1985); Allard & Gillis (1986); Gillis et al. (1987); Dodge (1989); Mikhail & Welch (1989); Renaud (1989a, 1989b); B. Morin (1990); Morin, Hudon, & Whoriskey (1991); D.B. Stewart et al. (1991); Renaud & Morrison (1992); Nielsen & Morin (1993); Andersen et al. (1994); D.B. Stewart (1994b); Bright, Grundy, & Reimer (1995); Carr et al. (1999); Evseenko & Pobalkova (2001); Jørgensen et al. (2005); Coulson, Marshall, Pepin, & Carr (2006); Evseenko et al. (2006); Carr & Marshall (2008); Hardie et al. (2008); Kulka (2011).

Lota lota

(Linnaeus, 1758) Burbot, lotte

common names: Local names are Chehluk (Gwich’in); Shulukpaoluk, Tiktaaliq, Tiktabek, Tiktailik, Tiktalaq, Tiktalik, and Titaliq (Inuktitut); Titaalirq (Inuvialuktun); and, in Alaska, Nätarrnaq, Titale, and Tittaalik (Inuktitut). Other common names are Eelpout, Lawyer, Ling, Loche, Lush, Maria, Methy, and many others. taxonomy: The genus and species names come from the French name for the fish, lotte. The species belongs to the subfamily Lotinae and was originally described under the genus Gadus. Authors have also placed it in the genus Enchelyopus. Synonyms include Gadus lacustris Walbaum, 1792, described from Hudson Bay; Gadus maculosus Lesueur, 1817, described from Lake Erie; Gadus compressus Lesueur, 1817, and Molva huntia Lesueur, 1819, both described from the Connecticut River, New England, United States; Lota brosmiana Storer, 1842, described from New Hampshire, United States; and Lota inornata DeKay, 1842, described from New York. Various authors have recognized up to eight subspecies, namely asiatica, kamensis, lacustris, leptura, lota, maculosa, obensis, and onegensis. Authors restrict this species to Europe and part of Siberia, while referring to populations

in eastern Siberia and North America as Lota maculosa, but do not provide an explanation for their action, other than to say that the genus is in need of revision.

description: This species is distinguished by having two dorsal

fins and one anal fin. The jaws are equal or the upper jaw is slightly longer than the lower jaw. The mouth is terminal and large. The teeth are small and present on the upper jaw (premaxillae), the lower jaw (dentaries), and the roof of the mouth (the head of the vomer, but not on the palatines). Anterior nostrils each have a prominent fleshy, tubebased flap. A chin barbel is present. The eye diameter is about 30% of snout length. There are two dorsal fins, the first being short with 7–16 similarly sized rays, and the second being long with 60–94 similarly sized rays. There is one long anal fin with 52–86 similarly sized rays. The origin of the anal fin is posterior to the origin of the second dorsal fin. The bases of the second dorsal fin and the anal fin touch the base of the caudal fin but form deep notches where they come into contact. The caudal fin is rounded. The pelvic fins are jugular (anterior to the pectoral fins), with 5–9 rays, the second ones being elongate. The pectoral fins have 15–24 rays. Total outer gill rakers on the first arch number 5–12 and are short. The branchiostegal rays number 6–7, rarely 8. Pit organs are present on the head instead of pores. The continuous lateral line ends where the second dorsal fin contacts the caudal fin. The scales are cycloid and extremely small. The pyloric caeca number 21–168. Drumming muscles are present on the gas bladder. The gas bladder has two short anterior horns. The total vertebrae (including the urostyle) number 50–67. The body colour is yellow, light tan, to brown with more or less well defined olive-green, dark-brown, or black vermiculations on the dorsal and lateral aspects as well as on the pectoral, dorsal, and caudal fins. The second dorsal fin, the caudal fin, and the anal fin have a dark submarginal band, while the margin is bright yellow or orange. The pelvic fins are pale. The species attains 180.0 cm in total length and 27–34 kg in weight, but is usually about 40–70 cm and 1–3 kg.

habitat: The species is benthopelagic, and, although normally found in fresh water, it can tolerate salinities up to 20‰ to feed – at the mouth of La Grande River, in James Bay, more than 12 km

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offshore of Great Whale River in Hudson Bay, up to 10 km off the mouth of the Mackenzie River, in tidal waters between Mackenzie River and Shingle Point (in 1843), and at the mouth of Coppermine River. In fresh waters it is usually found at the bottom of deep lakes down to 300 m and of large rivers with slow current, but it can also be found in small streams and in ponds, at depths as shallow as 0.3 m, and at temperatures of 0°C–18°C. In shallow waters it shelters under rocks, in crevices along the river banks, among the roots of trees, and in dense vegetation; in deep lakes it may be found in extensive burrows that it constructs in the substrate. Burbot stay in deeper water during the day and feed at night in shallow water. Burbot are generally sedentary outside of the spawning season. In Great Slave Lake 67% of tagged individuals were recovered within 10 km of their tagging site after an average of 581 days. One individual, however, was recovered 406 km away in the Slave River after only 90 days.

biology: Although primarily a freshwater species, it is known to

occur in brackish waters during the summer throughout the Canadian Arctic. Most aspects of its biology, therefore, refer to fresh waters. It is most active during winter and shows little activity during summer. Individuals under 50 cm total length feed mostly on a variety of invertebrates (amphipods, caddisfly larvae, crayfish, fingernail clams, opossum shrimp), and individuals greater than 50 cm total length feed almost exclusively on fishes (Alewife, Arctic Grayling, Burbot, ciscoes, Freshwater Drum, Kokanee, Logperch, minnows, Ninespine Stickleback, Northern Pike, Rainbow Smelt, Sculpins, Suckers, Trout-perch, Walleye, White Bass, whitefishes, Yellow Perch). Plant material, fish eggs, shield shrimps, and snails have also been reported from stomach contents. Burbot in brackish water have been reported feeding on amphipods, copepods, isopods, euphausiids, and mysids. Young fish reach 8–21 cm in total length at the end of their first year. Individuals of seven to nine years of age in the western Canadian Arctic attain total lengths of 50–55 cm. Longevity is 24 years. Sexual maturity is attained at ages three to four years and at a size greater than 24 cm in total length in southern populations, and at four to seven years in northern populations. Males mature about a year earlier than females. Adults do not spawn every year. Spawning migrations may vary from 17 to 255 km. Spawning occurs at night, between January and March (in Canada), over a two- to three-week period and at temperatures of 1°C–4°C. This usually occurs under ice, in lake shallows, or in areas of streams with slow current at depths of 0.3–10.0 m, over sand, gravel, or cobble. Males arrive on the spawning grounds first, followed three to four days later by the females. During spawning, 10–12 individuals (one or two females surrounded by males) come together to form a continuously moving ball. No nest is constructed, and no parental care is given. The semi-buoyant fertilized eggs are non-adhesive, about 1–2 mm in diameter, and possess a large oil globule. Fecundity ranges from 6,300 to 3,477,699 eggs, usually 300,000–400,000. Hatching occurs after about 30 days at 6°C, and 100 days or more at 0°C. The total length at hatching is 4 mm. The larvae are pelagic. At about 5–23 days the larvae begin to feed on their own.

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Parasites include protozoans, cestodes, nematodes, trematodes, acanthocephalans, leeches, mollucs, and crustaceans.

importance: It is of minor importance in Canada. The 1987 world

catch of 1,577 t was harvested in Finland (98%) and Sweden (2%), and an unreported catch was harvested in the European and Siberian parts of Russia. Its flesh is white and flaky and said to be tasty but a little dry. It is used mainly in salted form for human consumption or as pet food, but it is also processed as fish-meal. The liver is considered to be a delicacy and a good source of vitamins A and D and is sold smoked or canned in Europe. In northwestern Canada in the nineteenth century the eggs were mixed with flour to make biscuits. The Inuvialuit harvest this species for subsistence purposes (about 6,000–7,000 fishes yearly) out of Aklavik and Inuvik, Northwest Territories, principally during October–November. The winter harvest is caught by jigging through the ice. A small commercial fishery exists at Paulatuk, Northwest Territories. The Cree from five villages along the eastern shore of James Bay and Hudson Bay catch somewhat less than 3,000 kg yearly.

distribution: It is found in southern Hudson Bay and James Bay, western Coronation Gulf (Kugluktuk or Coppermine), Amundsen Gulf (Paulatuk), and the Beaufort Sea. It is found circumarctic in fresh waters between about 40° and 77° N.

Distribution of Lota lota

sources: Simpson (1843); Walters (1955); St-Charles & Roy (1974);

Milne & Smiley (1976); Hunter (1981); Lawrence et al. (1984); Dodge (1989); Gilbert (1991); Bernatchez & Giroux (2000); McPhail & Paragamian, in Paragamian & Willis (2000); Kottelat & Freyhof (2007).

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Other common names are Couch’s Whiting and poutassou.

163–181 small, cycloid scales in the lateral line. Total outer gill rakers on the first arch number 26–34. The branchiostegal rays number 7. The body colour is silvery blue dorsally, silvery on the sides, and white ventrally. Sometimes a dark blotch is present at the base of the pectoral fins. The peritoneum is black. The pyloric caeca number 8–15. Total vertebrae (including the urostyle) number 56–60. The species attains 50.0 cm in total length, but usually 30–35 cm, and 830 g in weight.

taxonomy: The genus comes from the Greek mikros (small),

habitat: The species is mesopelagic at depths from the surface to

Micromesistius poutassou (Risso, 1827)

Blue Whiting, merlan bleu

common names: A local name is Imaluunniit (Greenlandic).

mesos (middle), and istion (sail), in reference to the second dorsal fin. The species name is the Provençal word for this fish. The species belongs to the subfamily Gadinae and was originally described under the genus Merlangus. It was placed under the genus Boreogadus or the genus Gadus by authors in the late nineteenth and early twentieth century. Merlangus vernalis Risso, 1827, and Merlangus pertusus Cocco, 1829, both described from the Mediterranean Sea, and Merlangus albus Yarrell, 1841, described from off the United Kingdom, are synonyms.

description: This species is distinguished by having three dorsal fins and two anal fins, the first anal fin being much longer than the second anal fin and lying under the first and second dorsal fins. The lower jaw projects slightly beyond the upper jaw. Teeth are present on the upper jaw (premaxillaries), the lower jaw (dentaries), and the roof of the mouth (the head of the vomer, but not on the palatines). There is no chin barbel. The three dorsal fins are widely separated, the space between the second and third being wider than the length of the base of the first dorsal fin. The first dorsal fin has 11–15 rays, the second dorsal fin 9–14 rays, and the third dorsal fin 22–28 rays. The two anal fins are in contact at their base or have a narrow interspace. The first anal fin has 30–42 rays, and the second anal fin 22–30 rays. The origin of the first anal fin is slightly anterior to the origin of the first dorsal fin. The caudal fin is slightly forked. The pelvic fins are jugular (anterior to the pectoral fins), with 6 rays each. The pectoral fins have 18–23 rays. Lateral-line pores are present on the head. The lateral line is continuous to the base of the caudal fin and placed high above the midline. There are

3,000 m (usually 300–400 m) and at −0.3°C to 15°C (usually 1°C to 9°C) in full-strength sea water. It undertakes daily vertical migrations from near the bottom during the day towards surface waters at night.

biology: It feeds largely on pelagic crustaceans (amphipods,

copepods, euphausiids, and shrimps), but larger individuals feed also on small fishes (Lanternfishes) and cephalopods. It is preyed upon by Atlantic Cod. Early growth is fast, reaching 16–18 cm at one year of age, 24–25 cm at two years, 26 cm at three years, 29–30 cm at four years, 27–34 cm at five years, and 29–34 cm at ten years. Longevity is at least 20 years. First maturity is at three years of age. The spawning season in the western North Atlantic is from mid-March to mid-May and in the eastern North Atlantic from February to June (above the continental shelf of the western U.K. islands), after which the fish migrates north to the Faroe Islands, Iceland, and Norway. Spawning occurs at depths of 1,000 m or more, at temperatures of 8°C–9°C, and at salinity of 36.2‰ or more. Fecundity is highly variable at 6,000–150,000 eggs. Fertilized egg size is 1.04–1.28 mm in diameter, and eggs are without an oil globule. The size of larvae at hatching is 2.0–2.2 mm. Eggs, larvae, and young lead a pelagic mode of life.

importance: In the eastern North Atlantic and the Mediter-

ranean the 1987 landings amounted to 707,955 t; 98% of this catch was made in the eastern North Atlantic, and two-thirds of that were caught by fishing vessels of the former USSR and of Norway.

Micromesistius poutassou



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Since 1999 this species has surpassed Atlantic Cod in landings in the northeast Atlantic, reaching a peak of 2,400,000 t in 2004, but dropping to 640,000 t in 2009. The methods of harvesting are various and include trawls, long lines, trammel-nets, gill-nets, and purse seines mostly beyond the edge of the continental shelf, and it is therefore considered a deep-sea fishery. It is sold fresh and frozen and processed as oil and fish-meal. Its flesh is considered excellent.

distribution: The species is found in Davis Strait and in south-

Family Bythitidae Viviparous Brotulas, Donzelles vivipares

Brian W. Coad

west and southeast Greenland. It occurs in both the North American and the European parts of the North Atlantic Ocean. In the eastern North Atlantic it is found along the African coast at Cape Bojador (26° N), up to the southern coast of Iceland, and in the Norwegian Sea and the Barents Sea up to 79° N; it is also found in the Mediterranean Sea along the northern half as far as Greece. In the western North Atlantic it is found along the northeastern coast of the United States and along southeast Canada north to Davis Strait.

Viviparous or Livebearing Brotulas are found in the Atlantic, Indian, and Pacific Oceans. There are about 96 species, with four found in Canadian waters, one of which is on the Pacific coast and one in eastern Arctic waters. The maximum size is about 46 cm. The fins lack spines, and the pelvic fins are thoracic when they are present. The scales are usually present and cycloid or may be absent. The pelvic fin soft rays range from 0 to 2. Most members of this family have the anterior nostril close to the upper lip. A gas bladder and pyloric caeca are present. There is usually a strong spine on the operculum. There are usually less than eight long gill rakers on the anterior gill arch. Males have a penis or intromittent organ, and these fishes bear live young rather than shedding eggs. The skin is loose and is thick over the dorsal and anal fins. Livebearing Brotulas are found mostly in coastal waters near the shore or at moderate depths, but some are deep-sea fishes, and others are found in fresh water. They are secretive, hiding in crevices and caves or in burrows.

sources: Nielsen & Cohen (1973, 2002). Distribution of Micromesistius poutassou

sources: Kotthaus & Krefft (1957); Scott (1963); Jørgensen et al.

(2005); Medley, in FAO (2011); Tandstad, Shotton, Sanders, & Carocci (2011).

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importance: It is not economically important.

Bythites fuscus Reinhardt, 1837

distribution: The species has been found only off Baffin Island

Arctic Brotula, donzelle arctique

common names: A local name is Arktisk Brosmekvabbe (Danish/Greenlandic). The common name comes from Nielsen, Bertelsen, and Nystrøm (1992).

as a single collection at 61°56' N, 61°16' E, in the Canadian Davis Strait, and is also found at Fiskenæsset in west Greenland and in the Denmark Strait off east Greenland.

taxonomy: The genus name is the Greek bythites (an animal from

the bottom of the sea). The species name is the Latin fuscus (dusky). This species is known only from the holotype collected in 1834, two other specimens from the eastern Canadian Arctic caught 166 years later, and one fish from the Denmark Strait.

description: This species has a pair of large pores on the lower

jaw, and large lateral-line papillae are evident from the level of the anus to just anterior to the caudal fin. The dorsal fin-rays number 88–95, the anal fin-rays 66–70, and the pectoral fin-rays 29–31. Total vertebrae number 52–53. The lateral line runs near the dorsal profile from the anus anteriorly. The teeth are large and pointed. The head is covered by numerous papillae of two sizes. There are no developed gill rakers on the anterior gill arch, and the remaining gill rakers are separate knobs. The overall colour is brown. The species attains 14.6 cm in standard length, but according to Greenlanders it reaches the size of a seal.

habitat: A specimen has been caught at 526–531 m and 3.0°C–3.5°C in the Canadian Arctic.

biology: Although generally unknown, it is thought to be epiben-

Distribution of Bythites fuscus

sources: See the family sources and the bibliography.

thic. A 9.7 cm male specimen was ripe, while a 9.3 cm female was not.

Bythites fuscus



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Lophius americanus

Family Lophiidae

Valenciennes, 1837

Goosefish, baudroie d’Amérique

Goosefishes, Baudroies

Brian W. Coad

The Goosefishes or Monkfishes are anglerfishes found in the Arctic, Atlantic, Indian, and Pacific Oceans. There are 25 species, including one in Atlantic and Arctic Canada. The maximum size is 1.38 m. These fishes characteristically have a wide and flattened head with large spines, knobs, and ridges on top, and strong, curved, and depressible teeth in the large mouth. Flaps are present on the lower jaw and also run along the lower head and onto the body, aiding in camouflage. The pectoral fins are on stalks and are arm-like. There are no scales, and the skin is loose and slippery. The pelvic fins and a pseudobranch are present, and the frontal bones of the head are united. The fishing apparatus has a fleshy flap at the tip to attract prey close to the mouth. The second and third dorsal spines on the head are elongate but are not lures. Goosefishes live in shallow to moderately deep waters often in temperate parts of the world. Prey is engulfed by the huge mouth when it is in striking distance. As well as fishes and crabs, they have been recorded as eating small sharks and seabirds taken at the surface (hence Goosefish, though this seems unlikely). The females lay large pelagic egg masses or veils. The larvae have long pelvic fins that shrink as the fish grows. Despite their ugly appearance, they are economically important.

source: Caruso (1983).

common names: Other common names are Abbot, Allmouth, American Angler, Bellyfish, Fishing Frog, Greedigut, Molligut, Molykite, Monkfish, Mud-fish, Rape, Sea Devil, baudroie, diable de mer, lotte, and poisson-pêcheur.

taxonomy: The genus comes from the Greek lophos (crest or tuft). The species is named for America. Four collections identified as this species have been made off eastern Baffin Island, but no voucher specimens were kept. The species is included here because its distinctive appearance makes a correct identification likely. However, it should be noted that the related Lophius piscatorius Linnaeus, 1758, of Europe and Iceland is rarely reported (two specimens) from southwest Greenland. It is distinguished by having a bifid esca (versus a simple flag-like esca or “bait”), a long third dorsal fin spine at 11.6%–19.9% of standard length (versus 2.3%– 9.7%), and a fourth dorsal fin spine with length greater than snout width (versus less than snout width). description: The large, flattened head with a very wide mouth

full of recurved teeth, the flaps along the lower head and flank margin, and the fishing apparatus distinguish this anglerfish. The body tapers behind the pectoral fins. The lower jaw projects in such a way that the lower jaw teeth are exposed even when the mouth is closed. The first dorsal fin has three spines on the head, the first being the illicium or fishing apparatus. There is a series of three shorter spines connected by a membrane, and a second dorsal fin of 9–12 rays. The anal fin has 8–10 rays, and the pectoral fins have 19–28 rays. The colour is dark brown to tan above, with various blotches becoming lighter to white or dirty white on the belly. The spine membranes are black. The pectoral fins have a dark tip, and the pelvic fins are reddish. The eyes are green. The esca is greenish in life and has a dark spot at its base. The peritoneum is light. This fish may be able to match its colour and pattern to the bottom on which it rests. The young are mottled green and brown. The species attains 1.38 m in length and 31.5 kg in weight.

habitat: The Arctic records are from average depths of 1,040–1,221

m, deeper than the Atlantic Canadian and other captures, which are from the tide-line down to at least 840 m, although few are caught below 400 m. On the Scotian Shelf in Atlantic Canada captures are at 27–366 m. They are usually found demersally over gravel, pebble, shell, sand, or clay bottoms with cooler temperatures preferred, 3°C–6°C in winter and 5°C–9°C in summer, although they tolerate 0°C–24°C. There is a migration to deeper waters in winter. They may partially bury themselves in bottom sediment as concealment from prey.

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biology: The principal food is a wide variety of bony fishes,

including their own species, and also sharks and rays. Invertebrates are taken, especially when young, and adults will take such items as squids. The occasional unwary seabird at the surface is also captured. They are voracious feeders and can take a fish almost their own size. As a “sit-and-wait” predator, they catch most of their food using the fishing apparatus. The lifespan is a maximum of 30 years, females living longer than males, and growth is fast, with young reaching 11.4 cm by one year of age. In one study, maturity was attained at 3+ years (37 cm total length) in males and 4+ years (48.5 cm) in females. Spawning occurs in June–September in southern Canadian waters. The eggs are laid in a pink, violet-grey, or purple-brown veil or mucous sheet, up to 1.5 m wide and 12 m long. Each egg is 1.6–1.8 mm in diameter, and there are up to three million in the veil.

importance: The Goosefish is not heavily fished in Atlantic Can-

ada but may be taken incidentally in trawls and on long lines and used as fish-meal. It is considered good to eat and in the United States is landed as “tails only,” the large inedible head being discarded. The Canadian catch in 1985 was 1,845 t, and the catch in the Gulf of Maine to Mid-Atlantic reached a high of 26,800 t in 1996. It has been sold as “mock lobster” but is now marketed under its own name. Goosefish livers have also been marketed, reaching as high as $19 per pound. The species has been used by scientists as an experimental animal in physiological and biochemical research.

Distribution of Lophius americanus

sources: Staudinger (2006); A.K. Johnson, Richards, Cullen, & Sutherland (2008); R.A. Richards, Nitschke, & Sosebee (2008).

distribution: It has been found as four records off eastern Baffin Island in Baffin Bay (see the “Taxonomy” section). It is also found along the Atlantic coast of Canada and the United States south to Florida. Previous northern Canadian records were from the Gulf of St Lawrence and Newfoundland.

Lophius americanus



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Family Himantolophidae Footballfishes, Poissons-football

Brian W. Coad

The snout and chin papillae may serve to provide a poor surface for attachment when squid prey struggle to escape using their sucking discs. Footballfish may eject a luminous cloud of mucus from the esca to confuse predators that are trying to eat them.

sources: Regan & Trewavas (1932); Bertelsen (1951); Bertelsen & Krefft (1988); Pietsch (2009).

Himantolophus groenlandicus Reinhardt, 1837

Atlantic Footballfish, football fine-lampe

common names: A local name is Reinhardts Fakkelanger These deep-sea anglerfishes are found worldwide. Larvae, males, and juveniles are known from temperate to tropical waters, and large expatriate females can be found in sub-Arctic to Arctic waters. There are 18 species, including three recorded from Canada, of which one enters Arctic waters. The maximum size is 46.5 cm in females. The females of these fishes are globular in shape (hence, footballfish) with a blunt and short snout, have bony plates embedded in the skin, each with a median spine, and have the snout and chin covered with dense, wart-like papillae. The illicium or fishing rod is short and stout, and the esca or bait is large and bulbous and has numerous filaments. The body muscles are soft, and the bone spongy. The vomer bone in the roof of the mouth is broad and toothless. The pelvic bones are triradiate. The parietal bones in the skull are absent, in contrast to other Canadian anglerfishes. There are no pelvic fins, scales, or pseudobranch; the frontal bones on the head are not united; the lower pharyngeal bones are reduced and toothless; and there are 5–6 dorsal fin-rays, 4 anal fin-rays, 14–18 pectoral fin-rays, and 8–9 caudal fin-rays. Various additional osteological characters serve to distinguish the family. Its overall colour is black to dark brown. The pigmentation of the illicium or fishing apparatus changes with age and is highly variable between species. Adult males are not parasites on the females, as they are with some other anglerfishes, but are dwarfed, reaching 39 mm in standard length compared to 465 mm for females. The males are thought to locate females by species-specific pheromones emitted by the females. The males have well-developed nostrils and small jaws with 16–31 denticular teeth on the snout and 20–50 on the chin, each group of teeth being fused at their base to form upper and lower denticular bones. Adults are found down to 1,800 m or more but can be as shallow as 50 m, and they are occasionally washed ashore. The larvae are found mainly in the upper 50 m of the ocean. Food is attracted by light emitted from the esca and by movements of the filaments. The prey is sucked into the mouth when it approaches close enough.

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(Danish/Greenlandic). Other common names include Lightlamp Footballfish and football atlantique.

taxonomy: The genus comes from the Greek himas (thong or strap) and lophos (crest or tuft), in reference to the illicium and esca. The species is named for Greenland, where the type-specimen was found washed ashore at Godthåb. Himantolophus reinhardti Lütken, 1878, described from western Greenland and found dead at the surface, is a synonym. This species was identified aboard ship, and no voucher specimen was kept. Its distinctive shape and presence in neighbouring Greenland waters occasion its inclusion here. description: This species is distinguished by family characters and by esca or bait characters such as the possession of short filaments. The anterior escal appendage is usually simple, rarely divided into more than two branches, and the posterior appendage is simple, in two parts, or divided into 8–9 branches. The illicium and esca are heavily tuberculate. The dorsal fin-rays number 4–6, the anal fin-rays 4, and the pectoral fin-rays 14–18. There are 1–5 large spines on each lobe of the pectoral fin. The body has up to 4–60 similar spines, the number increasing with age. Adults are ashen to black or black brown overall, and young are a deep brown colour. The fins are greyish brown with blackish tips. The escal appendages are dark except for their tips, which are bright silver. Females reach 46.5 cm, and males are less than 4.0 cm. habitat: This species has been reported generally as deep as

1,830 m and as shallow as 137–146 m in southern Canadian waters. Northern captures are often in waters shallower than 200 m or are stranded on shore although the Canadian Arctic record was at 821 m. It is the commonest of the Footballfishes, comprising more than a third of all known specimens in the family.

biology: Its food includes fishes, cephalopods, and crustaceans.

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Himantolophus groenlandicus

importance: It is not economically important. distribution: It has been found in Davis Strait off southern Baffin Island as a single record from cruise data at 63.565º N, 59.885º W, and is also found from southwest Greenland. It occurs in the North Atlantic Ocean from Norway to South Africa and may occur in the western Indian Ocean. sources: See the family sources and the bibliography.

Distribution of Himantolophus groenlandicus



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Chaenophryne longiceps

Family Oneirodidae

Regan, 1925

Can-opener Smoothdream, doux-rêve ouvre-boîte

Dreamers, Rêveurs

Brian W. Coad

common names: A local name is Glathovedet Mareangler

(Danish/Greenlandic).

taxonomy: The genus comes from the Greek chaeno (yawn, gape) and phryne (toad), in reference to the mouth. The species name comes from the Latin longus (long) and -ceps (head). description: This species of deep-sea anglerfish is distinguished

Dreamers are found worldwide in the deeper parts of the oceans. There are about 60 species, including 18 occurring in Canadian waters, of which 3 are found in Arctic waters. Many species are quite rare and known only from a few specimens. The maximum size is about 37 cm in females, and males are dwarfs at less than 2.5 cm. Dreamers are deep-sea anglerfishes and often have a stout body, ranging from oval to globular in shape, but some are slender and compressed. The shapes are unusual enough to warrant their common name in allusion to the dream-like quality. The first dorsal fin spine is modified into the fishing apparatus and is directed forwards, sometimes with a joint. The esca or bait at the tip of the illicium or fishing rod contains luminescent bacteria. The esca lacks spines on its bulb. The second dorsal spine is minute and covered by skin. Females have naked skin, sometimes containing short and minute spines, but males are naked. The dorsal fin-rays number 4–8, and the anal fin-rays 4–7. The pelvic fins are absent. The teeth are depressible. The overall colour is dark brown to black except for the tip of the fishing apparatus. The mouth is darkly pigmented. The males lack jaw teeth but have denticles on the snout and lower jaw tips that are used for attachment to females. They probably do not attach permanently in most species, unlike some anglerfish males that become parasites on the females. The male olfactory organs are large to facilitate locating the female by smell. Their food includes fishes, crustaceans, and squids. The female fishing apparatus is used to entice prey close enough to be gulped down. The eggs may be laid in a jelly veil. The larvae are found in surface waters, but adults are mesopelagic to bathypelagic, down to about 3,000 m. They are not economically important. Dreamers are identified in part by details of the esca, a structure that requires some expertise to understand, may show variations with growth, and may be missing or damaged. Males and juveniles are often unknown, and males can often be identified only to the genus.

by the lack of sphenotic spines, a slightly concave operculum posteriorly, and high, cancellous bones, as well as details of the esca. The female Can-opener Smoothdream is separated from its relatives most readily by having pectoral fin-rays numbering 17–22, rarely less than 18. The esca or bait on the fishing apparatus has a pair of internally pigmented anterior appendages and a medial appendage or appendages, and the width of the escal bulb is 5.3%–11.4% of standard length. The dorsal fin-rays number 6–8, and the anal fin-rays 5–6. Some authors describe males as having 17–22 upper, and 23–27 lower, denticles on the jaws, while other sources state that no males are known or have been identified. The overall colour is dark brown to black. The mouth is unpigmented, as are parts of the esca. Females attain 20.7 cm in standard length. The attained length of males is unknown.

habitat: A single specimen was caught over a bottom depth of 976

m and a bottom temperature of 4°C in Davis Strait. Elsewhere this species is meso- and bathypelagic. The largest specimen known was taken off Newfoundland in 1968.

biology: It is mostly found below 850 m, where females use their fishing apparatus to catch fishes, cephalopods, and crustaceans.

importance: It is not economically important. distribution: It has been found in a single record from off Baffin Island in Davis Strait at 62º46' N, 59º06' W (ARC 8704914); it also occurs off southwest and southeast Greenland and worldwide.

sources: Regan & Trewavas (1932); Bertelsen (1951); Bertelsen & Pietsch (1977); Orr (1991); Pietsch (2009).

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Chaenophryne longiceps

Oneirodes sp. common names: Unknown. taxonomy: The genus comes from the Greek onerodes (dreamlike), alluding to the small, almost skin-covered eyes or to the general appearance being so unusual as to be dream-like (sources differ). This dreamer was not identified to species. A single specimen was referred to Oneirodes sp. with the esca resembling Oneirodes anisacanthus (Regan, 1925). However, the drawing of the esca does not clearly resemble any illustrated species of Oneirodes, although escae do vary considerably in form. The specimen is listed as Oneirodes eschrichtii Lütken, 1871, in the Zoologisches Museum an der Humboldt-Universität in Berlin (and the figure used here is of that species). More specimens of unidentified Oneirodes have been collected off Baffin Island in the Canadian Davis Strait. The Cosmopolitan Dreamtail, Oneirodes eschrichtii, is recorded from off western Greenland. description: The general structure of this species is summarized

in the family account. The single specimen has the dorsal fin with 6 rays, the anal fin with 4 rays, the pectoral fin with 17 rays, and the caudal fin with 9 rays. The upper jaw has 16–18 teeth on each side. and the lower jaw 12–13 teeth. The vomer bears 3 teeth on each side. The specimen is 19.5 cm long.

Distribution of Chaenophryne longiceps

source: Pietsch (1975).



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Oneirodes sp.

habitat: The single specimen was caught at 960–980 m, and three specimens in the year 2000 at 703–1,090 m and 3.0°C–3.5°C.

biology: Unknown. importance: It is not economically important. distribution: It is found in the southern Davis Strait off southern Baffin Island. Species in this genus are found worldwide in deeper waters.

Distribution of Oneirodes sp.

sources: Bertelsen (1951); Karrer (1973); Pietsch (1974).

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importance: It is not economically important.

Spiniphryne gladisfenae (Beebe, 1932)

distribution: It has been found in Davis Strait on the Can-

Prickly Dreamer, rêveur piquant

common names: A local name is Pigget Mareangler (Danish/

ada-Greenland border as a single record at 62°58' N, 57°42' W (ZMB 23470) and is found in the North and South Atlantic Ocean, the Western Indian Ocean, and the western Pacific Ocean.

Greenlandic).

taxonomy: The genus comes from the Latin spina (spine, thorn) and the Greek phryne (toad). The species is named after the tugboat Gladisfen used by William Beebe (1877–1962), the marine biologist and deep-sea explorer, in trawling around Bermuda. description: The Prickly Dreamer is unique within its family by

the presence of numerous, close-set skin spines covering the body and the fins, which are obvious without a microscope; by the details of the escal morphology; and by the number of dentary teeth. The dorsal fin-rays number 6–7, the anal fin-rays 4–6, and the pectoral fin-rays 15–16, rarely 17. The sphenotic spines are small but sharp, and the profile of the frontal bones on the head dorsally is convex. The dentary teeth number 21–45. The overall colour is black, red-brown to black in preservative. The tips of the distal escal appendages are dark red, and the posterior three-lobed appendage is a bright red-orange, dark in preservative. The distal appendage colour may be due to the presence of blood. Females reach 13.1 cm in standard length; the length of males and larvae is unknown.

habitat: Only a single Arctic specimen has been caught in the Davis Strait, at a depth of 1,950–1,955 m.

biology: This is a bathypelagic species probably feeding on fish and crustaceans.

Distribution of Spiniphryne gladisfenae

sources: Bertelsen (1951); Bertelsen & Pietsch (1975); Karrer (1976); Pietsch & Baldwin (2006).

Spiniphryne gladisfenae



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Ceratias holboelli

Family Ceratiidae

Krøyer, 1845

Northern Giant Seadevil, pêcheur à deux massettes

Seadevils, Poissons-pêcheurs

Brian W. Coad

Seadevils are deep-sea anglerfishes found worldwide. There are only four species, with three in Canadian waters, of which one is in the eastern Arctic. The maximum size is 1.45 m for females. The females have 2–3 rays in front of the soft dorsal fin that are modified as fleshy knobs or caruncles. These caruncles carry areas of bioluminescent tissue. Large mature females have reduced caruncles. Their mouth is vertical to strongly oblique. The eyes in large females degenerate and are covered by tissue, being hardly visible. The dorsal and anal fins have 3–5 rays, usually 4. The caudal fin has 8–9 rays. The esca or bait of the fishing apparatus is simple with only 1–2 filaments. The fishing apparatus or illicium can be withdrawn into a sheath of tissue on the back such that its rear end protrudes from a pore in front of the caruncles. This arrangement probably serves to draw the esca and the enticed prey closer to the mouth. In the skull the parietal bones are large, the frontal bones are not united, and sphenotic spines are absent. The lower pharyngeal bones are reduced and toothless. The skin and fin-rays have spines that are minute in smaller fish, and coarse and evident to the touch in larger fish. Pelvic fins, scales, and a pseudobranch are all absent. The males lack an external fishing apparatus, caruncles, and jaw teeth. Their eyes are well developed, and they depend on sight rather than smell to locate females; therefore, the olfactory organs are minute. The males are parasitic on the females. They have a pair of denticular teeth on the snout tip and two pairs on the lower jaw tip, which are used to attach to females, usually on the ventro-lateral body. The mouth parts fuse with the female, the gut degenerates, the circulatory systems fuse, and the male loses much of its appearance as a distinct fish. The larvae are hump-backed. Larvae can be found in surface waters, but adults are meso- or bathypelagic.

sources: Regan & Trewavas (1932); Bertelsen (1951); Pietsch (1986, 2009).

386

Ceratias holboelli

common names: A local name is Holbølls Storangler (Danish/

Greenlandic). Other common names include Deepsea Angler, Giant Seadevil, Kroyer’s Deepsea Angler Fish, Longray Seadevil, Northern Seadevil, and Twoclub Angler.

taxonomy: The genus comes from the Greek keras (horn) or

keratos (horned), in reference to the illicium. The species is named after Lieutenant Commander Carl Peter Holbøll (or Holboell) (1795–1856), who found a stranded specimen on a beach in Greenland in 1844. This species was identified aboard ship, and no voucher specimen was kept. Its distinctive shape and presence in neighbouring Greenland waters occasion its inclusion here.

description: This species is distinguished by its vertical to very

strongly oblique mouth, by two caruncles when young, and by the esca or bait of the fishing apparatus having one main filament, branched or not. There may be three short lateral filaments on each side of the esca. The two caruncles become minute in fish larger than 40 cm in standard length. The dorsal fin has 3–5 rays, usually 4 (not including caruncles), the anal fin 4 rays, the pectoral fin 15–19 rays, and the caudal fin 8–9 rays, usually the ninth and lowest reduced to a remnant and embedded in the skin (note that some rays in the figure are branched). There are 44–102 teeth on the lower jaw, in 2–3 irregular rows, which are much larger than the upper jaw teeth. The vomer has 1–6 teeth. The female body is covered in conical bony plaques in adults, or small spines in juveniles. The body and the mouth cavity are dark red brown to black, with skin pigment overlapping the bony plaques up to the tops in large females. The lower half of the esca is black, the upper half is white with a dark middle, and the middle process is light grey. Larvae and immature males have an

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Ceratias holboelli

unpigmented body; in mature males it is darkly pigmented. Mature males develop rough skin and become slender, in contrast to the loose-skinned, rounded immature males. Females reach 145.0 cm in length, large for a deep-sea angler, and males reach about 14 cm.

habitat: Apart from strandings, this fish is found from 120 m to 4,400 m, most commonly between 400 m and 2,000 m. Canadian Arctic specimens were from 580–934 m. The larvae are planktonic.

biology: Females may mature at five years and have nearly five

million immature eggs. The eggs may be in a gelatinous raft. Whales are known to eat this fish.

importance: It is not economically important. distribution: It has been found worldwide, including four rec-

ords from cruise data in Davis Strait off southern Baffin Island, and southwest and southeast Greenland and on the Atlantic and Pacific coasts of Canada.

Distribution of Ceratias holboelli

sources: See the family sources and the bibliography.



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sources: Regan & Trewavas (1932); Bertelsen (1951); Bertelsen, Pietsch, & Lavenberg (1981); Pietsch (2009).

Family Gigantactinidae Whipnoses, Tacts géants

Brian W. Coad

Gigantactis vanhoeffeni Brauer, 1902

Vanhoeffen’s Whipnose, tact géant de Vanhoeffen

common names: A local name is Vanhøffens Piskangler

(Danish/Greenlandic).

Whipnoses are found worldwide in deep-sea waters. There are about 22 species, with three occurring in Canada, of which one extends into the eastern Arctic. The maximum size is 43.5 cm in females. These anglerfishes are distinguished by having a fishing apparatus that is often at least as long as the body and can be up to four times as long. The fishing apparatus is not retractable as in the Ceratiidae and Oneirodidae families. Individual species are recognized by finray counts and the details of the esca or bait in the fishing apparatus. The body is elongate in females, with a slender head and an elongate caudal peduncle, unlike many other deep-sea anglerfishes. The eyes in females are very small and covered by a transparent skin layer. The upper jaw extends somewhat beyond the lower jaw. The jaw teeth are elongate and hooked, the lower jaw teeth being especially long and snaggly. The dorsal fin has 3–10 rays, the anal fin 3–8 rays, the pectoral fin 14–22 rays (usually 17–19), and the caudal fin 8–9 rays. Caudal fin-rays in adult females are unbranched, unlike those in many related families. The skin spines are minute or obvious and are also found on fin-rays. Males are free living with well-developed nostrils, minute eyes, and no teeth in the jaws, although there are 3–6 upper and 3–7 lower denticulate teeth outside the jaws. Both males and females lack sphenotic spines. These fishes also have frontal bones that are not united, lower pharyngeal bones that are reduced and toothless, and no scales, pelvic fins, or pseudobranch. The larvae have very large pectoral fins, about half the standard length of the fish. These fishes are predators on other mid-water fishes, cephalopods, crustaceans, and coelenterates, occurring generally between 1,000 m and 2,500 m. The fishing apparatus can be whipped back and forth or held steady while a vibration is passed along to the esca or bait to make it attractive to prey. A long fishing apparatus means that the fish must make a strong lunge to seize the prey, aided perhaps by the streamlined body and the strong tail fin. The lower jaws are connected by an elastic ligament and can be rotated outwards. The long and hooked teeth are used to grasp the prey, which is taken into the mouth by a rapid, inward twist of the lower jaws. Each lower jaw can be used alternately to push the prey down the throat.

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taxonomy: The genus comes from the Greek gigas (big, giant) and tactus (sense of touch, feeling), referring to the “giant touching apparatus” or fishing apparatus, or from the Greek gigantos (gigantic) and aktis (ray). The species is named after Ernst Vanhöffen (1858–1918), a German zoologist on the Valdivia expedition that caught these fish. This species was identified aboard ship, and no voucher specimen was kept. Its distinctive shape and presence in neighbouring Greenland waters occasion its inclusion here. description: This species is distinguished by the details of the

relatively short fishing apparatus (less than 70% of standard length) and by family characters such as the fishing apparatus emerging from the snout tip. The esca or bait has a black spiny extension at its tip, and the escal bulb and distal prolongation have distally flattened papillae. The esca is overall whitish in colour, tuberculate, and covered in small spines in small specimens. Males and larvae are unknown. Only 5–7 dorsal finrays are present, about equal in length; the anal fin-rays number 5–7, and the pectoral fin-rays 17–19. The ventral caudal fin-ray is reduced in size and embedded in the skin. The head and body are black with paired papillae, and the tips of the distal filaments of the esca are bright red. Females reach 62.0 cm in total length.

habitat: This species is bathypelagic and is caught usually between

700 m and 1,300 m, perhaps as deep as 5,300 m and as shallow as 300 m. The Arctic Canadian record was from 359 m.

biology: Unknown. importance: It is not economically important. distribution: It has been found as a single record based on cruise data at 61.1864º N, 64.2105º W, in Davis Strait off Resolution Island, but it is distinctive, and is also found off southwest and southeast Greenland and worldwide in tropical to subarctic waters including off the Atlantic coast of Canada.

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Gigantactis vanhoeffeni

Distribution of Gigantactis vanhoeffeni

sources: See the family sources and the bibliography.



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Scopeloberyx robustus

Family Melamphaidae

(Günther 1887)

Longjaw Bigscale, tête-à-crète robuste

Ridgeheads, Poissons-heaumes

Brian W. Coad

common names: A local name is Småøjet Kogleskælfisk (Dan-

ish/Greenlandic). The French common name was coined by Claude B. Renaud.

taxonomy: The genus comes from the Greek skopelos (said

to be an ancient name for a fish, used as a Lanternfish genus by Baron Georges Cuvier in 1816, now a synonym of Myctophum; the Greek word means “rock,” “peak,” among others) and beryx (an ancient name for a fish, unidentified, taken from the Greek dictionary of Varinus Phavorinus [1460–1537] by the Swiss naturalist Conrad Gesner [1516–65]). The species name is the Latin robustus (robust, stout). The Ridgeheads, Bigscales, or Bluebacks are small bathypelagic fishes of all oceans and seas except the High Arctic and the Mediterranean. There are about 36 species, of which 18 occur in Canada. One species is found in Arctic waters. The maximum size is about 15 cm. Family members have characteristic blunt heads that are sculptured and ridged. The bones and scales are paper thin, particularly in the deeper-water species. The head canals are highly developed, making cavernous structures in the superficial bones, are covered with epidermis rather than bone stretched between paper-thin bony partitions, and underlie sensory structures that appear as epidermal, embossed lines. The scales are large, cycloid, and easily detached. There is no true lateral line, and only 1–2 pored scales behind the head. The dorsal fin has 1–3 weak spines before 9–18 soft rays. The pelvic fins lie under the pectoral fins (thoracic or sub-thoracic position) and have 1 spine and 6–8 soft rays. The anal fin has 1 spine and 7–12 soft rays. Procurrent spines are developed where the caudal fin meets the body dorsally and ventrally, and number 3–4. There are 19–36 gill rakers, often with many spines, and 7–8 branchiostegal rays. The teeth are minute and found in bands or in single rows. The overall colour is dark brown to black. Ridgeheads are commonest below 1,000 m when they are adult, and are rarely found above 100 m except as young and larvae. They may descend below 4,000 m. Little is known about their biology, but they feed on zooplankton and gelatinous organisms and are eaten by marine mammals and the larger fishes. They are not economically important as they are only caught on scientific fishing expeditions using special deep-sea nets. These fishes are fragile, and preserved specimens are often damaged.

source: Ebeling & Weed (1973).

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description: This species is the only member of its family in Arc-

tic waters and is distinguished by the ridged and sculptured head. In this species the ridges are smooth and not crest-like. Scale rows number 24–27 from the end of the post-temporal bone to the caudal base (28–33 from the nape); scales are often lost, but scale pockets are distinct. Cheek scales number 3–4 but are also often lost. The dorsal fin has 1–3 spines and 9–13 rays, usually 11; the anal fin has 1 spine and 7–9 soft rays, usually 8; the pectoral fin has branched rays numbering 11–15; and the pelvic fin has branched rays numbering 6–8. Total gill rakers number 19–25, and branchiostegal rays 8. A supramaxillary bone is present, the preopercle is smooth, and the spine between the nostrils is inconspicuous. The eye measures more than nine times in head length, and the horizontal distance between the pectoral and pelvic fin insertions is 5% or less of standard length. The body is dark brown in colour, and the head, mouth cavity, and branchial cavity are black. The fins are pigmented. The species attains 10.0 cm in standard length.

habitat: Adults are mostly found mesopelagically and bathypelagically at 500–4,740 m, while younger fish are in shallower waters. The eggs and larvae are planktonic. The single Canadian Arctic specimen was caught at 1,408–1,417 m and 3.4ºC. biology: Little is known of the biology of this species. Maturity

may be attained at 24 mm in February–March in the western Atlantic Ocean, but some fish are not mature at over 60 mm.

importance: It is not economically important. distribution: The species was found in Davis Strait as a single record at 62°3'54" N, 60°6'24" W (CMNFI 2002-0047). It is also found in southwest and southeast Greenland and worldwide except for the High Arctic and the Mediterranean Sea.

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Scopeloberyx robustus

source: Kotlyar (2004b).

Distribution of Scopeloberyx robustus



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Anoplogaster cornuta

Family Anoplogastridae

(Valenciennes, 1833) Fangtooth, ogre

Ogrefishes, Ogres

Brian W. Coad

common names: A local name is Troldfisk (Danish/Greenlandic). Other common names include Common Fangtooth, Horned Fang-tooth Fish, and Ogrefish. taxonomy: The genus comes from the Greek ana (up) and oplon

(shield), or anoplos (unarmed) and gaster (belly). The species name comes from the Latin cornutus (horned).

description: This species is distinguished by obvious mucous

This family, also called Fangtooths or Sabretooths, is found bathypelagically in most oceans. There are only two species, of which one is found in Atlantic and eastern Arctic Canada. The maximum size is about 16 cm standard length. The head has an ogre-like appearance and is large, about onethird of body length. It is sculptured with large mucus cavities covered by a thin layer of skin. The short, deep body is covered with small, thin cycloid scales that have many spines. Each scale is on a short pedicel embedded in the skin. The lateral line is an open groove occasionally covered with scales at intervals. There are no spines in the fins except at the caudal base. The pelvic fin has 7 soft rays. The large jaws have characteristic and numerous fangs that are depressible in young but fixed in adults. There are 8–9 branchiostegal rays. Young are different from adults in having a big eye, long head spines, smaller teeth, and slender, lath-like gill rakers, and they were once described as a distinct species. These fish are remarkable for their ability to survive in aquaria for up to 23 days. Most meso- and bathypelagic fish die within minutes or hours of capture; therefore, most of our knowledge of them is based on dead specimens. They feed on crustaceans and fishes and are found down to 5,000 m. They are not economically important.

sources: Kotlyar (1986a, 2003).

cavities covered by skin on the head and by scales on pedicels. The dorsal fin has 16–20 rays, the anal fin 7–10 rays, and the pectoral fin 13–17 rays. Gill rakers number 14–23 and are long and slender in young, shorter and tooth-like in adults. The large lower-jaw fangs fit into sockets in the palate. Young fish have long parietal and preopercular spines, which gradually reduce in size with growth. Its colour is dark brown to black overall with a blue iris. The lateral line may be darker or lighter than the surrounding areas. The young are silvery with transparent fins. The species attains about 16.0 cm in standard length.

habitat: It is found between 2 m and 4,992 m, only larvae being

as shallow as 2 m, and sometimes in small schools or solitary. In the Canadian Davis Strait it has been taken at 703–1,090 m and 3.5°C–3.8°C, and as deep as 1,232 m. Adults live deeper than the young.

biology: Its food is large fish and crustaceans, and sometimes

squid, which are eaten whole and infrequently. Aquaria observations indicate that Fangtooths locate food simply by having to bump into it and sensing it chemically. They are “sit-and-wait” predators. A prompt reaction with the fangs secures the prey. The pectoral fins are used to fan water over the gills while the fish is digesting large prey that blocks the mouth. Fangtooths are eaten by larger fishes. They swim by sculling with the pectoral fins, only using the caudal fin when disturbed.

importance: It is not economically important. distribution: The species is found in Davis Strait off southern Baffin Island and southwest and southeast Greenland, and worldwide in warmer waters including the Atlantic and Pacific coasts of Canada.

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Anoplogaster cornuta



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Family Trachichthyidae Slimeheads, Hoplites

Brian W. Coad

Distribution of Anoplogaster cornuta

sources: See the family sources and the bibliography.

The Slimeheads or Roughies are found worldwide and have about 35 species, including four reported from Canada, one of which extends into the Arctic. The maximum size is about 60 cm. The head bears large mucous cavities covered by skin, hence their name. The cavities are separated by bony crests. There is an obvious spine at the angle of the preopercle bone and in the post-temporal bone of the side of the head. The belly has a midline ridge formed from modified scales or scutes. Lateral-line scales are enlarged also. The scales on the body are thin and cycloid or thick and covered with spines. The dorsal fin has 3–8 spines and 9–19 soft rays. The anal fin has 2–3 spines and 8–12 soft rays. The pelvic fin has 1 spine and 6–8 soft rays. The jaw teeth are in small bands. Some of these fishes are luminescent. The overall colour is reddish orange, pink, or dusky silver. These fishes are found bentho-pelagically, mostly in deeper water from 75 m to 1,000 m but may be found as shallow as 10 m and as deep as 1,900 m. They feed on fish and crustaceans. Some species are quite abundant and are sold fresh, dried, or salted, or they may be turned into fish-meal or oil. They are caught by deep trawling and line trawling but are easily overfished.

sources: Kotlyar (1986b, 1996).

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fin 1 spine and 6 soft rays. The lateral-line scales are enlarged and diamond shaped, being twice as high as they are wide, and number 28–34. The ventral scutes number 13–25 but may be indistinct in some fish. Total gill rakers number 17–22. Its colour is a bright brick red, sometimes a silvery pink, with the mouth, gill cavity, and peritoneum being bluish black. The fins are mostly clear. The species attains 75.0 cm in total length and 7.0 kg in weight.

Hoplostethus atlanticus Collett, 1889

Orange Roughy, hoplostète orange

habitat: A bathypelagic species, it is found from 180 m to 1,809 m

Hoplostethus atlanticus

common names: A local name is Orange Savbug (Danish/

Greenlandic). Other common names are Atlantic Slimehead, Deepsea Perch, Orange Ruff, Red Roughy, and hoplostète rouge.

taxonomy: The genus comes from the Greek hoplon (armour, weapon) and stethos (breast). The species name is the Latin atlanticus (of the Atlantic). description: The large mucous cavities on the head and the enlarged diamond-shaped lateral-line scales are distinctive. The dorsal fin has 4–7 spines and 15–19 soft rays, the anal fin 3 spines and 10–12 soft rays, the pectoral fin 15–20 rays, and the pelvic

over hard and rough bottoms. Most fish are caught at 4.5°C–6.5°C and 600–1,400 m commercially. In the northeast Atlantic Ocean early juveniles are found in relatively shallow habitats, juveniles are deep demersal, maturing adults rise in the water column, and adults become deeper with age. Canadian specimens north of 55° N were mostly caught below 700 m, increasing with depth to below 1,200 m at temperatures of 1.9°C–4.1°C. However, the eggs require temperatures greater than 6°C to complete development. This species forms dense spawning aggregations over steep continental slopes, ocean ridges, and seamounts. The brief larval dispersal period suggests that the adult migration of this long-lived species ensures gene flow over wide areas. Canadian captures occur in less than 3 sets per 1,000 observed, although they are more common, by about five times, north of 55° N than to the south.

biology: Orange Roughies feed on benthopelagic and mesopela-

gic fish, crustaceans, and squid. Juveniles feed mainly on crustaceans. However, isotopic analyses in the northeast Atlantic suggest more of a benthic diet, and metabolic rates are low, associated with their long lives. Maturity is attained at 5–32 years, and maximum age is reported as 149 years, one of the longest-lived

Hoplostethus atlanticus



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fish known. The minimum population doubling time is more than 14 years, and so they are easily overfished. The size at first maturity is 44–51 cm in total length elsewhere. Orange Roughies are synchronous annual spawners, with males appearing to spawn over a one- to two-week period, and females for up to one week. Spawning occurs from late January to early March in the northeast Atlantic and may well be the same for the Canadian populations. The eggs are orange red and pelagic but later sink to the sea bed, retaining young in the adult habitat.

Family Gasterosteidae Sticklebacks, Épinoches

Chantelle D. Sawatzky and James D. Reist

importance: This species is the focus of major and recent deep-

sea commercial fisheries off New Zealand, Australia, Namibia, and Chile, and there are some smaller fisheries elsewhere including off Iceland. Few of these fisheries have been exploited in a sustainable way, and this long-lived species is vulnerable to overfishing. It is marketed fresh and frozen. It is not economically important in Canada and is taken as a by-catch in bottom gear for Greenland Halibut and shrimps.

distribution: The species is found in Davis Strait off southern Baffin Island, southwest and southeast Greenland, and worldwide in the Atlantic, Pacific, and Indian Oceans. Most Atlantic coast captures in or near Canadian waters are centred at 63° N. However, inadequate developmental temperatures may mean that these fish are expatriates, dispersing as juveniles and adults from Iceland.

Distribution of Hoplostethus atlanticus

Sticklebacks are found in northern marine and fresh waters. Over 60 species have been described, but less than 12 are considered valid. There are 5 species in Canada, of which 2 are found in the Arctic. These fishes are small, less than 18.0 cm in total length, and usually much smaller. A high degree of polymorphism characterizes most taxa; thus, variation both within and among locations is very high. Most of this variation is considered to result from ecological adaptation and not from fundamental taxonomic differences. Thus, under a strict taxonomic species concept, variant groups are considered to be species complexes. General morphological characteristics include small size, a moderately elongate body, a slender caudal peduncle, bony scutes instead of scales often present on the sides, 3–16 spines in front of a dorsal fin with 6–14 rays, a pelvic fin with 1 spine and 0–3 rays, 12 caudal fin-rays, 3–4 branchiostegal rays, 28–42 vertebrae, and an incomplete circumorbital ring. The numbers of fin-rays, predorsal spines, body scutes or plates, gill rakers, and vertebrae are highly variable. The biology of these fishes has been studied extensively, with many papers and several books being written about them. Although they have no economic importance, their colour, body form, maximum age, development of spines and plates, speciation, and complex biology have attracted this research. Canada has giant unarmoured, white, black, and trophic forms, and all species of Sticklebacks have an intricate reproductive behaviour involving courtship and nest building. Some populations and species live entirely in fresh water or in sea water, while others are anadromous.

sources: Kulka, Themelis, & Halliday (2003); FAO (2011); Berg-

sources: Cox (1922); Wootton (1976, 1984); Coad (1981); Bell &

stad (2013); Trueman, Rickaby, & Shephard (2013).

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Foster (1994).

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Gasterosteus aculeatus

can be used locally to differentiate marine and anadromous forms from co-occurring freshwater life-history types. Typically this species has 3 dorsal spines, with the last one being short, although the range is 2–4. The second dorsal fin has 7–14 soft rays. The anal fin is shorter based than the soft dorsal fin, has 1 spine and 6–11 soft rays, and originates several rays posterior to the origin of the soft dorsal fin. The pectoral fin has 8–11 rays. The pelvic fin spine is long and serrated, and there is one soft ray. All spines are depressible and can be locked upright. The gill rakers number 17–27, and the vertebrae 29–35. Broad gill membranes are attached to the isthmus. The body coloration is silvery green to bluish black for marine and anadromous fish, mottled brown or greenish for freshwater fish, and yellow, white, or silver ventrally. Spawning coloration in males is blue or green, with bright blue eyes and a red or orange throat and belly. Females develop a pink tint on the throat and belly. The species attains 10.2 cm in total length but rarely exceeds 7.5 cm.

common names: Local names are Kakilaychok, Kakilishek,

habitat: The species is found in marine, brackish, and fresh waters;

Gasterosteus aculeatus Linnaeus, 1758

Threespine Stickleback, épinoche à trois épines

Kakilusuk, and Katilautik (Inuktitut); and Kakilisak, Pingasunik, and Kipinartulik (Greenlandic). Other common names include Banstickle, Common Stickleback, Eastern Stickleback, New York Stickleback, Pinfish, Saw-finned Stickleback, Tiddler, and Twospine Stickleback.

taxonomy: The genus comes from the Greek gaster (belly) and osteon (bone). The species name is the Latin aculeatus (spined), both genus and species names referring to the pelvic girdle and the spine. Gasterosteus insculptus Richardson, 1855, described from Northumberland Sound, is a synonym. Threespine Stickleback populations as a group exhibit a large degree of morphological variation that includes variations both within and among life-history types, and with respect to ecological niche differentiation. Thus, though treated here as a single species, this is actually a species complex that remains mostly unresolved at present. Taxonomic uncertainty exists for the relationship between the marine/anadromous fully plated form and the partially plated freshwater form. The two forms are generally reproductively isolated but may interbreed when occurring sympatrically (e.g., Little Campbell River in southwestern British Columbia). Additionally, the marine form has repeatedly colonized freshwater environments. These repeated colonizations across either space or time have resulted in many morphologically distinct populations. The disjunct distribution in the Atlantic and Pacific regions further confounds the issue as there is, at least presently, no possibility of gene flow between these regions. An analysis of allozyme variation between Asian, North American, and European populations has supported the recognition of a single taxonomic unit as a species complex. Future work may indicate that recognition of some subspecies is warranted. description: The body is somewhat elongate with a short snout.

The caudal peduncle of the anadromous fish has a bony lateral keel. Lateral bony plates are usually present in marine and anadromous forms, numbering up to 37, but only 0–9 in some freshwater forms. The presence of both lateral keels and large numbers of lateral plates



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only the seasonally anadromous fish are found in the marine waters of the Arctic and to 805 km offshore in the Gulf of Alaska. Generally it occurs in shallow vegetated areas over mud or sand substrates to a depth of 27 m, in both benthic and pelagic habitats. It is found more often in open water than amongst vegetation. Female fish in tidal marshes are most active during the day and are quiescent at night. Salinity tolerance varies between forms and populations and with water temperature and physiological state. Thermal tolerance also varies between populations.

biology: Its diet consists of copepods, cladocerans, ostracods,

chironomid larvae and pupae, ephemeropteran nymphs, oligochaetes, molluscs, algae, and the eggs and fry of their own species in fresh waters. In marine waters amphipods, isopods, and polychaete annelids are also preyed upon. Threespine Sticklebacks are preyed upon by fish-eating birds (gulls, terns, ducks, grebes, kingfishers, herons), piscivorous fish (trout, char, Northern Pike, Yellow Perch, Pikeminnow, cod, and other gadoids), and even seals. They are eaten by Herring Gulls in western Rupert Bay, James Bay. Spines and body plates are concentrated at the anterior end of the body where most predators strike. Even Threespine Sticklebacks caught by the tail are often manipulated by predators to be swallowed head first; the plates protect against injury while the spine serves to increase the apparent body size (thus fewer predators can take them) and to promote release. The lifespan varies by population with few individuals surviving past the age of two years. Anadromous fish likely mature by the age of one year. High post-spawning mortality occurs, but some fish may spawn a second year. No obvious sexual dimorphism is present other than coloration at spawning times. Reproduction is oviparous and occurs from April to October over the range of the species. The anadromous form spawns in fresh or brackish water from May to July in northern areas (the degree to which this occurs in Arctic ice-covered waters is unknown). Spawning substrates range from silt to rock. On the substrate the males construct barrel-shaped nests, open at each end, out

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Gasterosteus aculeatus

of vegetation and debris held together by a secretion from the kidney. The male courts the female by performing a zigzag courtship dance, leading her into the nest. A responsive female adopts a submissive head-up position revealing the egg-swollen belly. The male pokes his snout at the nest to indicate its position to the female, tipping his head sideways to display the bright red throat. The male jabs the female with his snout through the nest wall to stimulate egg release. Once the female has spawned, the male chases her away and then fertilizes the eggs in the nest. Males guard the nest, fan the eggs, and care for the fry. A male may have more than one set of eggs at different stages of development in the nest at one time. Sneaking and egg stealing by other males may occur, as confirmed by DNA “fingerprinting.” Females lay between 50 and 200 eggs at a time and usually spawn several times with different males in one season. A nest may have up to 1,026 eggs. The eggs are larger than those of other Sticklebacks (diameter about 2.0 mm) and are sticky. Females often cannibalize eggs, and the male lifespan is shortened through stress in defending against these and other attacks. Common parasites include trematodes, copepods, nematodes, molluscs, acanthocephalans, and cestodes. A study of parasites infecting Threespine Stickleback populations in Poland identified 51 parasitic species. The cestode Schistocephalus solidus reduces the reproductive potential of females.

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importance: The species is widely used by the scientific community in field and laboratory studies, the high variation and ecological associations being of importance in understanding evolutionary and adaptation processes in fishes. Ecological studies of this species in the Arctic are generally lacking especially in coastal areas. It is likely very important in coastal food webs as a forage species for piscivorous birds and fishes, including chars and other nearshore anadromous predators in marine waters. distribution: It is found in Cumberland Sound, Frobisher Bay, Ungava Bay, Hudson Strait and Bay, and James Bay. Only occasional and isolated occurrences are confirmed from the marine waters of the western Arctic including the Alaskan and Yukon north slopes and Northwest Territories. The easternmost record for the Beaufort Sea is Simpson Lagoon in Alaska. Accordingly, western Canadian Arctic records from the Beaufort Sea area require confirmation as to species (i.e., reports may represent Ninespine Stickleback). The westernmost records in the Canadian Arctic, from the Beaufort Sea, are from Byers and Kashino (1980), who also recorded, and presumably distinguished accurately, Ninespine Sticklebacks, in their study. These records are assumed to be vagrants if they are valid. The Northumberland Sound record is from Richardson (1855) (as Gasterosteus insculptus).

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Nearly circumpolar in distribution, it is absent, however, from the Arctic Siberian coast and much of the Arctic North American coast. The complex occurs in Europe (northern areas south to Syria) and North America, where it has a disjunct range in the latter. In the west the species occurs from Baja California north to the Aleutian Islands, St Lawrence Island, coastal southern and western Alaska, and the Chukchi Sea. It is present in eastern Arctic waters from Baffin Island and all the coasts of Greenland south to Chesapeake Bay. Freshwater populations occur inland to the Bristol Bay region of Alaska in western North America, and inland from the Atlantic coast, including Lake Ontario and the Ottawa and St Lawrence Rivers, north to Baffin Island.

Pungitius pungitius (Linnaeus, 1758)

Ninespine Stickleback, épinoche à neuf épines

Pungitius pungitius

common names: Local names are Kakidlautidlik, Kakilahaq, Kakilasak, Kakilishek, Kakilusuk, and Kakiva (Inuktitut); and Natagnak (Alaska). Other common names include Many-spined Stickleback, Pinfish, Tenspine Stickleback, and Tiny Burnstickle.

Distribution of Gasterosteus aculeatus

taxonomy: The genus and the species name are the Latin pungitius (pricking), in reference to the spines. Ninespine Sticklebacks exhibit a large degree of variation in morphology and life history. Although listed here as a single species, this is actually a species complex that for the most part remains unresolved with respect to the number of species and to their taxonomic limits and distributions. Some literature proposes that the North American Ninespine Sticklebacks be recognized as P. occidentalis (Cuvier, 1829), described from Newfoundland, based on a study of allozyme variation. A broader molecular study is needed before this name can be accepted. An alternative proposal recognizes the North American form as P. p. occidentalis based on morphology. As this is unresolved presently, traditional nomenclature is used here.

sources: Richardson (1855); Hynes (1950); Freeman (1965);

description: The dorsal spines number 6–13 (typically 9, usually

Hagen (1967); Byers & Kashino (1980); Audet, Fitzgerald, & Guderley (1985); Haglund, Buth, & Lawson (1992a); McKinnon & Rundle (2002); Hart (2003); Le Comber, Faulkes, Formosinho, & Smith (2003); Morozińska-Gogol (2006); Sawatzky et al. (2007).



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8–11) and are angled alternately left to right. The second dorsal fin has 8–13 soft rays. The anal fin has a stout spine and 6–11 soft rays. The pectoral fin-rays number 10–11, and the pelvic fin has 1 spine and 1 soft ray. Fish with reduced or absent pelvic skeletons or spines are not uncommon. The gill rakers number 10–14, and the vertebrae 30–35. The caudal fin is slightly forked. Small plates are present along the bases of the dorsal and anal fins. The gill membranes are joined and free from the isthmus. The small, rounded lateral plates number from 0 to 35, with completely plated fish having 28–35 including the caudal keel. The caudal peduncle has a well-developed lateral keel made up of 11–22 rectangular bony plates. The body coloration is brownish green dorsally and yellowish silver ventrally, with indistinct vertical bars on the sides. Breeding males turn black on the ventral surface and under the chin and have white or pale blue pelvic fin membranes. Breeding females are black or brown dorsally but silvery on the ventral surface. The species attains 9.0 cm in total length, but rarely exceeds 6.5 cm.

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Pungitius pungitius

habitat: Ninespine Sticklebacks are found in marine, brackish,

and fresh waters. Weedy habitat is preferred. In the marine environment (during summer) they occur abundantly in nearshore waters, marshes, and estuaries and are found in both benthic and pelagic habitats. They die when pools dry up and are then an important food for birds. In the Alaskan Beaufort Sea they were most abundant at collection stations within 2 km of the coast, and most frequent at temperatures of 2°C–4°C and a salinity greater than 27‰. Given the likely preference for warmer and freshened marine waters, during Arctic summers this species probably does not extend far into marine areas. Ninespine Sticklebacks, particularly in southerly locations, exhibit a wide range of life histories including wholly freshwater and anadromous types with fairly long seasonal occupancy of marine waters. In the Arctic, seasonal occupancy in marine waters corresponds with open water, likely during July and August. The anadromous form spawns in fresh water in spring to summer (May–July), migrates to marine waters for summer feeding, and returns to fresh water in fall to overwinter. Some Hudson Bay populations are believed to winter at sea.

biology: Much of the known biology of this species refers to

freshwater populations, even in Arctic waters. Its diet consists of small crustaceans and aquatic insects, occasionally fish eggs and fry. Predators include other fishes such as Arctic Char, and birds. It is fed on by Bowhead Whales in the Alaskan Beaufort Sea and by Herring Gulls in western Rupert Bay, James Bay. The species is short lived with a lifespan to age 4+ years and sexually mature at the age of one to two years. Females grow faster and live longer than males. Males rarely live past the age of three years due to high post-spawning mortality. Females may live to age 4+. Reproduction is oviparous. Males build nests in shallow water among weeds and off the bottom in densely vegetated areas. The nests consist of aquatic vegetation and debris held together by a

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secretion from the kidney. Each nest is a tunnel open at each end and about 3–4 cm long. Some nests are open only at one end, and the fish have to turn around to leave. Females are enticed into the nest to deposit their eggs. The male fertilizes the eggs, fans and aerates them, and guards the nest and subsequent fry until they are free swimming. Early fry leaving the nest are caught by the male in his mouth and spat back into the nest. Males may seize and drag away other small fishes and use charges, chases, nips, and bites to defend the nest and young. Both sexes may spawn multiple times each season with different individuals. In south-central Alaska, females were found to produce 63–267 eggs per clutch. The egg diameter is 1.5 mm. Ninespine Stickleback may harbour up to 33 species of parasites including protozoans, trematodes, cestodes, acanthocephalans, molluscs, branchiurans, and copepods. The cestode Schistocephalus pungitii is known to inhibit reproduction in both males and females.

importance: It is of limited importance to humans although it

has been used as human or dog food and as a source of oil. Similarly to Threespine Sticklebacks, morphological variation and ecological associations are scientifically important in understanding evolutionary and ecological processes. Ecological studies relevant to this species in the Arctic are generally lacking, particularly in coastal areas; however, it is likely very important in local food webs as a forage species when it is abundant. Predatory birds and fishes, including chars, prey upon them in nearshore coastal areas.

distribution: It is found in Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson Bay, James Bay, Foxe Basin, Dease Strait, Melville Sound, Bathurst Inlet, Coronation Gulf, Dolphin and Union Strait, Amundsen Gulf, Beaufort Sea, southern Cornwallis Island (Resolute Bay), western Victoria Island, and northern Banks Island.

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The species is circumboreal in distribution especially in waters inland to about 600 m elevation. It is found along the Arctic and Atlantic coasts south to New Jersey in the western Atlantic. This species is distributed in coastal waters from southwestern Alaska north and eastwards to eastern Canada, with adjacent distributions often extending far inland (e.g., much of Alaska, extreme northeastern British Columbia, Alberta, and the Mackenzie Valley to the northern Great Lakes). It occurs in northern Europe, Asia, and Eurasia from Siberia south to Korea and the Seas of Okhotsk and Japan. As noted, western Canadian records of Threespine Stickleback may be this species. Point distributions on the map likely under-represent the true Arctic distribution. At present, it likely does not occur much further north than the southern fringes of the northern Arctic islands at about 75° N; however, this may change with ameliorating climates.

Distribution of Pungitius pungitius

sources: Johansen (1926); McPhail (1963); Hanek & Threlfall

(1970); Griswold & Smith (1972); Cameron, Kostoris, & Penhale (1973); Eddy & Underhill (1974); Lowry & Burns (1980); Becker (1983); Curtis (1981); Thorsteinson, Jarvela, & Hale (1991); Haglund, Buth, & Lawson (1992b); Keivany & Nelson (2000); Heins, Johnson, & Baker (2003); Heins, Ulinski, Johnson, & Baker (2004); Sawatzky et al. (2007); Heins & Baker (2010); Aldenhoven, Miller, Corneli, & Shapiro (2010); Gallagher & Dick (2011).



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Family Scorpaenidae Scorpionfishes, Scorpènes

Brian W. Coad

The Scorpionfishes, Redfishes, or Rockfishes are found worldwide in temperate to tropical seas and rarely in fresh water. There are about 388 species, mostly in the Indo-Pacific Ocean. There are 51 species in Canada, mostly on the Pacific coast, with probably three in Arctic waters. The maximum size is 105 cm. Field identification of the Arctic species is difficult, and accurate identification requires dissection and/or molecular techniques. There is evidence of hybridization and introgression, which further complicates identifications. Old records under the name Sebastes marinus (Linnaeus, 1758) from Arctic waters may be any of the three species, but difficulties of identification in the field make these records uncertain. The head has ridges, spines, tabs, cirri, skin flaps, and tentacles on the top and sides. There are 1–2 (usually 2) opercular and 3–5 (usually 5) preopercular spines. A suborbital stay is usually attached to the preopercle. The eye is moderate to large in size. The mouth is large in a broad head. Teeth are villiform in both jaws and on the vomer. The colour is often reddish or brown. The dorsal fin is continuous with 7–18 spines, often separated by a notch from the 4–18 soft rays. The anal fin has 1–4 (usually 3) spines and 3–14 (usually 5) soft rays. The pectoral fin is large with 11–25 rays. There is 1 spine and 2–5 (usually 5) branched rays in the pelvic fin. The caudal fin is rounded or emarginate. The scales are usually ctenoid. These fishes are found from the shallows down to over 2,000 m. Most Scorpionfishes are fertilized internally. Some give birth to live young, and others lay eggs in a gelatinous balloon up to 20 cm in diameter. Sebastes species are live bearers, and their larvae stay near the surface until they are 3.0–5.0 cm long and move down in the water column. Larval drift and juvenile migrations carry redfish from the Irminger Sea off southeast Greenland into Davis Strait. Redfish larvae are an important food for Greenland Halibut and Atlantic Cod. In west Greenland the catch of redfish in the 1992 offshore shrimp fishery was second after the shrimp itself, at 157 kg wet weight/sq km. Many species are economically important. The three Atlantic coast redfishes (Acadian, Deepwater, and Golden) ranked fourth

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Sebastes sp.

in importance in Canada among commercial fishes by weight, 73,500 t in 1988. The Atlantic Canada total in 2011 was 13,800 t, worth $10,577,000. Annual catches in the northwest Atlantic (FAO Statistical Area 21) have been as high as 215,000 t in 1960–9 and as low as 22,000 t in 2008. Commercial fishing of these species occurs off the northern tip of Labrador. In western Baffin Bay and Davis Strait 26 t were caught in 1975, and in NAFO Subarea 0A off Baffin Island in 1976, 126 t were caught.

sources: Barsukov (1968); Desrosiers, Sévigny, & Chanut (1999);

Pikanowski, Morse, Berrien, Johnson, & McMillan (1999); COSEWIC (2010b); FAO (2011).

Sebastes fasciatus Storer, 1854

Acadian Redfish, sébaste acadien

common names: A local name is Amerikap Suluppaagaa

(Greenlandic). Another common name is Labrador Redfish. Ocean Perch and Redfish are market names, and the three redfish species are not distinguished for commercial purposes. This species is referred to as Beaked or Sharp-beaked Redfish when grouped with the Deepwater Redfish and not distinguished.

taxonomy: The genus comes from the Greek sebastos (magnificent, venerable). The species name is the Latin fasciatus (banded). description: This species is distinguished from other northwest

Distribution of Sebastes species

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Atlantic and eastern Arctic redfish species by a combination of overlapping external characters. It is easily misidentified when its range overlaps with the two closely related species from eastern Canada. One useful character is internal and requires dissection to see. This involves the muscles attaching to the gas bladder. In this species there are one or two muscle heads with tendons passing between the third and fourth ribs, or more rarely (5%) the fourth and fifth ribs;

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Sebastes fasciatus

the muscle is narrow; and the posterior tendon has three branches and is attached to the eighth, ninth, and tenth vertebrae. This species is a less robust redfish and has a larger eye than the Golden Redfish. The dorsal fin has 13–16 spines and 12–16 soft rays, usually 13 or 14. The anal fin has 3 spines and 6–8 soft rays, usually 7. The pectoral fin-rays number 18–20, usually 19. The tip of the pectoral fin usually does not reach the anus. The gill rakers usually number 36 or more although the range is 32–38. The total vertebrae number 29–31, modally 29. The scales in series along the flank just below the lateral line number about 60–70, and the lateral-line scales are larger. The downward angle of the third preopercular spine is usually less than 50°. The occipital–nuchal ridge is usually fused on both sides. The knob at the tip of the chin is long and sharp. Larvae have 26–42 ventral row melanophores. The overall colour is a bright orange red with green-black blotches on the body and iridescent green flecks or markings. The pelvic and anal fins are a deep red, with other fins being a lesser red. The belly is a pale red but fades to white after death. The species reaches 45.7 cm, usually 30.0 cm, in length and 11.2 kg in weight.

habitat: This is a shallower-water species than its two relatives,

from 1 m down to 592 m, generally at 150–300 m. It is found over rocks or other hard bottoms and over clay-silt or mud, and infrequently on sand. Vertical migrations of larvae may be less extensive than those of Deepwater Redfish. Larvae are found at preferred depths of 11–30 m by day and 10 m or less by night but can be observed in the upper 200 m of the water column. Juveniles in the Gulf of Maine prefer 5ºC–10ºC temperatures, while adults prefer 4.5ºC–7.0ºC. Juveniles remain pelagic until they attain 40–50 mm, after about four to five months of growth. Smaller fish live in shallower waters (75–175 m) and, as they grow, occupy deeper water.



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biology: The biology of redfishes has not yet been fully worked

out because of the difficulties in separating the species. Biology in the Arctic is not known, and the data come from Atlantic Canada and the northern United States. Often the biology of the “beaked” redfishes, S. fasciatus and S. mentella, are combined in the literature. Its food is crustaceans, in particular euphausiids and decapods, and, when larger, it takes fishes at night when it rises in the water column to capture pelagic prey. It is eaten by a wide variety of other fishes. This is a slow-growing and long-lived species that may attain about 58 years per one report, 75 years per another. Its growth is slower in northern parts of the range. Females grow faster than males after age 10 years. Acadian Redfish are late maturers at about 7–10 years, although in southern waters some mature as early as 2 years for males and 3 years for females. Strong year classes occur every 5–12 years. Spawning occurs in March–July in southern waters, and about 15,000–20,000 live young are produced. Absolute fecundity for this species, and its relative the Deepwater Redfish, may reach 107,000 larvae.

importance: It is economically important in the Georges Bank

and Gulf of Maine area. The catch in 1959 was 389,000 t (presumably including all redfish species) but is now less as quotas have been imposed to protect stocks; since 1994 the landings in the Gulf of Maine have been limited to 500 t. The Committee on the Status of Endangered Wildlife in Canada has assessed it as “Threatened.”

distribution: Distribution is centred further south than that of its relative S. mentella but does reach southern Arctic waters as defined here, off the northern tip of Labrador and in Hudson Strait according to literature reports. It is rare off the Labrador Shelf and southeast Greenland. The southern distribution is the Gulf of Maine.

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It is also found rarely off Iceland. Redfishes are mapped without distinction of species because these are often not clearly identified in the literature. No S. fasciatus are included in the general map under the family account.

sources: See under Sebastes mentella.

Sebastes mentella Travin, 1951

Deepwater Redfish, sébaste atlantique

common names: A local name is Suluppaagaq Itisoormiu (Greenlandic). Another common name is sébaste du nord. Ocean Perch and Redfish are market names, and the three redfish species are not distinguished for commercial purposes. This species is referred to as Beaked or Sharp-beaked Redfish when grouped with the Acadian Redfish and not distinguished. taxonomy: The species name comes from the Latin mentum (chin) and -ella (small). description: This species is distinguished from other northwest

Atlantic and eastern Arctic redfish species by a combination of overlapping external characters. It is easily misidentified when its range overlaps with the two closely related species from eastern Canada. One useful character is internal and requires dissection to see. This involves the muscles attaching to the gas bladder. In this species there are one or two muscle heads with tendons passing between the second and third ribs, the muscle is narrow, and the posterior tendon is not branched and is attached to the seventh vertebra. The dorsal fin has 14–15 spines and 12–16 soft rays, usually 14 or 15. The anal fin has 3 spines and 7–11 soft rays, usually 8 or more. The lower unbranched pectoral fin-rays are usually 8 or less (rarely 9), total rays number 19–20, and the tip of the pectoral fin mostly extends beyond the anus. The gill rakers usually number 35 or less although the range is 32–37. The total vertebrae number 30–32, modally 30. The downward angle of the third preopercular spine is usually more than 50°. The occipital–nuchal ridge is usually separated on one or both sides. The knob at the tip of the chin is long and sharp. Larvae have 5–10 ventral row melanophores. The overall colour is a bright scarlet red. The regions between the pectoral base and the operculum, and the cheeks and the pelvic region, are silvery. Young fish are silvery pink and have a dark band between the soft dorsal and anal fins that tapers anteriorly. The species reaches 58.0 cm, usually 40.0 cm, in length and 8.5 kg in weight.

habitat: This bathypelagic redfish is found further north, further offshore, and in deeper water than related species and is the principal species in Arctic waters. The more fusiform morphology

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of this species may enable it to move further than Acadian Redfish. It is found over rock or clay and silt bottoms from 100 m to 1,100 m, being both benthic and bathypelagic, but it also appears to make daily vertical feeding migrations. It is predominant along the Labrador coast at 200–700 m, while in north and central Labrador and the regions of Baffin Island it lives on the shelf and continental slope at 100 m and deeper. Near Baffin Island it is found at 230–400 m depths in the first two years of life. It has been caught in Hudson Strait at 260–360 m and in Davis Strait and southern Baffin Bay at 293–1,443 m and 0°C–5.6°C. Specimens identified as S. marinus (presumably S. mentella) have been caught at 140–530 m in Hudson Strait in 94 collections, and at 225–649 m in Ungava Bay in 26 collections. Off Labrador it is found at 2.5°C–4.0°C in depths of 274– 457 m in commercial quantities. Generally this species is found at 2°C–8°C, and therefore off Baffin Island it is found at greater depths because of an overlying cold current. The largest catches off Baffin Island were caught at 1.8°C–3.8°C. This species rises higher in the water column at night to feed than do the other two redfish species of Atlantic Canada. It is abundant in Davis Strait and eastern Hudson Strait but not in western Hudson Strait and Ungava Bay. A non-maturing group of redfish is found in Davis Strait and off the Labrador coast. They lack sexual dimorphism, and the reproductive organs are undeveloped. This is called a pseudo-population, formed from eggs and larval drifts from the Irminger Sea off southeast Greenland. Larvae are found in surface waters but may make vertical migrations in some parts of the range. Juveniles are pelagic for about four to five months.

biology: The biology of redfishes has not yet been fully worked

out because of the difficulties in separating the species. Often the biology of the “beaked” redfishes, S. fasciatus and S. mentella, is combined in the literature. Their food is generally pelagic items such as crustaceans (euphausiids, copepods, amphipods, hyperiids, and shrimps), siphonophores, ctenophores, molluscs, polychaetes, cephalopods, arrow worms, and, when they are larger, fishes. Fishes include Myctophidae (Glacier Lanternfish), Osmeridae, Pleuronectidae, Ammodytidae, Stomiidae, and Paralepididae as well as its own species. Diurnal vertical migrations occur in pursuit of prey. This species is a basic prey item for Greenland Halibut in Davis Strait, for Black Dogfish and, as small specimens, for Atlantic Cod in Ungava Bay. Thick-billed Murre chicks are fed this fish at Akpatok Island in Ungava Bay (0.04% of diet), and it is also eaten there by Brünnich’s Murres. Narwhals eat redfish (identified as Sebastes marinus but presumably S. mentella) in Mittimatalik (Pond Inlet), Baffin Island. Hooded Seals also eat redfish in the Canadian Arctic, again presumably this species. This is a slow-growing and long-lived species that may attain about 75 years. They are late maturers at 10–15 years. Half the population is mature at 18.5 cm for males and 29.5 cm for females. After age 10 years, females grow faster than males do. Strong year classes occur only every 5–12 years. Growth is slower in northern parts of the range, and the size at maturity of males in Davis Strait and Baffin Bay is greater than that of males in more southern waters such as the Gulf of St Lawrence.

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Sebastes mentella

Copulation occurs probably from October to January, but fertilization only takes place in February to April. Spawning occurs in March–August, and this species is the first of the redfishes to do so in the deeper water it inhabits in Newfoundland waters (other reports suggest that the Golden Redfish spawns earliest). The west Greenland shelf area is an important nursery for this species. About 15,000– 20,000 larvae of 6–9 mm length are released by each female each year. Larvae live in surface waters, perhaps for many years in northern areas, but descend to deeper waters as they grow. In southern areas the descent occurs by early fall in their first year of life.

by-catch in north Davis Strait had been 1,950.9 t. The excluder is not effective for small fish (10–20 cm). The estimated biomass and abundance in NAFO Division 0A was 1,226 t and 10.29 million fish, and in Division 0B was 3,448 t and 4.2 million fish, based on 2001 and 2000 trawl surveys. The Committee on the Status of Endangered Wildlife in Canada assessed the northern population as “Threatened” in April 2010. Directed fisheries have been closed in parts but are open in other areas. The by-catch in shrimp fisheries could affect population recovery even with separator grates. It is sold fresh or frozen and eaten fried, broiled, or baked.

importance: It is economically important and makes up the bulk

distribution: The species is found near Mittimatalik (Pond Inlet), in Baffin Bay, Davis Strait, Hudson Strait, sporadically in Ungava Bay, northwest, southwest, and southeast Greenland, and south to Nova Scotia. It is also found in the eastern Atlantic Ocean. This is the most wide-spread North American Sebastes, and most records in Arctic waters are presumed to be this species. Redfishes are mapped without distinction of species because these are often not clearly identified in the literature. The map is mostly of fish identified as S. mentella, including the northern and western limits.

of the catch in Newfoundland and Labrador waters. The commercial fishery is a recent development with a maximum catch of 389,000 t in 1959. Catches since then have been less as quotas were imposed to prevent overfishing. In 1979 the catch was worth $15.5 million and weighed 81,587 t. All three redfish species are managed as a single unit and often appear on the market, fresh or frozen, as “ocean perch.” The catch extends into Davis Strait and is taken primarily by southern Canadian fishers and foreign vessels, although Baffin Island and northern Québec Inuit are increasing their participation. Redfish are taken as a by-catch in the shrimp fisheries in north Davis Strait and Ungava and Hudson Bays. In 1998 as much as 21.63% of the by-catch was redfish in north Davis Strait, but with an excluding attachment, the Nordmore grate, it was less (8.47%). In 1992 the



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sources: Dunbar & Hildebrand (1952); Tuck & Squires (1955);

Tuck (1960); Mansfield (1967); Templeman (1967b); Zakharov & Chekhova (1972); Litvinenko (1974, 1980); Berth, Schultz, & Vaske (1979); Dunbar & Moore (1980); Ni (1981a, 1981b, 1982a, 1982b, 1983);

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Finley & Gibb (1982b); Kenchington (1983, 1986); LGL Ecological Research Associates (1983); McGlade, Annand, & Kenchington (1983); Misra & Ni (1983); Ni & McKone (1983); Ni & Sandeman (1984); Gaston (1985); Ni & Templeman (1985); Power & Ni (1985); Chumakov & Podrazhanskaya (1986); Konchina (1986); Morin & Dodson (1986); Atkinson (1987); Strong (1988); Trottier, Rubec, & Ricard (1988); Hudon (1990a); Rubec, McGlade, Trottier, & Ferron (1991); Troyanovsky (1992); Clarke (1993); Kulka (1995, 1999); Gorelova & Borodulin (1997); Orr & Bowering (1997); Alekseev (1999); Roques, Sévigny, Bernatchez, & Power (2000); Treble et al. (2000); Born & Böcher (2001); Chambers & Dick (2007).

Sebastes norvegicus (Ascanius, 1772)

Golden Redfish, sébaste orangé

common names: A local name is Suluppaagaq Angisooq

(Greenlandic). Other common names are Bream, Brim, Brin, Norway Haddock, Pinkbelly Rosefish, grande sébaste, sébaste atlantique, and sébaste doré.

taxonomy: The species was described originally from Norway (norvegicus). The species name norvegicus is a replacement name for Sebastes marinus (Linnaeus, 1758), which was based on a member of the Serranidae (Sea Basses). description: This species is distinguished from other northwest

Atlantic and eastern Arctic redfish species by a combination of overlapping external characters. It is easily misidentified when its range overlaps with the two closely related species from eastern Canada. A key internal character is the muscles attaching to the gas bladder. In this species there are three or four muscle heads with tendons passing between the second and third ribs or both the second and third ribs, and the third and fourth ribs; the muscle is wide; and the posterior tendon usually has six branches. This is the most robust of the northwestern Atlantic and eastern Arctic redfishes. It has the smallest eye. The dorsal fin spines number 13–17, usually 15; the anal fin soft rays number 7–10, usually 8. The lower unbranched pectoral fin-rays are 9 or more (rarely 8). The total vertebrae number 30–32, usually 30–31. The downward angle of the third preopercular spine is usually more than 40°. The knob at the tip of the chin is blunt and weak. Larvae have 9–24 ventral row melanophores. The overall colour is orange, orange yellow, or golden yellow. This colour is developed at about 15.0 cm length and an age of four to five years. The species attains 1.0 m in total length and 15.0 kg in weight, but usually 35–55 cm and 6.4 kg. The world all-tackle angling

Sebastes norvegicus

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record weighed 2.65 kg and was caught in 1982 at Holsteinsborg, Greenland.

habitat: It is found in water less deep on average than the habitat of the preceding species but with major overlaps. Generally it is found over rock or clay and silt bottoms from less than 300 m to over 750 m at 3°C–7°C. It is caught less frequently than the other redfish species even in southern waters, making assessment of biology difficult.

Family Cottidae Sculpins, Chabots

Brian W. Coad

biology: The biology of redfishes has not yet been fully worked

out because of the difficulties in separating the species, especially young fish. The presence and distribution of this species in Arctic waters is uncertain, and its biology there is poorly known. It forms only a minor portion of the total redfish catches in northern Canadian waters generally. As with the above species, its food is crustaceans and, when the redfish is larger, fishes; it rises in the water column at night to feed on pelagic prey. This too is a slow-growing and long-lived species that may attain greater than 75 years. The size at maturity for males is similar to that of the other redfish species, but females are larger. Other aspects of biology are similar, with some divergence in maturation patterns in southern waters. It may be the earliest spawner in Newfoundland waters of the three redfish species.

importance: See the previous species for a summary of the fisheries for redfishes. distribution: Its presence in Arctic waters as defined here may be confused with that of the previous two species. Old records of Sebastes marinus became S. norvegicus with the change in nomenclature, but these old records may have been mostly S. mentella. It is found on both sides of the North Atlantic Ocean, including Atlantic Canada, and may extend into Arctic waters, but this requires confirmation. It is recorded, sparsely, from northern Labrador. Golden Redfish is said to be much less common off North America than the other two species. It is common in southwest Greenland according to literature reports. Redfishes are mapped without distinction of species because these are often not clearly identified in the literature (although the map indicates mostly S. mentella, and fish identified as S. norvegicus are found in Davis Strait, Hudson Strait, and Ungava Bay). sources: See the sources under Sebastes mentella.



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Sculpins and Bullheads are found principally in marine and fresh waters of the northern hemisphere. There are about 300 species, including 61 in Canada, 14 of which occur in Arctic waters. The maximum size is 99 cm although most are smaller. Sculpins have a naked body or have scales, plates, prickles, or spines. The head is large and blunt, and the body tapers posteriorly. The head often has spines or knobs, particularly 1–4 preopercle spines. The eyes are large and dorsally placed. The mouth is large, but the teeth are generally small. There is one lateral line with 1–2 pores on the caudal fin base. The dorsal fin usually has separate short spinous and longer soft portions. Pelvic fins (rarely none) have 1 spine (often embedded and hard to detect) and 2–5 soft rays. There are no anal fin spines. The pectoral fin is large and fan shaped. The caudal fin is usually rounded or truncate. Adults lack a gas bladder. The males of some species have a penis-like urogenital papilla to deliver sperm internally to females. The eggs are few, large, and demersal and may be guarded by the male. The coloration is usually mottled and drab but can be colourful. Most Sculpins are small shallow-water, bottom-living species abundant in tide pools and shallows, although some are found down to 2,000 m. The larvae are planktonic. Adult food is bottom invertebrates such as molluscs, crustaceans, polychaete worms, insects, and small fishes. Sculpins in the Grande Baleine estuary of James Bay produce fewer and larger larvae after ice break-up that feed on larger food items, in contrast to the strategy of the small, numerous Sand Lance larvae that hatch before ice break-up and feed on small prey. They are not economically important although some are used as bait, occasionally as a change of diet, or in times of scarcity for dog and cat food, or they can provide young anglers with sport. About 10,000 kg of Sculpins were harvested in northern Québec in 1987–8. They are taken in the by-catch in northern Québec but most often by hook and line. Across the Arctic they are taken in native harvests, often incidentally in nets set for more important species like Arctic Char, and often by children. Some may be discarded unused. In the Belcher Islands they have been broiled and eaten with berries. Sanikiluaq residents in

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the Belcher Islands harvested about 1,176 Sculpins in 1984, eating them roasted on fires, boiled, or raw. Shorthorn Sculpins have been used for food in Ungava Bay in the past when other sources of food were absent, and for dog food. Dogs themselves are reported to catch this fish in tide pools. A marine test fishery at Wemindji in James Bay from 1987 to 1989 caught large numbers of Sculpins, meant for local fox and mink fur farms. Catches of Sculpins in the Baffin region ranged from none to 1,022 fish at Kangiqtugaapik (Clyde River), and at seven of twelve localities catches were less than 100 fish. In north Baffin Island many hunters have never eaten Sculpins and have only fished as children for them. Two hunters said they have “the devil’s likeness” and were therefore inedible. In Tuktoyaktuk some Sculpins are harvested in summer, fall, and winter. The Copper Eskimos in the central Arctic, in both historical and prehistorical times, fashioned double-pointed spears for stabbing Sculpins. Triglops species are a food item for Black Guillemot and Thick-billed Murre chicks at Prince Leopold Island off Somerset Island and are taken in shallow water. Sculpins are eaten by Bearded Seals at Aujuittuq (Grise Fiord) and at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River on Baffin Island). Belugas eat them off southeastern Baffin Island. “Kanayuk” and variant spellings is an Indigenous name referring usually to Myoxocephalus species.

sources: Jensen (1944); McAllister (1963a); McTaggart Cowan (1968, 1971, 1972a); Khan (1972); Neyelov (1979).

Artediellus atlanticus

Jordan and Evermann, 1898 Atlantic Hookear Sculpin, hameçon atlantique

common names: A local name is Atlantikup Kanajua (Greenlandic). Other common names are Arctic Sculpin, Hookhorn Sculpin, and chaboisseau atlantique. taxonomy: The genus is a diminutive form of Artedius, named for

the Swedish naturalist Peter Artedi (1705–35), called the Father of Ichthyology because Linnaeus used his notes for fish descriptions. The species name is the Latin atlanticus (of the Atlantic).

description: This fish and its relatives, the Rough and Snowflake

Hookears, are distinguished from other Sculpins in the Arctic by having the upper preopercular spine pointed and strongly hooked upwards; the margin of the first dorsal fin is often dark; and both dorsal fins in males are spotted or barred. This species has 22–28 caudal fin-rays, two small pores between the eyes anteriorly, no parietal cirri, 6–24 teeth on the vomer bone, and 4–39 teeth on the palatine bone, both bones being in the roof of the mouth. The first dorsal fin spines number 7–9, the second dorsal finrays 12–16, the anal fin-rays 9–13, and the pectoral fin-rays 18–24.

Artediellus atlanticus

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Artediellus atlanticus

The lateral-line pores number 17–28. The overall colour is a light green brown with dark brown to black or reddish blotches. The caudal peduncle has a dark band. The dorsal fins are dark with whitish streaks and spots or broad dark bars separated by thin light ones. The caudal fin has irregular dark bars that may be thin or broad. The iris may be reddish. Males have a higher first dorsal fin than that of females, and the white spots, streaks, or bars on it are prominent. Mature males have darker fin bars than those of females. The species reaches 16.8 cm in standard length, perhaps 25 cm, but is usually much smaller at around 5.3 cm in standard length on average in the Gulf of Maine, for example.

exceeds 17 years. Spawning takes place from May to November, perhaps as early as March to April in the Gulf of St Lawrence, and as late as the end of November in northeast Greenland, with each female having up to 216 eggs of 4.2 mm in diameter. The egg numbers may be as low as 40 because of their large size and the small body size of this species, but can be up to 350. The eggs develop for more than 200 days at 0°C, and the larvae hatch at an advanced stage and are similar to adults in appearance and behaviour. Males possibly guard the eggs.

habitat: This species is usually found on soft bottoms from shallows to deep water. In Davis Strait and Baffin Bay it has been caught at 151–1,365.5 m, elsewhere as shallow as 11 m, and as deep as 1,479 m in Davis Strait. The temperatures preferred range from 0°C to 4°C but as low as −1.7°C and up to 5.6ºC are tolerated. It is closely associated with Liparis fabricii and Boreogadus saida in high-latitude, low-temperature areas of Davis Strait and Baffin Bay.

distribution: This species is found in Baffin Bay, Davis Strait, Hudson Strait, and northern Ungava Bay and south to Cape Cod. It is found also on all Greenland coasts (as far north as 77º28' N on the Greenland side of Baffin Bay) and across the northern coast of Eurasia as far east as the Chukchi Sea.

importance: It is not economically important.

biology: Its food is bottom invertebrates such as polychaete

worms, and molluscs; and, in some populations, rarely crustaceans; in others, commonly amphipods. They are eaten by various other fishes including Atlantic Cod. The males are larger than the females. Mature females are 6.0 cm in total length. The maximum age



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description: This fish and its relatives, the Atlantic and Snow-

flake Hookears, are distinguished from other Sculpins in the Arctic by having the upper preopercular spine pointed and strongly hooked upwards; the margin of the first dorsal fin is often dark; and both dorsal fins in males are spotted or barred. This species has cirri and conical tubercles on the nape and the anterior back. Absent or weak nasal spines, used in some keys, do not appear to be consistent characters. The first dorsal fin spines number 7–9, the second dorsal fin-rays 12–14, the anal fin-rays 10–13, and the pectoral fin-rays 18–23. The lateral-line pores number 21–31. The head is greyish brown to reddish brown, and the flanks are lighter with large irregular dark or reddish spots that appear as bands in some fish. The male spiny dorsal fin has a black spot at the rear. All fins have yellow to orange bars, which may be wavy, irregular, or blotchy. The pectoral fin tips are white, and the fin has bars of yellow spots. The caudal fin base may have a dark bar. The species reaches 8.9 cm in total length.

habitat: This sculpin is found on mud-sand or gravel bottoms

Distribution of Artediellus atlanticus

sources: Jensen (1952b); Karrer (1973); Van Guelpen (1986); Hudon (1990a); Nielsen et al. (1992); von Dorrien (1993, 1996); Treble et al. (2000); Jørgensen et al. (2005); Chambers & Dick (2007); Mecklenburg, Byrkjedal, Karamushko, & Møller (2014).

often in brackish waters (10‰–15‰, or as low as 5.39‰ in the Kara Sea) at 0–100 m, rarely to 290 m, and usually less than 50 m in the Kara Sea. Its preferred temperatures are around 0°C, up to 6°C in Canada, and as low as −1.48°C. It may burrow into bottom sediments. The young are pelagic. There are an estimated 1,406,000 Hamecons in Liverpool Bay, making this species one of the most abundant in the Beaufort Sea.

biology: Its food is various bottom invertebrates such as crusta-

Artediellus scaber

ceans, including gammarid and hyperiid amphipods, copepods, mysids, euphausiids, cumaceans and isopods, and polychaete worms. Females about three to four years old are mature. The lifespan is about seven years. Spawning probably occurs in August– September with up to 336 eggs of 2.7 mm in diameter being produced.

Hamecon, hameçon rude

importance: It is not economically important.

Knipowitsch, 1907

Artediellus scaber

common names: Another common name is Rough Hookear.

distribution: The species is found at eastern Somerset Island, Dease Strait, Melville Sound, Amundsen Gulf, Beaufort Sea, Prince of Wales Strait between Banks and Victoria Islands, and the south coast of Melville Island. It is found also west to the Barents Sea across northern Eurasia and through the Alaskan Beaufort Sea, Chukchi Sea, and south to the Bering Sea. A fish from Killiniq, Ungava Bay (ROM 23385), was identified as this species by several competent workers, but it is the only eastern Canadian record. The relevant Royal Ontario Museum accession-book listing mentions “Canadian Arctic Expedition,” which worked in the western Arctic, and the labelling in the jar is also suspect (Erling Holm, personal communication, 5 April 2013), so this record is not accepted. It has not been recorded from Greenland or Atlantic Canada.

taxonomy: The species name is the Latin scaber (rough).

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Artediellus scaber

sources: Andriyashev (1961); Hunter, Jones, & Rich (1980); Frost & Lowry (1983); Percy et al. (1985); Ponomarenko (1995b).

Distribution of Artediellus scaber



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Artediellus uncinatus (Reinhardt, 1835)

Arctic Hookear Sculpin, hameçon neigeux

Artediellus uncinatus

common names: A local name is Imaviup Kanajua (Greenlandic). Other common names are Hookhorn Sculpin, Snowflake Hookear, Snowflake Sculpin, crapaud de mer, and crochet arctique.

taxonomy: The species name is the Latin uncinatus (hooked). description: This sculpin and its relatives, the Atlantic and

Rough Hookears, are distinguished from other Sculpins in the Arctic by having the upper preopercular spine pointed and strongly hooked upwards; the margin of the first dorsal fin is often dark; and both dorsal fins in males are spotted or barred. This species has 19–23 total caudal fin-rays, no pores between the eyes anteriorly, cirri on the parietal region (sometimes rubbed off), 0–12 teeth on the vomer bone, and 0–15 teeth on the palatine bone, both bones being in the roof of the mouth. The first dorsal fin spines number 7–8, the second dorsal finrays 10–14, the anal fin-rays 10–12, and the pectoral fin-rays 19–24. The lateral-line pores number 20–30. The body is blotched or mottled in dark or pale brown, or red. The belly is creamy in colour. The dorsal fins are dark brown with white to bluish spots. White spots in rows may be found on the head, body, pectoral, and caudal fins, hence the name “snowflake.” The caudal fin has a basal blotch. Males have elongated first dorsal fin spines, free of the membrane at their tip, and the first dorsal fin is larger than that of females, with

Artediellus uncinatus

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prominent black-edged white spots. The species reaches 10.0 cm in total length.

Gymnocanthus tricuspis

habitat: This sculpin is found on various bottom types from mud

Arctic Staghorn Sculpin, tricorne arctique

to gravel at 13–598 m. In Hudson Strait it has been caught at 115–366 m for 6 collections, in Ungava Bay at 158–458 m for 31 collections, in southern Davis Strait at 146–598 m for 40 collections, and in northern Davis Strait and Baffin Bay at 175–576 m for 7 collections. On the Labrador coast it is found at temperatures as low as −1.2°C.

(Reinhardt, 1830)

biology: It is assumed that its food is various invertebrates and

that spawning takes place in late summer. The eggs are deep orange and 3 mm in diameter. They may be spawned in clumps attached to hydroids. The egg numbers exceed 100. Mature females are found at only 4.0 cm total length. The lifespan is about four years. Gymnocanthus tricuspis

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, the

mouths of Cumberland Sound and Frobisher Bay, and in Hudson Strait and Ungava Bay. Literature records (Johansen, 1926; Holeton, 1974) and cruise data from Dolphin and Union Strait, the Beaufort Sea, and Resolute Bay on southern Cornwallis Island are unconfirmed by specimens or are misidentifications and are not mapped (Mecklenburg et al., 2002). There are no Alaskan Beaufort Sea records. It is found also from the Greenland coasts (and north to 76º32' N in Baffin Bay), south to northern Nova Scotia off Cape Breton, and possibly to Cape Cod, Massachusetts.

common names: A local name is Glatulk (Danish/Greenlandic). Another common name is crapaud de mer.

taxonomy: The genus comes from the Greek gymnos (naked) and akantha (spine or thorn). The species name comes from the Latin tres or tri (three) and cuspis (cusp). The genus is sometimes misspelled Gymnacanthus. Gymnocanthus tricuspis hudsonius Vladykov, 1933, is described from Nottingham Island and Killiniq (Port Burwell), Hudson Strait. This could be a distinct species, Gymnocanthus hudsonius, if diagnosable, but this taxon is not generally recognized and has not been re-examined recently. Acanthocottus patris Storer, 1850, described from Great Mecatina Island Harbor and Red Bay, Straits of Belle Isle, Labrador, and Gymnacanthus tricuspis groenlandicus Schmidt, 1927, described from the Arctic, are also synonyms. description: This species is characterized by the upper preopercu-

lar spine being broad at the tip or with two to three spinules, and the vomer and palatine bones in the roof of the mouth lacking teeth. The first dorsal fin spines number 10–12, the second dorsal finrays 12–17, the anal fin-rays 15–19, and the pectoral fin-rays 16–21. There are no scales except under the pectoral fins. There are 39–45 lateral-line pores. The top of the head has prickles or warts. Males have a large genital papilla and larger first dorsal and pelvic fins. The back is dark brown to grey in colour and is clearly separated from the yellowish to creamy or white belly. The flanks have blackish-brown bands formed from dark blotches. The dorsal and pectoral fins have brown or black bands, and the pectoral fin has a yellow tip or the fin is generally yellow. The anal and pelvic fins have yellow rays or are hyaline. The caudal fin-rays are brown with colourless membranes. The coloration is brightest in adult males that have roundish white spots on the darkish first dorsal fin, on the belly, on the inside of the pectoral fins, and on both sides of the pelvic fins. These spots are yellowish on the posterior part of the body. The species reaches 30.0 cm in total length.

Distribution of Artediellus uncinatus

sources: Jensen (1952b); Holeton (1974); Finley & Evans (1983); Van Guelpen (1986).



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habitat: This species is common in Arctic and Labrador waters and rare to the south. It was, with Shorthorn Sculpins, the

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Gymnocanthus tricuspis

commonest fish species in northwest Baffin Bay and eastern Lancaster Sound. There are an estimated 14,492,000 Staghorn Sculpins in Liverpool Bay, for example, making this species one of the most abundant in the Beaufort Sea. In the Chukchi Sea they number from 13 to 8,050 fish/sq km. They have been caught as shallow as 0.5 m in Brentford Bay, Boothia Peninsula, and at 7.0 m in Ikpiarjuk (Arctic Bay), Adam Sound, but are usually below 18 m down to 556 m on rock, sand or sand-mud, clay or gravel, and shell bottoms. At Cape Hatt, smaller specimens were most prominent in dense filamentous algae, mid-sized fish preferred Fucus distichus areas, and the largest fish showed no preference. The preferred temperatures are −1.84°C to 12.83°C, usually at or below 0°C. Studies at Qeqertarsuaq in Greenland show that this species has a lower metabolic scope and reduced competitive ability than Myoxocephalus scorpius and M. scorpioides at 9°C, indicating that climate change may alter species composition in Arctic waters. In Ungava Bay they are found deeper than the shore zone occupied by Shorthorn Sculpins. Salinity may be as low as 16‰ and as high as 35‰. They may burrow into sand and other soft substrates and enter brackish water. On the Beaufort Sea shelf they typify the < 50 m fish assemblage, presumably because they have a wider tolerance of temperature and salinity than does the Arctic Alligatorfish, which typifies > 50 m depths. When approached by a diver, young specimens in Resolute Bay, Cornwallis Island, swam away for up to 6.4 m. The fry are pelagic.

biology: As many as 86 food species have been recorded from

the stomachs of this species in the Beaufort Sea and the waters of southeast Baffin Island. Amphipods dominated. Other foods are small fishes like Sand Lances, cumaceans, clam siphons, gastropods, larvaceans, and polychaete worms. Some diet items are planktonic

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and must have been captured off the sea bed. At Cape Hatt, northern Baffin Island, the pelagic pteropod Limacina helicina was the major part of the prey at 49% by number and 56% by biomass. There is seasonal variation in diet, with the fish feeding on amphipods in December and on mysids and polychaete worms in June in Resolute Bay, Cornwallis Island. Diets also vary with locality, with gammarids dominating in the Chukchi Sea and polychaetes in the Beaufort Sea. Larval and post-larval Staghorn Sculpins feed on copepods such as Acartia clausi, Acartia longiremis, Cyclops sp., Eurytemora herdmani, Calanus glacialis, Calanus hyperboreus, Jaschnovia tolli, Limnocalanus macrurus, Metridia longa, Microcalanus pygmaeus, Microsetella norvegica, Microsetella rosea, Oithona similis, Oncaea borealis, Pseudocalanus minutus, Tisbe furcate, and ectinosomatids; on amphipods such as Apherusa glacialis, Onesimus glacialis, Parathemisto abyssorum, and Paroediceros propinquus; on diatoms such as Coscinodiscus sp.; on mysids such as Mysis litoralis; on sea butterflies (pteropod molluscs) such as Clione limacina and Limacina helicina; on rotifers such as Keratella cochlearis; on narcomedusans such as Aeginopsis laurentii; on larvaceans such as Fritillaria borealis and Oikopleura vanhoeffeni; and on polychaetes, bivalves, cirripedes, podocop ostracods, eggs, and even insects in the Beaufort Sea. This sculpin is eaten by Harp Seals; by Black Guillemots at Digges Sound, Hudson Bay; by Atlantic Cod at Killiniq (Port Burwell), Ungava; by Brünnich’s Murres and Thick-billed Murre chicks (0.04% of diet) at Akpatok Island, Ungava Bay; and by Thick-billed Murre chicks at Coats Island and Cambridge Point, Coburg Island. In Barrow Strait, Nunavut, it is eaten by Thick-billed Murres and Black Guillemots. It is probably food for Arctic Char in Hudson Bay and is eaten by char at Bernard Harbour. Ringed Seals, Bearded Seals, Arctic Cod, Polar Eelpout, and Bering Flounder also eat this species in Arctic Alaska.

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Sexual maturity is attained towards the age of 4 years. Males mature at 2–3 years (60–70 mm), and females at 3–4 years (about 90 mm) in the Chukchi Sea. Males mature at about 3–4 years (8 cm), and females at 3–4 years (13 cm) at Cape Hatt, northern Baffin Island. The lifespan at eastern Baffin Island and at Cape Hatt is 8 years, and in the Chukchi Sea is 9 years. A study in the Chukchi Sea indicated that their life in Arctic waters is unpredictable as the age structure changes from year to year. The reasons can be recruitment failures, mass mortalities, or dispersal of individuals. Spawning occurs in October at Newfoundland, but fish in spawning colours are seen at Killiniq (Port Burwell), Ungava, in mid-September. The yellow-pink or white eggs are up to 2.0 mm in diameter and number 35,000 in large females, with up to 3,723 mature eggs. The eggs are laid in algae, and the male guards the eggs. Larvae appear in the plankton from the end of April to the end of June. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae occur from May to July, suggesting a winter spawning. This species needs 80–90 days for hatching of eggs at 1.5°C.

importance: In northern Québec this species is considered a desirable food fish and is caught by hook and line.

distribution: It is found in the nearshore and shore waters of Nares Strait, Baffin Bay, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, Foxe Basin, the Arctic islands, Dease Strait, Melville Sound, Bathurst Inlet, Dolphin and Union Strait, Amundsen Gulf, and the Beaufort Sea. It is found also on all Greenland coasts and south to Maine in the eastern Atlantic Ocean; in the northern Bering and the Chukchi Seas; and on the Arctic coast of Eurasia.

shev (1948); Dunbar & Hildebrand (1952); Jensen (1952b); Tuck & Squires (1955); Ellis (1960); Tuck (1960); MacInnis (1973); Wilson (1973); Thomson, Cross, Bain, & Patterson (1978); Bradstreet (1980); Hunter et al. (1980); Green (1983); Gaston et al. (1985); Percy et al. (1985); Gaston et al. (1987); Crawford (1989); Gaston (1989a, 1991); Atkinson & Percy (1992); Pavlov, Burykin, & Kublik (1992); Ponton et al. (1993); Ponomarenko (1995b); R.L. Smith, Barber, Vallarino, Gillispie, & Ritchie (1997); Gray & Norcross (2013); Majewski, Lynn, Lowdon, Williams, & Reist (2013); Seth et al. (2013).

Icelus bicornis (Reinhardt, 1840)

Twohorn Sculpin, icèle à deux cornes

common names: A local name is Kanajoq Kapinartulik Nalinginnaq (Greenlandic). taxonomy: The genus comes from the Greek ikelos (Icelus, son of Hypnos, the god of sleep), because of the sluggish movements. The species name is the Latin bicornis (two horned). description: This species and its relative, the Spatulate Scul-

pin, are distinguished from other Sculpins in the Arctic by having a forked or trifurcate uppermost preopercular spine that is not strongly hooked upwards and by having a row of spiny plates below the dorsal fins. There are four preopercular spines, the lower three being directed downward. The species is separated from its relative by having a pair of spines at the base of the caudal fin. The lateral line is incomplete (rarely to the posterior edge of the hypural plate, the end of the vertebral column); caudal peduncle scales are usually present above and below the midline of the caudal peduncle; axillary pectoral scales number 9–30; a row of small scales is often present between the dorsal scale row and the lateral line; a row of scales is often present above the anal fin; and the male urogenital papilla is cylindrical, its tip elongate and tapering and longer than caudal peduncle depth. The first dorsal fin spines number 7–10, the second dorsal fin-rays 17–23, the anal fin-rays 12–17, and the pectoral fin-rays 15–19. Lateral-line scales number 23–43. The overall colour is yellowish brown with darker spots and blotches dorsally, and white on the belly. A dark bar passes through the eye but may be diffuse in some fish. There is a dark spot at the pectoral fin base. The caudal, second dorsal, and pectoral fins are thinly barred. The species reaches 15.7 cm in standard length.

habitat: Twohorn Sculpins are abundant and wide spread in marine waters offshore. They are found at −1.8°C to 8.8°C from the shallowest water to 930 m on mud, sand, shale, pebbles, or rock

Distribution of Gymnocanthus tricuspis



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sources: Johansen (1926, 1927a); Vladykov (1933a); Andria-

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Icelus bicornis

bottoms or, in the Russian Arctic, in algae. In Davis Strait they have been caught at −1.35°C to 3.96°C. Baffin Bay fish have been caught at 41–930 m and Ungava Bay fish at 115–279 m. In the Canadian Arctic they are in shallower water than in warmer, southern, Atlantic waters, at 10 m in Eclipse Sound near Mittimatalik (Pond Inlet), for example. Salinity may be as low as 25.4‰. In the Alaskan Beaufort Sea and northeastern Chukchi Sea this species, along with Arctic Cod and Polar Eelpout, accounted for 65% of all trawled fishes caught. Twohorn Sculpins have anti-freeze peptides in their blood, which help them resist low temperatures.

biology: This species has 91 prey species recorded from the

stomach contents of fish from the Beaufort Sea and the waters of southeastern Baffin Island. They feed on amphipods predominantly, and on such other crustaceans as mysids, copepods, isopods, euphausiids, cumaceans, and shrimps, and less importantly on polychaete worms. There is seasonal variation in diet, with feeding on amphipods in December and on mysids and polychaete worms in June in Resolute Bay, Cornwallis Island. It is eaten by Brünnich’s Murres and Thick-billed Murre chicks (0.15% of diet) at Akpatok Island, Ungava Bay, and at Cambridge Point, Coburg Island. Arctic Char eat this species at Bernard Harbour, Nunavut, and in Dolphin and Union Strait.

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Females are larger than males, which reach a maximum size of about 7.0 cm. Females mature at four years of age at 6.0 cm. In northern waters spawning occurs in August to October, and females have 700–1,000 eggs with a diameter of 3.1 mm. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae, possibly of this species, occur from May to June.

importance: It is not economically important. distribution: The species is found in Nares Strait, Baffin Bay, Cumberland Sound, Frobisher Bay, Davis Strait, Hudson Strait, Ungava Bay, northern Hudson Bay, the Arctic islands including the northern tip of Ellesmere Island, Dease Strait, Bathurst Inlet, Amundsen Gulf, and the Beaufort Sea including the Alaskan Beaufort, to all Greenland coasts and south to the southern Scotian Shelf in Atlantic Canada. General reports for James Bay are not confirmed by specimens. It is possibly found in Richmond Gulf. The species occurs on the Arctic coast of Eurasia but not in the Chukchi or East Siberian Seas.

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description: This species and its relative, the Twohorn Scul-

Distribution of Icelus bicornis

sources: Günther (1877b); Jensen & Volsøe (1949); Tuck & Squires (1955); Ellis (1960); Tuck (1960); Frost & Lowry (1983); Green (1983); Craig (1984); Nelson (1984); Atkinson & Percy (1992); Ponton et al. (1993); Wöhrmann (1997).

Icelus spatula

Gilbert and Burke, 1912 Spatulate Sculpin, icèle spatulée

pin, are distinguished from other Sculpins in the Arctic by having a forked or trifurcate uppermost preopercular spine and a row of spiny plates below the dorsal fins. There are four preopercular spines, the lower three being directed downward. The species is separated from its relative by lacking the pair of spines at the caudal fin base; the lateral-line scales end on the posterior edge of the hypural plate; the caudal peduncle scales are absent or, if present, are only above the lateral line; axillary scales number 1–14; there are no scale rows between the dorsal scale row and the lateral line; there is no scale row above the anal fin; and the male urogenital papilla is spatulate, with a short, curved, or hook-like tip, its tip being shorter than the caudal peduncle depth. The first dorsal fin spines number 7–11, the second dorsal fin-rays 17–22, the anal fin-rays 13–18, and the pectoral fin-rays 16–20. The lateral-line scales number 33–47. The head is covered with prickles. The male has a curved, spatulate genital papilla. The overall colour is mottled light brown, yellow tan, or pinkish brown to olive, with four to five darker saddles over the back; the saddles are often obscured in Arctic fish by the darker back or are not present in some fish. The belly is whitish to creamy. The first dorsal fin has one to two large blotches or pigment along the rays. The second dorsal fin is barred irregularly. The caudal and pectoral fins have four to seven bars, and there is a white area on the anterior three pectoral fin-rays. The caudal base has a brown patch. There may be two large brown spots below the eyes. The species reaches 21.0 cm in length.

habitat: This sculpin is found from shallow water down to about

930 m over sand and mud or occasionally rock, stones, and pebbles. In Baffin Bay it has been recorded from 41 m to 930 m. On the Labrador coast its preferred temperatures are as low as −1.85°C, usually below 0°C, and elsewhere to 7.8°C. In Exeter Sound the bottom temperature was −1.58°C at 75–200 m. The salinity range is 24.7‰– 34.2‰. The species is common in Resolute Bay, Cornwallis Island, and juveniles have been observed swimming at 0.5–1.0 m above the bottom and actively swimming away from divers.

biology: The diet is dominated by mysids and amphipods in the

Icelus spatula

common names: A local name is Spatel-tornulk (Danish/

Greenlandic).

taxonomy: The species name is the Latin spatula (small spoon or sword, a blade).



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Beaufort Sea and the waters of southeastern Baffin Island. Other foods are crustaceans such as copepods, euphausiids, cumaceans and shrimps, polychaete worms, and molluscs. Some planktonic food, such as arrow worms and mysids in Resolute Bay, Cornwallis Island, is taken by swimming up to 1 m off the bottom. In Barrow Strait, Nunavut, this fish is eaten by Thick-billed Murres and Black Guillemots. At Coats Island it is fed to Thick-billed Murre chicks by the parent birds. It is probably eaten by Bearded Seals at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River) on Baffin Island. Atlantic Cod also feed on this species in Ungava Bay. Females grow larger than males. Males only reach 7.5 cm in length. The lifespan is five years at eastern Baffin Island, and up to seven years elsewhere. Spawning occurs in August–September in the Bering Sea. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae, possibly of this species, occur from May to June. Females carry up to 9,100 eggs with diameters up to 2.5 mm.

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Icelus spatula

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, the mouth of Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, sparsely in Hudson Bay, the Arctic islands, Dease Strait, Bathurst Inlet, Amundsen Gulf, and the Beaufort Sea. It is found also in the Alaskan Beaufort Sea, Chukchi Sea, Sea of Okhotsk, and Bering Sea, northwest and southwest Greenland, and southward to Browns Bank in Atlantic Canada. General reports for James Bay are not confirmed by specimens. It occurs also on the Arctic coast of Eurasia. sources: Jensen & Volsøe (1949); Edwards (1961); Bradstreet

(1980); Hunter et al. (1980); Finley & Evans (1983); Green (1983); Gillis & Allard (1984, 1986); Nelson (1984); Gaston et al. (1987); D.B. Stewart et al. (1991); Atkinson & Percy (1992); Ponton et al. (1993).

Distribution of Icelus spatula

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of the spiny dorsal fin, and two smaller saddles under the second dorsal fin. The fins have thin bars, and the caudal fin may have only the rays pigmented. Males have contrasting black and cream areas on the body and fins, especially the pelvic fins, during the spawning season. The species attains 19.4 cm in total length.

Myoxocephalus aenaeus (Mitchill, 1814)

Grubby, chaboisseau bronzé

common names: Other common names are Little Sculpin and crapaud de mer nain.

taxonomy: The genus comes from the Greek myoxos (dormouse) or possibly myos (muscle) and kephale (head), both rather obscurely. The species name comes from the Latin aeneus (bronze or brassy) and is sometimes misspelled aeneus. description: This species and its four Arctic relatives in the

genus Myoxocephalus are distinguished from other Sculpins by the upper preopercular spine being a simple straight point; the vomer bone in the roof of the mouth bears teeth, but palatine teeth are absent; there are no oblique folds on the lower flank; the lateral line lacks plates; and the anal fin-rays are 17 or less. It is separated from its relatives by having three preopercular spines (one pointing forward to downward, two pointing backward); the uppermost preopercular spine is less than twice as long as the one below; and the spiny dorsal fin origin is in front of the rear edge of the operculum. The first dorsal fin spines number 8–11, the second dorsal fin-rays 12–17, the anal fin-rays 8–14, and the pectoral fin-rays 14–17. There are a few scales along each side of the lateral line. The overall colour is grey, sometimes greenish grey to brown, with darker mottles or saddles on the flanks, and pale grey to white below. There is often a pale band on the lower caudal peduncle. The colour tends to match the bottom where the fish lie. There is a dark, wide saddle under the front

habitat: Grubbies are found abundantly on mud, sand, gravel, or

rock bottoms from tide pools usually down to about 130 m in coastal waters at a wide range of temperatures (0°C–21.1°C) and salinities. Five collections from Hudson Strait were at 0–357 m. They may enter estuaries and are found among eelgrass. They have anti-freeze peptides in their blood, which help them resist low temperatures, a necessity for a shallow-water, winter-spawning species. The antifreeze protein increases in November to a peak level in January.

biology: Its food includes crustaceans, molluscs, worms, sea

squirts, sea urchins, and a variety of small fishes such as Sand Lances and Sticklebacks. They are also scavengers. This species is eaten by Atlantic Cod and Longhorn Sculpins, among other fishes. Spawning takes place in winter to spring, and possibly summer. At Cape Cod some females spawn during their first year of life. The eggs are up to 2.0 mm in diameter and can be clear or coloured red, green, or yellow. The eggs are laid in clumps that stick to seaweed, shells, bryozoans, or rocks. Grubbies can make growling sounds by vibrating the pectoral muscles.

importance: It is not economically important. distribution: The species is found sparsely in Hudson Strait, Ungava Bay, and northern Hudson Bay and south to New Jersey in the Atlantic Ocean.

Myoxocephalus aenaeus



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Myoxocephalus octodecemspinosus (Mitchill, 1814)

Longhorn Sculpin, chaboisseau à dix-huit épines

Myoxocephalus octodecemspinosus

common names: Other common names are Gray Sculpin, Great Sculpin, Hacklehead, Long-spined Sculpin, Muddler, Pig-fish, Plug Eye, Rubbish, Scopy, Scully, Scummy, and Scumpy. Distribution of Myoxocephalus aenaeus

sources: Reisman, Fletcher, Kao, & Shears (1987); Hudon (1990a); Wöhrmann (1997).

taxonomy: The species name comes from the Latin octodecim (eighteen) and spinosus (spined). description: This species and its four Arctic relatives in the

genus Myoxocephalus are distinguished from other Sculpins by the upper preopercular spine being a simple straight point; the vomer

Myoxocephalus octodecemspinosus

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bone in the roof of the mouth bears teeth, but palatine teeth are absent; there are no oblique folds on the lower flank; and the lateral line lacks strong plates. The species is separated from its relatives by having three preopercular spines (one pointing forward to downward, two pointing backward), and the uppermost spine is about four times longer than the one below. The first dorsal fin spines number 7–10, the second dorsal finrays 15–17, the anal fin-rays 12–15, and the pectoral fin-rays 16–19. The lateral line has weak plates. The overall colour is dark olive grey, greenish brown, or greenish yellow, fading to a white belly. The colour will vary with the background on which the fish lives; it may be plain if that background is uniformly coloured as when it is sandy or muddy, and it may be even whitish with dark-grey blotches when the bottom comprises pebbles with white corallines. The flanks are blotched and may have three to four irregular bars. The first dorsal fin is dusky, the second dorsal fin has three to four bars, and there are dark bars on the caudal, anal, and pectoral fins. The fins have a pale yellowish background colour. The iris is red. The species reaches 45.7 cm in total length.

habitat: Longhorn Sculpins are shallow-water residents in spring

and summer and retreat to deeper water in winter. They are recorded down to 342 m in Ungava Bay. They can be found in harbours. The preferred depth on the Scotian Shelf is 53–90 m. The winter migration avoids very low temperatures, and this species lacks the blood anti-freeze found in some other Sculpins, such as the Grubby. It is found at a wide range of temperatures, −0.5ºC to 19ºC.

Distribution of Myoxocephalus octodecemspinosus

source: Hudon (1990a).

biology: Its food is crustaceans, molluscs, worms, squids, sea

squirts, and small fishes such as Herrings, Smelts, Sand Lances, and Cods. It also scavenges in harbour areas. It is eaten by Atlantic Cod and a variety of other fishes and is a cannibal. This sculpin may be seen sitting on the sea bottom with its mouth gaping. Longhorns make sounds by the effect of contraction of the lateral muscles on the pectoral girdle. Sexual maturity is attained in the third year of life, and the lifespan is nine years. Spawning takes place in winter from December to January, perhaps November to February. The eggs are adhesive and are laid in masses on sponges and in depressions on hard surfaces, such as shells and rock cavities. The eggs are up to 2.3 mm in diameter after swelling in water, number as many as 8,000, and can be green, orange, chocolate brown, or red.

Myoxocephalus quadricornis (Linnaeus, 1758)

Fourhorn Sculpin, chaboisseau à quatre cornes

importance: This sculpin has been used for fish-meal and oil and in pet food in southern Atlantic Canada.

distribution: It is found in the Hudson Strait, Ungava Bay, and possibly eastern Hudson Bay – the latter from Halkett (1920) and needing verification – and south to Virginia in the Atlantic Ocean. Myoxocephalus quadricornis

common names: Local names are Kanajuk, Kanayuk, Kanayuq,

Kaneeok, Kan-ny-yoke, and Kanyok (Inuktitut); and Hornulk (Danish/Greenlandic). Other common names are Devil Fish, Fourspined Sculpin, and chaboisseau quadricorne.



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Myoxocephalus quadricornis, with variation in head spine development (top left)

taxonomy: The species name comes from the Latin quatuor (four) and cornu (horn), hence four horned. Cottus hexacornis Richardson, 1823, described from the mouth of the Tree River near the Coppermine River, and the Arctic Sea about Herschel Island, is a synonym and is regarded as a valid subspecies of Myoxocephalus quadricornis for the western Arctic by some authors. Acanthocottus labradoricus Girard in Storer, 1850, described from the Labrador coast, is a synonym (and has been regarded as a synonym of M. scorpius) and is regarded as a valid subspecies of Myoxocephalus quadricornis for the eastern Arctic by some authors. The subspecies were distinguished on the basis of relative development of the frontal and parietal head spine (large and spongy in hexacornis, and weak to absent in labradoricus). Cottus polaris Sabine, 1824, described from North Georgia, the east side of the Boothia Peninsula, is a synonym but has been used by some authors as a subspecies name for American Arctic populations of Fourhorn Sculpin, with the type subspecies (Myoxocephalus quadricornis quadricornis) in Europe.

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This species may be placed in a distinct genus, Triglopsis Girard, 1851, but recent molecular studies indicate that it should be retained in Myoxocephalus.

description: This species and its four Arctic relatives in the

genus Myoxocephalus are distinguished from other Sculpins by the upper preopercular spine being a simple straight point; the vomer bone in the roof of the mouth bears teeth, but palatine teeth are absent; there are no oblique folds on the lower flank; and the lateral line lacks plates. The species is separated from its relatives by having four preopercular spines (two pointing forward and two backward), the lateral line is usually incomplete; mandibular pores are difficult to see; and, uniquely there are four blunt, rough-edged spines on top of the head, two between the eyes and two further back (the frontal and parietal spines). The first dorsal fin spines number 6–10, the second dorsal fin-rays 11–17, the anal fin-rays 12–17, and the pectoral fin-rays 14–18. There is a row of rough scales from below the first dorsal fin origin to the

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caudal fin, and a second row below the second dorsal fin and sometimes below the first dorsal fin. The colour is dark brown to dark green brown or grey on the back, the flanks are brassy to golden brown, and the belly is white. There are four to seven saddles, or the colour is more uniform. The soft dorsal, anal, caudal, and pectoral fins are barred, the pectoral fins having up to four bars. The second dorsal fin membranes are often darker than the rays. Males develop a rosy colour under the head, on the lower pectoral fin, and on the anal and pelvic fins, and a canary-yellow colour on the lower sides and belly. The second dorsal fin and pelvic fins are larger in mature males, and males have tubercles on the second dorsal and pectoral fins. The species reaches 39.6 cm, and possibly 60.0 cm, in total length.

habitat: This sculpin is commonly found in shallow coastal

areas on rocky and sand-mud substrates, usually near or in estuaries, and seldom below 20 m, but as deep as 45 m and perhaps to 100 m. Stream mouths and shallow protected areas are important for juveniles. The species is common in tide pools and is the commonest sculpin along the coast of southeastern Baffin Island and the Yukon coast. It was the only species found in tide pools at Cape Hatt, northern Baffin Island. In Mould Bay, Prince Patrick Island, it is extremely common and tangles gill-nets unless these are set about 0.3 m off the bottom. It has been reported lying on Laminaria algae fronds in Resolute Bay, Cornwallis Island. The spring freshet of the Mackenzie River is avoided, and this species moves offshore; however, it may enter rivers for some distance, including up to 200 km in the Mackenzie River. Some populations are land-locked relicts in lakes. Overwintering occurs in bays and inlets and possibly offshore. It is scarcer than other Myoxocephalus species in Ungava Bay, apparently preferring colder waters, but in the Beaufort Sea nearshore areas, their fry are often the only fish species to be found. This species, and the Arctic Flounder, are the most important demersal species in the Beaufort Sea. It is the second most abundant species, after Arctic Cisco, around Herschel Island. In the Beaufort Sea, adults are wide spread and abundant in brackish nearshore areas, at temperatures of −1.7°C to 13.5°C and salinities of 0‰–32‰. This species comprises 80% of the winter catch of marine fishes in the Beaufort Sea and 75% of the biomass of fish (6,000–9,500 kg/sq km) in trawl surveys near Tuktoyaktuk. In Hudson and James Bays it is a true estuarine species, spending its whole life in this habitat, and here too it may be found in tide pools. It dominates in the estuary at Wemindji, James Bay, during summer, along with Slender Eelblennies and juvenile Cisco and Lake Whitefish. In James Bay near the Eastmain River it is often the second most abundant species after Cisco. Larvae in Rupert Bay and nearby James Bay have been caught mostly at 13°C–15°C and 0‰–4‰ from June to August. In James Bay, adults have a plasma-freezing point of −0.85°C. In the Eastmain River estuary in James Bay it is found at 6.5°C–16.0°C and a salinity of 1‰–17‰ throughout the year. In eastern Hudson Bay this species is more common in shallower and more brackish water (7.9‰–28.0‰) than Shorthorn Sculpins (16‰–24‰). Fry are pelagic as well as benthic. Males are territorial and may not be caught as easily as females. It is diurnal from November to April and largely nocturnal the rest of the year.



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biology: Its food consists of crustaceans such as gammarids, isopods, mysids, ostracods, copepods, cumaceans, molluscs (such as the limpet Acmaea testudinalis and the gastropod Margarites helicinus in east Hudson Bay), various aquatic insects, polychaete worms, sipunculids, hydrozoans, ascidians, mammal remains, plant fragments, fish eggs, and small and juvenile fishes such as Sand Lances, Capelin, Smelts, Fourhorn Sculpin, Arctic Cisco, Arctic Cod, and Threespine and Ninespine Sticklebacks. In the Beaufort Sea, larger fish prefer the giant isopod Saduria entomon, and isopods can comprise 99.3% of the average diet. In Cambridge Bay, food is primarily bivalves, sea urchins, polychaetes (especially Pectinaria sp.), amphipods, and other invertebrates. Larval Fourhorn Sculpins in the Beaufort Sea eat such copepods as Calanus glacialis, Cyclopina sp., Eurytemora sp., Limnocalanus macrurus, Metridia longa, Oncaea borealis, and Pseudocalanus minutus; branchiopods such as Podon leuckarti; mysids such as Mysis litoralis; and eggs, bivalves, and polychaetes. At Cape Hatt, northern Baffin Island, the pelagic pteropod Limacina helicina was the major part of the prey biomass. The species is a major predator on eggs of other fishes and on its own eggs. It is an important forage fish. In Nunaluk Lagoon on the Yukon coast it is food for Mew Gulls, whitefishes, Burbot, Arctic Sculpin, Eelpouts, and Arctic Char. It is fed on by Bowhead Whales, Inconnu, and Arctic Char in the Canadian Beaufort Sea; by Ogac at Wemindji, James Bay, and on the coast of Shepherd Bay on the Boothia Peninsula; by Atlantic Cod at Killiniq (Port Burwell), Ungava; by Arctic Cod in Wakeham Bay, Ungava; and by Brook Trout in Richmond Gulf. At Bernard Harbour, Nunavut, it is eaten by Arctic Char, Saddled Eelpouts, and Shorthorn Sculpins. In Barrow Strait, Nunavut, it is eaten by Thick-billed Murres and Black Guillemots. Near the Nuvuk Islands, Nunavut, it is eaten by Black Guillemot chicks. Black-legged Kittiwakes also eat this fish. It is probably eaten by Bearded Seals at Aujuittuq (Grise Fiord) and probably also at Mittimatalik (Pond Inlet) on Baffin Island. A subadult male Polar Bear was seen to dive and capture this species in the Union River, Creswell Bay, Nunavut. Its lifespan is 16 years, with maturity at three to eight years off the Yukon coast. Off the Alaskan coast, maturity is attained as early as age two years, but by age four years in males and at age four to six years in females. Males mature before females on the Yukon coast. Females grow faster, live longer, and are larger than males. Spawning occurs from late summer and autumn to winter and as late as spring, and it may occur under ice as in the brackish Mallik Bay on the Mackenzie Delta. Timing varies with temperature, salinity, and locality. On the Canadian Beaufort Sea coast they spawn in mid-winter, and the eggs hatch before the break-up of the ice. Spawning may occur more than once in each season in the Alaskan Beaufort Sea. Late September or October appears to be the time at Shepherd Bay on the Boothia Peninsula. The presence of larvae in Rupert Bay and nearby James Bay indicates spawning in May and June. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae, possibly of this species, occur from May to June. The eggs are up to 3.0 mm in diameter, number up to 17,968, and can be green, brown green, or brown. A nest is excavated by the male in mud or algae, where the female lays adhesive clumps of eggs. The male guards the

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eggs. This species needs 80–90 days for hatching of eggs at 1.5°C. Larvae are pelagic and abundant in May to June in shallow, pelagic areas. High levels of cadmium have been found in livers of this species at Tuktoyaktuk, measuring 0.56 ng/g. In Cambridge Bay near a dump, Fourhorn Sculpins contain up to 1,950 ng/g PCBs in pooled liver system, or 220 ng/g in whole tissue excluding liver. These are the highest levels recorded for Arctic marine fishes and approach those seen in marine mammals. Populations of the Fourhorn Sculpin in fresh waters and euryhaline lakes of the Arctic Archipelago are assessed as “Data Deficient,” and those of Garrow Lake, Cornwallis Island, are in danger of extirpation or have already been lost because of lead-zinc mining waste.

importance: This sculpin is said to have been used as food along

sources: Johansen (1926, 1927a); Berg & Popov (1932); Vladykov

(1933a); Dunbar & Hildebrand (1952); MacDonald (1954); Ryder et al. (1964); Macpherson (1971); Khan & Faber (1973); Kendel et al. (1975); Percy (1975); Craig & McCart (1976); McAllister & Aniskowicz (1976); Simard & Legendre (1977); Talbot (1977b); Barber (1978a); Berkes (1979); Neyelov (1979); Bradstreet (1980); Craig & Haldorson (1980); Dutil & Power (1980); Lowry & Burns (1980); McCart (1980); Morin et al. (1980); Cairns (1982); Foy (1982); Lambert & Dodson (1982a); Finley & Evans (1983); D.B. Stewart & Bernier (1983); Craig (1984); Lawrence et al. (1984); Bond & Erickson (1987, 1989); Goldberg, Yasutake, & West (1987); Baker (1988); Crawford (1989); Houston (1990); Jensen & Jensen (1991); Morin et al. (1991, 1992); Ponton et al. (1993); Whoriskey, Audet, & Hudon (1994); Bright, Dushenko, Grundy, & Reimer (1995); Kontula & Väinölä (2003); Dyck & Romberg (2007); Burn (2012); Niemi et al. (2012).

the shores of Hudson Bay and is occasionally caught for sport. It was often used for dog food and sometimes as bait for foxes when caught in coastal gill-nets set for Arctic Char. In James Bay subsistence fisheries they are killed and discarded because they foul nets and are considered inedible.

Myoxocephalus scorpioides

distribution: The species is found in Nares Strait and at the tip

Arctic Sculpin, chaboisseau arctique

of Ellesmere Island, northern Baffin Island, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, Foxe Basin, Gulf of Boothia, the Arctic islands, Queen Maud Gulf, Dease Strait, Melville Sound, Bathurst Inlet, Dolphin and Union Strait, Amundsen Gulf, around Banks and Victoria Islands, and the Beaufort Sea. It is found also in the Alaskan Beaufort Sea, Chukchi Sea, and Bering Sea, northwest and northeast Greenland, and northern Eurasia.

(Fabricius, 1780)

common names: Local names are Kanajuk, Kanayuk, and Tivaqiq; and Falsk Ulk (Danish/Greenlandic). Another common name is Northern Sculpin. taxonomy: The species name comes from the Greek skorpios (scorpion), the root of “sculpin” in English, and eidos (likeness, form). description: This species and its four Arctic relatives in the

Distribution of Myoxocephalus quadricornis

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genus Myoxocephalus are distinguished from other Sculpins by the upper preopercular spine being a simple straight point; the vomer bone in the roof of the mouth bears teeth, but palatine teeth are absent; there are no oblique folds on the lower flank; and the lateral line lacks plates. The species is separated from its relatives by having three preopercular spines (one pointing forward to downward, two pointing backward); the uppermost preopercular spine is less than twice as long as the one below; the spiny dorsal fin origin is not obviously in front of the rear operculum edge; the pectoral fin-rays number usually 14–16 (range 14–18); the total count of dorsal, anal, and pectoral fin-rays is 35–38; the frontal and parietal spines are not well developed and lack accessory spines at the base; papillae on top of the head are large and closely spaced; and there are tabs or cirri over the eyes. The first dorsal fin spines number 8–10, the second dorsal finrays 13–17, and the anal fin-rays 10–14. The lateral-line pores number 36–42. The colour is a dark olive to blackish brown or purplish brown with darker mottles or bands. The second dorsal and anal fins are dusky with light spots. The caudal fin base has a pale blotch. The pectoral fin has broad dark bands in females, or four to five rows of clear

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Myoxocephalus scorpioides

spots in males. Other fins have bands or white blotches. Males have white or silvery spots with dark borders on the flank under the pectoral fin and above the anal fin. The breast and belly have dark spots in males, and there is a broad white stripe on the reddish-orange belly running from the pelvic fins to the anal fin. Males have spots on the lower lip. The species reaches 30.0 cm in standard length.

habitat: This sculpin is found in shallow coastal waters, most

commonly between tides on rocky bottoms among algae and under stones. It may descend to 280 m but is usually in or just below the tide zone. It survives under ice cover at −1.4°C because its blood contains an anti-freeze protein that blocks ice formation in the tissues at temperatures down to −2.0°C. The protein concentration in the fish increases as temperatures in the sea fall. Common in brackish waters such as the Koksoak River of Ungava Bay, this is one of three abundant marine species at Wemindji, James Bay, in the coastal waters, along with Shorthorn Sculpins and Ogac, where temperatures seldom rise above 15°C in summer. In Hudson and James Bays it uses estuaries on a seasonal basis.

biology: Its food is known to be various crustaceans, but other aspects of its biology have not been studied. Amphipods are the



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primary food in Ungava Bay and in Creswell Bay, Somerset Island. It is eaten by Black Guillemots in Digges Sound, Hudson Bay; by Arctic Char in Hudson Bay and Ungava Bay; by Atlantic Cod at Killiniq (Port Burwell), Ungava; and by Ogac at Wemindji, James Bay. It is probably eaten by Bearded Seals at Aujuittuq (Grise Fiord), and probably at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River) on Baffin Island. It is eaten by Brünnich’s Murres and Thickbilled Murre chicks (0.07% of diet) at Akpatok Island, Ungava Bay, and by Thick-billed Murre chicks at Cambridge Point, Coburg Island. Spawning takes place in the autumn, and the eggs are demersal and up to 1.3 mm in diameter. The male guards the eggs. Eggs, some hatched, believed to be of this species were found at Wakeham Bay, Ungava, in late August. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae, possibly of this species, occur from May to June.

importance: It is not economically important. distribution: The species is found in Baffin Bay at eastern

Devon Island and southern Ellesmere Island, northern Baffin Island and its Davis Strait coast, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, Foxe Basin, Gulf of

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Boothia, Dease Strait, Dolphin and Union Strait, Amundsen Gulf, northern Banks Island, and Beaufort Sea. It is found also south to the Gulf of St Lawrence in Atlantic Canada, in northwest and southwest Greenland, the Alaskan Beaufort Sea, Chukchi Sea, East Siberian Sea, and the northern Bering Sea.

common names: Local names are Kanajuk, Kanayuk, and

Qanirkuutuk; and Kanajoq Nalinginnaq (Greenlandic). Other common names are Bull Rout, Daddy Sculpin, European Sculpin, Father-lasher, Goat Sculpin, Greenland Bullhead, Greenland Sculpin, Horny Whore, Pig-fish, Scully, Scummy, Scumpy, Sea Scorpion, Short-spined Sea Scorpion, chaboisseau commun, and chaboisseau de mer commun.

taxonomy: The species name comes from the Greek skorpios (scorpion), the root of “sculpin” in English. Myoxocephalus scorpius groenlandicus (Cuvier, 1829), described from Greenland, is sometimes recognized as a distinct subspecies in Greenland and eastern North America. Cottus porosus Valenciennes in Cuvier and Valenciennes, 1832, was described from Baffin Bay, between west Greenland and Baffin Island, and is a synonym. Acanthocottus labradoricus Girard in Storer, 1850, from the Nelson River near York Factory in Hudson Bay, is a synonym (or a synonym of Myoxocephalus quadricornis, as authorities differ). Cottus glacialis Richardson, 1855, described from Northumberland Sound, 76°53' N, is a synonym. description: This species and its four Arctic relatives in the

Distribution of Myoxocephalus scorpioides

sources: Johansen (1927a); Vladykov (1933a); Dunbar & Hildebrand (1952); Tuck & Squires (1955); Tuck (1960); Hargens (1972); Sekerak, Thomson, Bain, & Acreman (1976); Morin, Dodson, & Power (1980); Finley & Evans (1983); Gaston et al. (1985); Morin, Hudon, & Whoriskey (1991, 1992); D.B. Stewart et al. (1991); Ponton et al. (1993); Wöhrmann (1997).

Myoxocephalus scorpius (Linnaeus, 1758)

Shorthorn Sculpin, chaboisseau à épines courtes

genus Myoxocephalus are distinguished from other Sculpins by the upper preopercular spine being a simple straight point; the vomer bone in the roof of the mouth bears teeth, but palatine teeth are absent; there are no oblique folds on the lower flank; and the lateral line lacks plates. The species is separated from its relatives by having three preopercular spines (one pointing forward to downward, two pointing backward); the uppermost preopercular spine is less than twice as long as the one below; the spiny dorsal fin origin is not obviously in front of the rear operculum edge; the pectoral finrays number usually 16–18 (range 14–19); the total count of dorsal, anal, and pectoral fin-rays is 39–43; the frontal and parietal spines usually have stout accessory spines at the base; the papillae on top of the head are small and scattered; and there are spines over the eyes. The first dorsal fin spines number 7–12, usually 9–11; the second dorsal fin-rays number 12–20, usually 14–17; and the anal fin-rays number 9–16, usually 11–14. The lateral-line pores number 39–45. There is a row of plates above, and a row below, the lateral line. The taxon groenlandicus has 37–39 vertebrae, as opposed to 34–36 in scorpius. The overall colour is dark green to greenish or reddish brown or yellow to almost black with dark mottling, always brown-based. The belly is white to yellowish or light orange in females, and reddish orange or cherry red with white spots in mature males. The fins are brown, green, or yellow and have paler spots or bars. Males have large pale white to yellowish spots on the lower flank. They are more brightly coloured and have larger fins than females in the breeding season. The species reaches a reputed 90.0 cm in total length, but other records give 60.0 cm in total length and the fish are commonly smaller.

habitat: Shorthorn Sculpins are found in coastal waters and on Myoxocephalus scorpius

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banks offshore down to 494 m in Arctic Canada. In Hudson Strait, 54 collections had a depth range of 157–382 m. Elsewhere it is

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Myoxocephalus scorpius

found from the shallows to 550 m. They are usually caught on mud, sand, pebble, stone, gravel, shell, or weed bottoms and are common around harbours and wharves and in tide pools. They may be more abundant than bottom trawls indicate because of an ability to hide in rock crevices and dense vegetation. This species has been described as sluggish and swims slowly. Large specimens were abundant under Laminaria fronds in Resolute Bay, Cornwallis Island, and could be picked up by hand. At Cape Hatt in northern Baffin Island, as size increased, the fish showed a preference for greater algal canopy cover. With Arctic Staghorn Sculpins, they are the commonest fish species in northwest Baffin Bay and eastern Lancaster Sound. At Cape Hatt this species was the most abundant at 29% of the total number of specimens collected in subtidal areas. They are common and widely distributed in coastal habitats of the Beaufort Sea but are not abundant below 15–20 m. Near Tuktoyaktuk 75% of the fish biomass in trawl surveys is this species. They have been caught in a wide range of temperatures and salinities. This is one of three abundant marine species at Wemindji, James Bay, in coastal waters, along with Arctic Sculpins and Ogacs. It is the dominant marine species in estuaries of James Bay. In Hudson and James Bays it uses estuaries on a seasonal basis, moving offshore as temperatures increase. In Ungava Bay they are very common in the shore zone and



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in tidal pools. In the Beaufort Sea they may overwinter in nearshore areas but move into slightly deeper water as nearshore areas freeze. This sculpin in Arctic waters – Aujuittuq (Grise Fiord), Ellesmere Island – has a biological anti-freeze year-round that lowers the freezing point of body fluids below that of sea water. Its blood viscosity at Resolute Bay is less than that of fish species from more temperate waters, an advantage in low-temperature environments since blood flows more freely. At Baffin Island the plasma freezing point is −1.60°C, and in James Bay the species had high freezing-point depressions and could tolerate −1.86°C.

biology: Its food includes benthic, epibenthic, and pelagic

organisms. The diet comprises crustaceans such as gammarids (Onisimus and Gammarus), mysids, cumaceans, decapods, and isopods; sea urchins; molluscs such as pelecypods and gastropods (Margarites helicinus in Dolphin and Union Strait, for example); polychaete worms; kelp; and such fish as Capelin, Herrings, Eelpouts such as the Fish Doctor, Sculpins such as Fourhorn Sculpin and its eggs, Arctic Staghorn Sculpin, Triglops sp., Snailfishes, Atlantic Spiny Lumpsucker, and small Arctic Cod. The food choice is often governed by availability and by environmental conditions. At Killiniq (Port Burwell) in late August food was almost exclusively

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pteropods (Limacina). In Cumberland Sound, Baffin Island, Shorthorn Sculpins were found to be feeding on planktonic amphipods (Parathemisto libellula and Pseudalibrotus glacialis), while in nearby Pangnirtung Fiord they were eating mainly benthic snails (Littorina saxatilis and Margarites umbilicalis) and clams (Modiolaria discors). Drifting ice in the Sound probably reduced visibility, making it difficult for the Sculpins to locate food on the sea bed, but the illuminated and brightly coloured plankton were easily detected. In Frobisher Bay the diet comprises mostly (99.5%) two to three amphipod species (Onisimus and Gammarus spp.). In Cambridge Bay its food is primarily bivalves, sea urchins, polychaetes (especially Pectinaria sp.), amphipods, and other invertebrates. In the Beaufort Sea and the waters of southeastern Baffin Island, fish is the most important diet item, mostly by cannibalism. At Cape Hatt, northern Baffin Island, the pelagic pteropod Limacina helicina form about 50% of the diet. In Brevoort Harbour and nearby waters of eastern Baffin Island, stomach contents include Lycodes sp., Arctic Staghorn Sculpin, Spatulate Sculpin, Shorthorn Sculpin, Triglops sp., and Liparis sp. In Bathurst Inlet the Fish Doctor is eaten. Shorthorn Sculpins are eaten by Arctic Char in Hudson Bay, in Frobisher Bay, and at Herschel Island in the Beaufort Sea, and by Black Guillemots in Digges Sound, Hudson Bay, and the Alaskan Beaufort Sea. Arctic Cod and Arctic Char eat them in Wakeham Bay, and Atlantic Cod at Killiniq (Port Burwell), Ungava. In Barrow Strait, Nunavut, they are eaten by Thick-billed Murres and Black Guillemots, and Thick-billed Murres feed this fish to their chicks at Coats Island, Nunavut. They are eaten by Bearded Seals at Aujuittuq (Grise Fiord), and at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River) on Baffin Island, and by Ringed Seals in Cumberland Sound. Young fish are eaten by Inconnu, ciscoes, whitefishes, char, and gulls. Females are larger and older than males at maturity. Off southeastern Baffin Island, sexual maturity is reached at age three years for males and age three to four years for females. At Cape Hatt males matured between three and six years of age (15 cm), and females began to mature at three to four years (17 cm). Its lifespan may exceed 20 years, but most large fish in the Beaufort Sea are eight to nine years old, and collections in southeastern Hudson Bay are mostly four to eight years old. Fish at Cape Hatt reach nine years of age, and at Peterhead Inlet and the Sylvia Grinnell estuary in Frobisher Bay reach 19 years for females and 15 years for males. Fish caught in September in Richmond Gulf were ripening, and eggs were seen at Wakeham Bay, Ungava, in late August. Late September or October appears to be the spawning time at Shepherd Bay on the Boothia Peninsula. Spawning takes place in November and December at Newfoundland when temperatures are about 2.7°C–3.2°C. Depth range is 6–11 m, and adhesive eggs are laid in V-shaped crevices. The eggs are up to 2.5 mm in diameter, pale pink to purplish pink or a reddish yellow in colour, and number up to 60,976 in each female. Males guard the eggs until hatching. The fry are pelagic but may also be found near the sea bed. In Alaska the Fourhorn Sculpin spawns during summer. Larvae are found in the plankton from the end of April to the start of July. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae, possibly of this species, occur from May to June.

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Arsenic levels in muscle and liver tissues are 8–16 times higher than the maximum permissible level for this species in Strathcona Sound near a mine. This species is caught easily by young anglers but is a bait stealer and a nuisance to anglers after more desirable species. It will grunt when taken out of the water.

importance: It has been utilized for human food on a local basis

in Labrador and was once important as bait for use in lobster traps. Occasionally it is caught for sport. It forms part of the by-catch in the Killiniq fishery of Ungava Bay and is the most important sculpin there.

distribution: The species is found in Baffin Bay, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, Foxe Basin, Gulf of Boothia, through the Arctic islands, Queen Maud Gulf, Dease Strait, Melville Sound, Bathurst Inlet, Dolphin and Union Strait, Amundsen Gulf, and the Beaufort Sea. It is found also from the Alaskan Beaufort Sea, Chukchi Sea, and to northern British Columbia on the Pacific coast; all coasts of Greenland; and south to New York on the Atlantic coast. It occurs also in northern Europe and along the Eurasian coast.

Distribution of Myoxocephalus scorpius

sources: Kumlien (1879a); Johansen (1926, 1927a); Vladykov

(1933a); Hildebrand (1948); Dunbar & Hildebrand (1952); Grainger (1953); Scholander, van Dam, Kanwisher, Hammel, & Gordon (1957); M.S. Gordon, Amdur, & Scholander (1962); Blanc & Hureau (1968); Macpherson (1971); MacInnis (1973); Moore & Moore (1974); Talbot & Legendre (1977a); Bohn & Fallis (1978); den Beste & McCart (1978); Bradstreet (1980); Morin, Dodson, & Power (1980); Fletcher, Addison, Slaughter, & Hew (1982); LGL Limited, Toronto,

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& Environmental Sciences Limited, Vancouver, BC (1982); Fabijan (1983); Finley & Evans (1983); D.B. Stewart & Bernier (1983); Gaston et al. (1985); M.S. Graham, Fletcher, & Haedrich (1985); Kao, Fletcher, Wang, & Hew (1986); Morin & Dodson (1986); Gaston et al. (1987); Crawford (1989); Hudon (1990a); D.B. Stewart et al. (1991); Atkinson & Percy (1992); Morin et al. (1992); Ponton et al. (1993); Whoriskey et al. (1994); Bright, Grundy, & Reimer (1995); Wöhrmann (1997); Dick, Chambers, & Gallagher (2009); Divoky, Chilton, & Czapanskiy (2012).

Triglops murrayi Günther, 1888

Moustache Sculpin, faux-trigle armé

habitat: This sculpin is benthic to epibenthic on sandy bottoms at

Triglops murrayi

common names: A local name is Murrays Knurulk (Danish/ Greenlandic). Other common names are Mailed Sculpin and fauxtrigle maillé.

taxonomy: The genus comes from the Greek trigla (the classical name for the unrelated European fish Mullus barbatus) and ops (appearance). The species is named after the pioneering Scottish oceanographer, marine biologist, and limnologist Sir John Murray (1841–1914), born in Cobourg, Ontario, who was a naturalist on the Challenger expedition (1872–6) to survey the deep oceans and was editor of the ensuing reports. Triglops ommatistius terraenovae Gilbert, 1913, described from Newfoundland (45°09'30" N, 49°48'30" W), is a synonym. description: This species and its relatives the Bigeye Sculpin

and the Ribbed Sculpin are distinguished from other Sculpins in the Arctic by having the upper preopercular spine a simple point without spinules; the vomer is toothed, but the palatine lacks teeth; the lateral-line plates have backward-pointing spines; the anal finrays number 18 or more; and the lower flank has oblique skin folds with serrated edges. It is distinguished from its relatives by having the jaws equal or the upper jaw extending slightly beyond the lower; the breast and sides of the abdomen are unpigmented; the peritoneum is unpigmented or with widely spaced melanophores; the body is light in colour, with 4–5 saddles (1 beneath the spiny



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dorsal fin, 2–3 beneath the soft dorsal fin, and 1 on the caudal peduncle); the pectoral fin-rays number 16–21, usually 16–19; the third pelvic fin-ray is longer than the second ray; the orbit diameter is small, equalling the snout length and less than the postorbital distance; the dorsal row of scutes is well developed; the caudal fin has 3–6 narrow, dark bars; the second dorsal fin-rays number 18–26, usually 19–24; the lateral-line plates number 44–49; the second pelvic fin-ray is longest in males; and males have a black spot at the rear of the spiny dorsal fin. The first dorsal fin spines number 9–12, and the anal fin-rays 18–27. The lower 6–7 pectoral fin-rays are finger-like. The middle and inner pelvic fin-rays are equal in length, or the middle ray is shorter. There are small spiny scales below the dorsal fin. The overall colour is dark brown, olive, pinkish or grey silver with yellowish, orange, or white lower flanks and belly. The back has large saddles or blotches, small spots or blotches, or both. The lower, posterior flank may have dark spots or short bars. The head is often spotted. The black iris of the eye is rimmed with a fine gold-coloured line. The first dorsal fin has a black spot at the margin between spines 1 and 2. The dorsal, pectoral, and caudal fins have thin dark bars. In males the dorsal fin spot is well developed, there is a separate spot in the posterior part of the same fin, and white patches develop on the breast and flanks. There is a black bar above the upper jaw (the “moustache”). The lower part of the head may be reddish below and posterior to the moustache. The peritoneum is lightly spotted and silvery. The species reaches 20.0 cm in standard length.

0°C–12°C, usually in marine conditions but sometimes in brackish water down to 23.6‰ in Europe. Fifty-two collections from Ungava Bay had a depth range of 115–470 m, and 109 collections from Hudson Strait a range of 0–530 m. The maximum depth in the eastern Canadian Arctic is 823 m. Larvae are pelagic under ice from the beginning of December to mid-May and have been caught at −1.4°C to 3.6°C in Europe.

biology: Its food is various crustaceans such as amphipods,

mysids, copepods, cumaceans, decapods, euphausiids, and isopods; chaetognaths; ascidians; pelecypods; and polychaete worms, some taken above the bottom. Copepods dominated in a study of Beaufort Sea and southeastern Baffin Island fish, with euphausiids second in importance. It is eaten by Atlantic Cod in Ungava Bay, by Thick-billed Murres and Black Guillemots chicks in Digges Sound, Hudson Bay, and at Coburg Island, and by Thick-billed Murre chicks at Coats Island, Hudson Bay. Females grow larger than males. Spawning occurs in summer and fall, perhaps into winter. In the Gulf of Maine in October, ripe females have amber eggs numbering up to 2,739. There may be several spawnings in a season. Each egg clutch has 19–20 eggs in the White Sea. The yellow-pink eggs may be up to 2.2 mm in diameter, 2.5 mm in a clutch. At Kuujjuarapik in southeastern Hudson Bay, Triglops post-larvae, possibly this species, occur in May and June.

importance: It is not economically important.

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Triglops murrayi

distribution: The species is found sparsely in Baffin Bay, including at Nirjutiqavvik (Coburg Island), and in northern Davis Strait and is more common in southern Davis Strait and in Frobisher Bay, Hudson Strait, Ungava Bay, and Hudson and James Bays. A specimen from Kugmallit Bay, Beaufort Sea (CMNFI 1989-0462) keys out to this species but is the only record west of Hudson Bay. An anomaly in collection date may indicate confusion, and this record is neither accepted nor mapped. The species is found in the western Atlantic Ocean south to the Gulf of Maine and in southwest and southeast Greenland and northern Europe. sources: Andriashev (1949); Musick & Able (1969); Gaston et al. (1981); Gaston et al. (1985); Gaston (1987); Hudon (1988, 1990a); Atkinson & Percy (1992); Pavlov et al. (1992); Ponton et al. (1993); Pietsch (1994).

Distribution of Triglops murrayi

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Triglops nybelini Jensen, 1944

Bigeye Sculpin, faux-trigle aux grands yeux

common names: A local name is Nybelins Knurulk (Danish/ Greenlandic). Other common names are Mailed Sculpin and fauxtrigle de Nybelin. taxonomy: The species is named after Ovar Nybelin, director of

the Naturhistoriska Museet, Göteborg, and author of Våra fiskar.

description: This species and its relatives the Moustache Sculpin and the Ribbed Sculpin are distinguished from other Sculpins in the Arctic by having the upper preopercular spine a simple point without spinules; the vomer is toothed, but the palatine lacks teeth; the lateral-line plates have backward-pointing spines; the anal finrays number 23 or more; and the lower flank has oblique skin folds with serrated edges. It is distinguished from its relatives by having the lower jaw extending beyond the upper jaw; the breast, sides of the abdomen, and the peritoneum (dissection required) have dense melanophores; the body is dark, but saddles beneath the dorsal fins and on the caudal peduncle are absent or faint; the pectoral fin-rays number 20–23; the second ray of the pelvic fin is longest; the orbit diameter is large, exceeding snout length and equal to or longer than postorbital distance; and the dorsal row of scutes reduced. The first dorsal fin spines number 9–12, the second dorsal finrays 24–29, and the anal fin-rays 23–28. The middle pelvic fin-ray is the longest. Plate-like lateral-line scales number about 45–52. The

back and top of the head are dark brown in colour, and the flanks are lighter brown. Much of the head and body has dense melanophores. The body has two to five characteristic oblique pigment lines below the mid-flank. The caudal fin base has dorsal and ventral spots. The fins are mostly clear except for thin bars on the lower pectoral fin-rays. In males the flank lines merge into two stripes, and the pectoral fin has black markings. The peritoneum is densely spotted and black. The species reaches 17.0 cm in standard length.

habitat: It is sometimes found inshore on mud bottoms but is

usually bathyal, generally below 135 m down to 1,279 m, with juveniles rarely in shallow water less than 37 m deep. Temperatures are below 0°C, down to −1.8°C. Salinity is usually high. Capture depth in Davis Strait and southern Baffin Bay is 146–1,353.5 m at 0.18°C to 5.4°C, and in Hudson Strait is 0–356 m. Bigeye Sculpins have antifreeze peptides in their blood that help them resist low temperatures.

biology: Its food is crustaceans such as amphipods (hyperiids)

and mysids, including planktonic species. It is eaten by Thick-billed Murre and Black Guillemot chicks in Digges Sound, Hudson Bay, and at Coburg Island and by Thick-billed Murre chicks at Prince Leopold Island off Somerset Island and at Cape Hay, Bylot Island. It is also taken by Atlantic Cod and, in Davis Strait, probably by Ringed Seals. Females grow larger than males. Spawning takes place in summer, and females may have up to 1,000 pink or amber eggs of up to 3.0 mm in diameter.

importance: It is not economically important.

Triglops nybelini



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Triglops nybelini

distribution: The species is found in Smith Sound, at Nirjutiqavvik (Coburg Island), in Baffin Bay, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, the Arctic islands, Dease Strait, Bathurst Inlet, Amundsen Gulf, and the Beaufort Sea. It may also be in the Alaskan Beaufort Sea, but this has not been confirmed. It is found also on all coasts of Greenland, south to the Gulf of Maine as larvae, on the northern coast of Eurasia, and probably circumpolar. sources: Jensen (1944); Andriashev (1949); Musick & Able (1969); Karrer (1973); MacLaren Marex (1979a, 1979b); Gaston et al. (1981); Gaston & Nettleship (1981); Gaston et al. (1985); Gaston (1987); Pietsch (1994); Ponomarenko (1995b); Wöhrmann (1997); Treble et al. (2000); Jørgensen et al. (2005).

Distribution of Triglops nybelini

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Triglops pingelii Reinhardt, 1837

Ribbed Sculpin, faux-trigle bardé

common names: A local name is Pingels Knurulk (Danish/

Greenlandic).

taxonomy: The species is based on a personal name (source unknown). Triglops pleurostictus Cope, 1865, described from Godhaven, western Greenland, is a synonym. description: This species and its relatives the Bigeye Sculpin and

the Moustache Sculpin are distinguished from other Sculpins in the Arctic by having the upper preopercular spine a simple point without spinules; the vomer is toothed, but the palatine lacks teeth; the lateral-line plates have backward-pointing spines; the anal fin-rays number 20 or more; and the lower flank has oblique skin folds with serrated edges. It is distinguished from its relatives by having the jaws equal or the upper jaw extending slightly beyond the lower jaw; the breast and sides of the abdomen are unpigmented; the peritoneum is unpigmented or with widely spaced melanophores; the body is light in colour with 4–5 saddles (1 beneath the spiny dorsal fin, 2–3 beneath the soft dorsal fin, and 1 on the caudal peduncle); pectoral fin-rays number 16–21; the third pelvic fin-ray is longer than

the second ray; the orbit diameter is small, equalling snout length and less than the postorbital distance; the dorsal row of scutes is well developed; the caudal fin is without bars (rarely with a single faint bar, and the upper lobe may have a black tip); the second dorsal finrays number 20–28, usually 23–26; the lateral-line plates number 47–51; the third pelvic fin-ray is longest in males; and males lack a black spot at the rear of the spiny dorsal fin. The first dorsal fin spines number 9–13, the anal fin-rays 20–28, the lower pectoral fin-rays 7–8, which are weakly finger-like, and the gill rakers 5–9. There is a sharp nasal spine and four blunt preopercular spines. The male anal papilla is long and curved forward. The back is olive brown to red brown in colour with 4–5 dark saddles that may be poorly developed. The belly is white to silvery. The dorsal fins may have thin bars formed by small spots on each fin-ray. The tail fin has small blotches dorsally and ventrally at the base and, in males, at the distal ends of the fin lobes. There is a black bar and a silvery area in front of the pectoral fin. The pectoral fins have 4–6 weak, curved bars. Males have a silver line below the lateral line, with a narrow black stripe below. In females the stripe is broken into dark blotches. The species probably reaches 24.2 cm in total length.

habitat: The Ribbed Sculpin is a benthic to epibenthic species that

is found over mud, silt, sand, shell, pebble, gravel, and rocks in colder waters at 4–100 m but as deep as 930 m. It is caught at 260–360 m in Hudson Strait and at 100–738 m in eastern Arctic Canada generally. It may be found at temperatures as low as −1.8°C at Resolute Bay,

Triglops pingelii



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Cornwallis Island, where juveniles have been observed swimming 0.5–1.0 m above the bottom, actively swimming away from divers. It has been reported at −2.0°C in Hudson Bay and to 6.0°C elsewhere in Arctic Canada. Salinities are as low as 16‰ in Russia, and it has been reported from the estuary of the Churchill River in Hudson Bay, albeit from a zooplankton tow, at 31.6‰. Larvae are pelagic. It actively forages in the water column when young and to some extent when adult.

biology: Its food is crustaceans, with amphipods and mysids often

dominating, but also including isopods, arrow worms, and more rarely some polychaete worms and fishes such as Ribbed Sculpins, Myoxocephalus sp., Arctic Cod, and Liparis sp. Mysids and fish are important food items off southeastern Baffin Island. At some localities, such as southeastern Baffin Island, larval ascidians, copepods, and pteropods dominate. In Shepherd Bay on the Boothia Peninsula kelp is found in stomach contents. In Resolute Bay, Cornwallis Island, arrow worms are almost an exclusive diet item in December, but mysids and copepods are found in June. It actively forages in the water column on arrow worms, mysids, and other species, avoiding competition with other benthic feeders. Thick-billed Murre chicks eat this sculpin in Digges Sound, Hudson Bay; at Coats Island, Nunavut; Coburg Island; Akpatok Island (26.69% of chick diet); and Prince Leopold Island, off Somerset Island. It has been recorded as being eaten by Ringed Seals and Atlantic Cod at Killiniq, Ungava Bay, and by Arctic Char in Frobisher Bay and Ungava Bay. It is probably eaten by Bearded Seals at Aujuittuq (Grise Fiord) and probably at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River) on Baffin Island. It is eaten by Brünnich’s Murres at Akpatok Island, Ungava Bay. Its lifespan is at least nine years near Baffin Island. Males predominate in populations and attain a size of up to 14 cm, but all the largest fish are female. Each female has a mean fecundity of 1,800 eggs with a diameter of 3.0 mm that may be spawned in autumn or early winter. Off southeastern Baffin Island the eggs per female numbered up to 444. The eggs are a milky red and demersal. Larvae are found in the plankton in April to June. At Kuujjuarapik in southeastern Hudson Bay, Triglops post-larvae, possibly this species, occur in May and June.

Distribution of Triglops pingelii

sources: Jensen (1944); Andriashev (1949); Grainger (1953); Tuck

& Squires (1955); Tuck (1960); den Beste & McCart (1978); Hunter et al. (1980); Gaston et al. (1981); Gaston & Nettleship (1981); Finley & Evans (1983); Green (1983); D.B. Stewart & Bernier (1983); Gillis & Allard (1984); Gaston et al. (1985); Gaston (1987); Gaston et al. (1987); Hudon (1988); D.B. Stewart et al. (1991); Atkinson & Percy (1992); Pavlov et al. (1992); Ponton et al. (1993); Pietsch (1994); Lawrence & Baker (1995); Tokranov (1995); Pietsch & Orr (2006).

importance: It is not economically important. distribution: The species is found in Nares Strait, Baffin Bay,

Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, sparsely in Hudson and James Bays (possibly in Richmond Gulf), Foxe Basin, the Arctic islands, Dease Strait, Bathurst Inlet, Dolphin and Union Strait, Amundsen Gulf, and the Beaufort Sea. It is found also in North Korea and Japan; the Alaskan Beaufort, Chukchi, and Bering Seas south to Washington State; in the North Atlantic Ocean including all coasts of Greenland; through Atlantic Canada south to Massachusetts; and in the White Sea.

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and all fins, unusually in fishes, lack branched rays. The pectoral fins are large and fan-like, sometimes with the lower rays elongated. The head may have various spines, and the area around the mouth may have various barbels. Males usually have larger pelvic and anal fins than those of females. The anus is between the pelvic fins. These are bottom-living fishes at shallow to moderate depths, with some down to 1,290 m. The barbels are used to help locate such food as crustaceans and polychaetes. The eggs are demersal and adhesive, and the young are pelagic for two to three months. They have no commercial importance although some are sold as dried souvenirs.

Family Agonidae Poachers, Poissons-alligators

Brian W. Coad

sources: Jensen (1942a); Kanayama (1991); Busby (1998); Sheiko & Mecklenburg (2004).

Poachers, Alligatorfishes, Starsnouts, and Rockheads (also called Agones in French) are found mainly in the North Pacific Ocean, with a few species being in the North Atlantic Ocean. There are about 50 species, including 21 in Canada, of which 3 are in Arctic waters. The maximum size is 42 cm although most species are much smaller. They are easily recognized by their elongate bodies covered in bony plates that are non-overlapping, fused together in rows, and often spiny or toothed. The cheek is covered by bones. The pelvic fins are thoracic (under the pectorals) and have one spine and two soft rays. There may be one or two dorsal fins. The anal fin lacks spines,

Aspidophoroides monopterygius (Bloch, 1786)

Alligatorfish, poisson-alligator atlantique

common names: A local name is Almindelig Krokodilleulk (Danish/Greenlandic). Other common names are Atlantic Alligatorfish and Sea Poacher. taxonomy: The genus comes from the Greek aspis (shield), phoreo (to bear), and eidos (likeness, form). The species name comes from the Greek monos (single) and pterygion (fin).

Aspidophoroides monopterygius



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Aspidophoroides groenlandicus Valenciennes, 1840 (Greenland), and Aspidophoroides borealis Valenciennes, 1841 (North Atlantic), are unneeded new replacement names.

description: This alligatorfish is distinguished from its relatives

by having a very slender body with one dorsal fin, no barbels, 9–11 pectoral fin-rays, and 45–53 plates in the dorsal row. There are eight plate rows anteriorly, decreasing to six posteriorly. The plates are smooth dorsally. The cheek plates are tightly arranged. There is a ridge on the mid-line of the breast. There are two spines on the snout tip. The dorsal fin has 4–6 soft rays, the anal fin 4–6 soft rays, and the pelvic fin 1 spine and 2 soft rays. The body colour is a dark brown to olive green with 4–5 dark bars along the flank and 2–3 in front of the dorsal fin. The belly is whitish. There are bars on the dorsal and pectoral fins. The caudal fin is dusky with a white margin. The dorsal, pelvic, and lower pectoral fins develop white pigmentation, probably at spawning. The species reaches 18.0 cm in total length.

habitat: It is found over pebbles, sand, mud, shell, and some-

times rock bottoms at 0–695 m and temperatures below 3°C, down to −1.07°C. An upper temperature limit is probably about 9–11°C. Fifteen collections from Hudson Strait were taken at 0–280 m, and off west Greenland they have been found at 332 m. Larvae may enter the brackish water of estuaries. This species rests with the pectorals splayed and braced on the bottom.

biology: This species is eaten by Atlantic Cod at Killiniq (Port

Burwell), Ungava, and by Haddock and halibut, among other fishes, elsewhere. Its biology is not well known, but crustaceans are an important food item, and a medium-sized female carries about 600 eggs of 1.25 mm in diameter. In the Gulf of St Lawrence their putative lifespan is up to seven years. Larvae are found in Passamaquoddy Bay in Atlantic Canada from April to June, suggesting a late fall to winter spawning season there. Gulf of St Lawrence fish are thought to spawn in mid- to late-autumn, and fecundity reaches 3,255 eggs; they feed on caprellids, hyperiids, gammarids, isopods, mysids, euphausiids, copepods, calcareous algae, and pteropods.

Distribution of Aspidophoroides monopterygius

sources: Johansen (1927a); Hildebrand (1948); Nielsen et al. (1992); Arbour, Avendaño, & Hutchings (2010).

Aspidophoroides olrikii Lütken, 1877

Arctic Alligatorfish, poisson-alligator arctique

importance: It is not economically important. distribution: The species is found from Davis Strait, Hudson Strait, and Ungava Bay, and northwest and southwest Greenland south to Massachusetts, and rarely New Jersey. There are no confirmed records from Hudson Bay and southern Ungava Bay despite some literature reports. It is found also in the North Pacific Ocean, Bering Sea, and eastern Chukchi Sea but not the Alaskan Beaufort Sea.

Aspidophoroides olrikii

common names: A local name is Olriks Panserulk (Danish/ Greenlandic).

taxonomy: The species is named after Christian Søren Marcus Olrik (1815–70), the Danish director of Greenlandic commerce and a justice of the Supreme Court. It was previously in the genus Ulcina. description: This alligatorfish is distinguished from its relatives

by having a stout body with one dorsal fin, one pair of barbels at the

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Aspidophoroides olrikii

corner of the mouth, 12–16 pectoral fin-rays, and 33–40 plates in the dorsal row. There are eight plate rows anteriorly, six posteriorly. The cheek plates are loosely arranged. The dorsal and anal fins each have 5–7 soft rays, usually 6. The pectoral fin is without finger-like modification of the lower rays, although lower membranes are more incised than upper membranes. The gill membranes are not joined to the isthmus. The dorsal part of the body is dark grey, brownish grey, or greenish, and the flanks have several vague bars, with only one in front of the dorsal fin. The belly is lighter. The dorsal fin has two dark greenish bars. The pectoral fin is barred with rows of spots. There is a bar from the eye running back and downward. Males develop milky-white areas on the dorsal and anal fins. The caudal fin has a pale edge and a pale central area. The species attains 8.6 cm in total length.

habitat: The species is found on the bottom over sand, mud, or

rock at 0–632 m. Seventy-one collections from Hudson Strait were at 0–486 m, 67 collections from Ungava Bay were at 161–436 m, and 35 collections from southern Davis Strait were at 146–632 m. They live in temperatures usually around 0°C, but as low as −1.85°C or as high as 7.5°C. Salinities of 23‰–28‰ are tolerated. It is reported from the estuary of the Churchill River in Hudson Bay, albeit from a zooplankton tow, at 31.6‰ and 27.2 m. On the Beaufort shelf they typify the > 50 m fish assemblage, presumably because they have a lesser tolerance of temperature and salinity than does the Arctic Staghorn Sculpin, which typifies < 50 m depths.

culans; turbellarians; and nemertean worms. A major food item in deeper waters of the Beaufort Sea and southeast Baffin Island was found to be siphons of the clam, Macoma calcarea, nipped off as they protrude. This alligatorfish is eaten by halibut and, at Killiniq (Port Burwell), by Arctic Char. Up to 260 demersal eggs of 1.7 mm in diameter are laid. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae occur in June.

importance: It is not economically important. distribution: The species is found in Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson Bay, James Bay, Foxe Basin, Dease Strait, Bathurst Inlet, Beaufort Sea, and Prince of Wales Strait between Banks and Victoria Islands; northwest and southwest Greenland; and south to Newfoundland. There is one record from Slidre Fiord, Ellesmere Island (J.G. Hunter, pers. comm.), which needs to be verified with specimens. It is found also off northern Eurasia, the Alaskan Beaufort Sea, Chukchi Sea, and the northern Bering Sea.

biology: Its food is often small crustaceans such as gammarid

amphipods, isopods, copepods, ostracods and cumaceans; sipun-



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Leptagonus decagonus (Bloch and Schneider, 1801)

Atlantic Poacher, agone atlantique

Leptagonus decagonus

common names: A local name is Kanajorlak (Greenlandic). Distribution of Aspidophoroides olrikii

sources: Frost & Lowry (1983); Atkinson & Percy (1991, 1992);

Ponton et al. (1993); Lawrence & Baker (1995); Majewski et al. (2013).

Other common names are Atlantic Sea Poacher and Northern Alligatorfish.

taxonomy: The genus comes from the Greek leptos (slender or thin), a- (without), and gonia (joint or angle), that is, rigid. The species name comes from the Greek deka (ten) and gonia (angle). It was also placed in the genus Agonus. description: This poacher is distinguished from its relatives by

having two dorsal fins and five pairs of barbels around the mouth. The barbel at the rear of the jaw is branched. There are 41–48 keeled or spiny plates in the dorsal row. A supra-lateral plate row is present. There are 4–8 first dorsal fin spines and 5–8 second dorsal fin soft rays. The anal fin has 5–8 soft rays, and the pectoral fin has 13–17 rays. There are strong spines on the head and the back. The overall colour is a yellowish grey to brownish grey or reddish grey, with two to three darker bars along the flank. The dorsal, anal, and pectoral fins

Leptagonus decagonus

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Leptagonus decagonus

are brown black distally. The caudal fin is dark brown. A black band runs from the tip of the snout through the eye and fades posteriorly on the head. The species attains about 25.0 cm in length.

habitat: This species is benthic, possibly bathypelagic, and is

usually found over mud, clay, and sand, often with some stones or pebbles, from 2 m to 968 m, with deepest records in Baffin Bay. In Davis Strait it is reported at 412–820 m and 0.5–1.1°C. Young of the year are caught in open-water trawls off southeastern Baffin Island in late summer. It lives in colder waters down to −1.85°C on the Labrador coast, and probably similar temperatures in the Arctic, but as warm as 5°C elsewhere. It may be found in salinities as low as 27.4‰.

off Point Barrow in the Alaskan Beaufort Sea, in the Chukchi and Bering Seas, and the Sea of Okhotsk. It is found also on all coasts of Greenland, south to the Grand Banks, rarely Sable Island in Atlantic Canada, and in northern Europe. General reports for James Bay are not confirmed by specimens.

biology: Its food is various crustaceans both pelagic and ben-

thic, molluscs, polychaete worms, and other bottom organisms. As there are some records of pelagic crustaceans in the diet, it may well swim off the bottom too. This species is eaten by Thick-billed Murres at Coats Island in northern Hudson Bay and by Bearded Seals at Aujuittuq (Grise Fiord) and at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River) on Baffin Island. It is also food for Greenland Halibut in Davis Strait. Spawning is suspected to occur in spring and summer. The eggs are pink, number up to 1,750, and may be 2.2 mm in diameter. Fry are pelagic.

importance: It is not economically important. distribution: The species is found in Smith Sound, near Aujuittuq (Grise Fiord); Ikpiarjuk (Arctic Bay) and Mittimatalik (Pond Inlet) on northern Baffin Island; Baffin Bay and Ellesmere Island, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, sparsely in Hudson Bay (the north and southeast), Dease Strait, Bathurst Inlet, Amundsen Gulf, and the Beaufort Sea;



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Distribution of Leptagonus decagonus

sources: den Beste & McCart (1978); Finley & Evans (1983);

Chumakov & Podrazhanskaya (1986); Nielsen et al. (1992); Ponomarenko (1995); Braune et al. (2014).

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Family Psychrolutidae Fathead Sculpins, Chabots veloutés

Brian W. Coad

best known from the North Atlantic and North Pacific Oceans and off South Africa. Their food is both benthic and planktonic invertebrates and fishes. One species, unidentified, is eaten by Glaucous Gulls at Seymour Island in the Canadian Arctic. Their eggs sink and are quite large (2.0–4.5 mm in diameter). They have no commercial importance.

sources: Jensen (1952b); Nelson (1982); Jackson & Nelson (1998).

Cottunculus microps Collett, 1875

Polar Sculpin, cotte polaire

Fathead, Soft, or Blob Sculpins are found in the Atlantic, Indian, and Pacific Oceans. There are about 30 species, including 8 species in Canada, of which 3 are in the Arctic. The maximum size is about 70.0 cm. They are characterized by scaleless bodies that may have plates covered in prickles; there is a loose skin over a clear and gelatinous layer; skin often covers the dorsal and anal fin-rays; there is a reduced lateral line with less than 21 pores; the pelvic fins have 1 spine and 3 soft rays; the spiny dorsal fin is small and partly obscured by skin; the dorsal fins are usually continuous (not separated into spiny and soft parts as in Sculpins), although the fin bases are nearly separate in some species; and various skeletal features. These fishes are benthic on soft bottoms and are found from shallow water down to 2,800 m. They are rare in tropical waters and are

common names: A local name is Almindelig Paddeulk (Danish/

Greenlandic). Another common name is Arctic Sculpin.

taxonomy: The genus comes from the Greek kotta (head) and kottos (the old name for the European freshwater Cottus gobio) and from the Latin -unculus (little). The species name comes from the Greek mikros (small) and ops (eye). Cottunculus sadko Essipov, 1937 (Sadko Sculpin, cotte de Sadko), is a synonym, based on younger specimens. description: This species is distinguished by having spines

on the top of the head (occipital spines that are usually blunt, and the rearmost of two spines at the upper rear of the eye are strongly developed) and on the preopercle; the skin has minute spines (the spiny area is greater than the naked area); the head is rounded in

Cottunculus microps

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Cottunculus microps

dorsal view, not compressed anteriorly, and the head width is usually greater than the head length; 9–11 anal fin-rays; and 3–4 bands on the body. The dorsal fin spines are weak and number 6–9, soft rays 13–16; the pectoral fin-rays number 17–20, and the pelvic fin-rays 3. The lateral-line pores number 11–15. The cranial arches are usually as wide as they are long, and the tabular bones have contiguous bases. The overall colour is pale, but the bands are broad and dusky, extending onto the spiny and soft dorsal fins. The head is blotched and spotted and often has a dark band across the snout. The pectoral fin has small dark spots, and the caudal fin has irregular bars or spots. The species reaches 30.0 cm in standard length but is usually smaller.



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habitat: It is usually caught at 165–895 m – Davis Strait and Baf-

fin Bay fish were taken at 290 m to 1,342 m and −1.29°C to 3.9°C, somewhat cooler than its relative – and generally down to 1,450 m in Davis Strait. In Ungava Bay 13 collections were caught at 158–649 m. It has been recorded from 350–500 m in the Canadian Beaufort Sea.

biology: Elsewhere in its distribution it is reported to eat

polychaete worms, amphipods, mysids, and sea spiders. Spawning in the Barents Sea takes place in June and July, and probably earlier, and 124–220 eggs of 4.5–5.0 mm in diameter are released. Spawning may be extended or intermittent. Females grow to larger sizes than males.

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importance: It is not economically important. distribution: The species is found from Smith Sound at the

north of Baffin Bay, Davis Strait, eastern Hudson Strait, western Hudson Strait (a single record), Ungava Bay, the Beaufort Sea, all coasts of Greenland, and south to New Jersey. It is found also around Iceland and Jan Mayen Island, in northern Europe, and in the Kara and Laptev Seas. Chukchi and Alaskan Beaufort Sea records (Mecklenburg & Mecklenburg, 2012) were thought to be C. sadko, but this species is now a synonym of C. microps.

Cottunculus thomsonii (Günther, 1882)

Pallid Sculpin, cotte blême

common names: A local name is Thomsons Paddeulk (Danish/ Greenlandic).

taxonomy: The species is named after Sir Charles Wyville Thompson (1830–82), the scientific director on the deep-sea voyage of the HMS Challenger. description: This species is distinguished by having spines on

the top of the head and on the preopercle; the skin is smooth with only scattered spines (the area covered by spines is less than the naked area); there are no bands on the body; the eyes are relatively large (the orbit diameter is equal to or greater than the bony interorbital width); and anal fin-rays number 12–14. The dorsal fin spines number 6–8, are flexible, and in large fish are reduced in size, and the dorsal soft rays number 14–17. The pectoral fin-rays number 21–23. The lateral-line pores number about 18–19. The terminal chin pore is usually a paired structure. The head is knobbed and ridged. The overall colour is a grey brown to a reddish brown with translucent skin. The body is dark posteriorly. The tips of the pectoral fin membranes are black. The species reaches about 43.0 cm in length.

habitat: The species is reported generally at depths of 100–1,600 m and in Davis Strait at 365–1,369 m. It is caught on sand and ooze bottoms at temperatures of 3.12°C–4.4°C in Davis Strait, somewhat warmer than the habitat of its relative.

Distribution of Cottunculus microps

sources: Karrer (1973); Hudon (1990a); Ponomarenko (1995);

Treble et al. (2000); Jørgensen et al. (2005); Byrkjedal, Hadler-Jacobsen, Rees, & Orlov (2014).

biology: The biology is mostly unknown although its food is small benthic invertebrates.

importance: It is not economically important.

Cottunculus thomsonii

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Cottunculus thomsonii

distribution: The species is found in Baffin Bay, Davis Strait, southwest and southeast Greenland, and south to South Carolina. It is found also in the eastern North Atlantic Ocean. source: Karrer (1973).

Distribution of Cottunculus thomsonii



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Psychrolutes phrictus Stein and Bond, 1978

Giant Blob Sculpin, chabot maculé

common names: Other common names are Blob Sculpin and

Giant Blobsculpin.

taxonomy: The genus comes from the Greek psychros (cold) and loutron (bathe), that is, one who bathes in cold water. The species name comes from the Greek phriktos (causing one to shudder), in reference to its grotesque appearance. description: This species is distinguished by the lack of head

and preopercle spines, an interorbital width that is greater than twice the exposed eye diameter, and a very large head, 41%–61% of standard length.

The body is very globular with relatively tough and thick skin. The head and body have small scattered cirri. Small fish (< 5.0 cm in standard length) have prickles as well as cirri on the head and body. There are no head or preopercular spines. The upper jaw overhangs the lower jaw, and the maxilla extends back to mid-eye level or beyond. The gill membranes are broadly attached to the isthmus without a posterior fold. Vomerine and palatine teeth are absent. The anus is midway between the anal and pelvic fin bases. Small fish have a proportionately larger head, more-exposed eyes, and a narrower interorbital space. The spiny and soft dorsal fins are continuous, with the spiny section buried in tissue. The dorsal and anal fins do not extend onto the caudal fin base. The dorsal fin spines number 7–9, and soft rays 19–20; the anal fin-rays number 12–14; and the pectoral rays number 22–26, usually 24–25. The lateral-line pores number 12–14, total gill rakers 9–13, and vertebrae 33–36. The overall colour is greyish to blackish, and it may be indistinctly mottled with white. There are no bands or other patterns on the body.

Psychrolutes phrictus

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Psychrolutes phrictus

The snout and lower head are paler than the rest of the body. Large fish have a white head anteriorly. The peritoneum, stomach, opercular cavity, and mouth are pale. The species attains 70.0 cm in total length and 9.5 kg in weight.

habitat: This species has been caught in Canadian waters at 998

m and 1,001 m on the bottom and generally at 480–2,800 m. It is found on middle and lower continental slopes. Larvae and juveniles are thought to be pelagic.

biology: Its food comprises sea pens, crabs, molluscs, ophiuroids,

and cnidarians. Otoliths of pelagic fishes have also been found in one fish, but the prey may have been swimming near the bottom. Octopus beaks in the stomach and sucker marks on the skin show that cephalopods are successfully eaten. A plastic bag was found in one fish. Other aspects of biology are poorly known.

importance: It is not economically important. distribution: The species is found in the Beaufort Sea, based on two records caught in August 2012 at 71º51'09" N, 131º29'25" W (CMNFI 2013-0029) and 71º51'16" N, 131º29'57" W (CMNFI 20130030). It is found also from the Bering Sea south to California in the east and Japan in the west.

Distribution of Psychrolutes phrictus

sources: Stein & Bond (1978); Matarese & Stein (1980); Peden & Ostermann (1980); Yabe, Maruyama, & Amaoka (1983).



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Cyclopteropsis jordani

Family Cyclopteridae

Soldatov, 1929

Smooth Lumpfish, petite poule de mer douce

Lumpfishes, Poules de mer

Brian W. Coad

common names: None. taxonomy: The genus comes from the Greek kyklos (round), pteron (fin), and opsis (similar to, shape).The species is presumably named after David Starr Jordan (1851–1931), an American ichthyologist and a co-author of The Fishes of North and Middle America. description: This species is distinguished from its relatives by

Lumpfishes or Lumpsuckers are found in cold to temperate marine waters of the northern hemisphere. There are 28 species, including 8 in Canada, of which 4 are in the Arctic. The maximum size is 70 cm. They are distinguished from the related Snailfishes by having fewer vertebrae (29 or less, compared to 36 or more), a globular or globose body, usually two short dorsal fins (the spiny first dorsal fin may be embedded in thick skin or absent), and a short anal fin. The body is often covered with tubercles of various sizes on a wrinkled skin. There are no scales. The pelvic fins are modified into an adhesive, sucking disc. There are two pairs of nostrils. The gill slit is small and usually above the pectoral fin base. The teeth are small and conical and arranged in rows or bands. Young Lumpfishes are often hard to identify as they differ in appearance from adults, lacking some of the key characters. The position and development patterns of tubercles are sexually dimorphic, and males and females have been identified as distinct species. Tubercles may develop with age or disappear. Some species have few available specimens to clarify this situation. The adhesive disc is used to cling to rocks and seaweeds. A force of 13.3 kg is necessary to overcome the suction of a 39 cm Lumpfish (Cyclopterus lumpus). They can inflate their bodies with air or water, perhaps as a defensive mechanism. Some species live pelagically in deep waters and migrate long distances to spawn on a coast. Adult Lumpfishes are found as deep as 900 m but are usually at 100–400 m. Their food is various crustaceans, molluscs, and worms. Pelagic species feed on slow-moving prey such as medusae and ctenophores. Females lay eggs in spongy masses that adhere to rocks and seaweeds, and males of some species guard the eggs. Some species migrate long distances to lay their eggs. The Lumpfish is used as a source of “caviar” because the eggs are large. The flesh of male Lumpfish is tasty when smoked.

sources: Soldatov & Popov (1929); Jensen (1944); Lindberg &

Legeza (1955); Ueno (1970); Able & Irion (1985); Mecklenburg & Sheiko (2003); Hatano, Abe, Wada, & Munehara (2015).

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having the gill opening above the pectoral fin; about five or more large blunt tubercles restricted to the rounded anterior half of the body; usually no supplemental pores above the lateral line; the first dorsal fin rounded with rays unequal (the middle one being the longest); and a postorbital tubercle row present, extending onto the body. The dorsal fin spines number 6, and soft rays 12–13; the anal fin-rays number 10–12, and the pectoral fin-rays 23–24. Each anterior tubercle has a few blunt prickles. The first two tubercles of the postorbital row are obvious, but the rest are weakly developed. A postbranchial row runs from the gill opening, arcing down posteriorly to four large tubercles of the circumpectoral row. A pair of tubercles is in front of the anal fin and between the dorsal fins. Young lack tubercles. The species attains about 6.1 cm in standard length.

habitat: This is a sedentary species, attached by the adhesive disc,

and found down to 95 m. It is reported from over mud and clay bottoms in the Canadian Arctic at 68 m.

biology: As there are few specimens, little is known of its biology.

Food items include polychaete worms, crustaceans, and molluscs. The species is oviparous, with large, adhesive eggs laid in protected areas and guarded by an adult. It may be eaten by Ringed Seals at Admiralty Inlet and by Thick-billed Murres at Prince Leopold Island, off Somerset Island.

importance: It is not economically important. distribution: The species is found in Frobisher Bay, Ungava

Bay, Admiralty Inlet off Lancaster Sound (as the exact location is not known, it is not mapped), at Prince Leopold Island off Somerset Island, and at the type locality in the Kara Sea. General reports for Hudson Strait are not confirmed by specimens. The Prince Leopold Island locality is based on a literature record (Gaston & Nettleship, 1981) and on the fish being a food item (see above), and so it could be a misidentification. The Ungava Bay specimen (CMNFI 1960-0449) has been lost. The Admiralty Inlet specimen (University of Washington 42927, formerly University of British Columbia BC 59-350) was found dead in a seal hole, and its identity has been verified by several workers on Lumpfishes.

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Cyclopteropsis jordani

Cyclopteropsis mcalpini (Fowler, 1914)

Arctic Lumpsucker, petite poule de mer McAlpine

common names: A local name is Dværgstenbider (Danish/ Greenlandic).

taxonomy: The species is sometimes spelled macalpini, incorrectly. It is named after Charles Williston McAlpin (1865– 1942), Secretary of Princeton University, who helped to secure the collection for the university. This species was identified aboard ship, and no voucher specimen was kept. Its presence in neighbouring Greenland waters occasions its inclusion here. description: This species is distinguished from its Arctic and

Distribution of Cyclopteropsis jordani

sources: Ellis (1962); Gaston & Nettleship (1981); Gaston (1985).



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Atlantic relatives by the gill opening lying above the pectoral fin; the compressed, anterior half of the body having 5–6 large and blunt tubercles that bear many minute prickles; usually no supplemental pores above the lateral line; a rectangular first dorsal fin with rays equal in length; and a postorbital tubercle row absent. The head is pointed, and the mouth is strongly oblique. The dorsal fin spines number 7, the soft rays 11, the anal fin-rays 12, and the pectoral finrays 26. The species attains 7.5 cm in total length.

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Cyclopteropsis mcalpini

habitat: This species is benthic over mud generally at 174 m or more, with Canadian Arctic records at 109–329 m.

biology: Its biology is little known. About 60–70 eggs, around 5.0 mm in diameter, are laid in shells or protected areas and guarded by the male.

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, the mouth of Frobisher Bay, Hudson Strait, Ungava Bay, and off northern Labrador. It is known from Ulriks Bay (sic, thought to be the type locality) in northwest Greenland and from the Barents Sea. The Canadian records are from cruise data, and specimens have not been seen.

Distribution of Cyclopteropsis mcalpini

sources: See the family sources and the bibliography.

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Cyclopterus lumpus Linnaeus, 1758

Lumpfish, grosse poule de mer

common names: Local names are Lepisuk, Nipisa, and Qorkshuyoq; and Angusatdluk (male), Arnardluk (female), and Nipisa (Greenlandic). Other common names are Lump, Lump Sucker, and lompe. taxonomy: The genus comes from the Greek kyklos (circle) and pteron (fin). The species name comes from the English lump (to look disagreeable, ungainly). Cyclopterus lumpus var. hudsonius Cox, 1920, was described from Fort Churchill, Manitoba, in Hudson Bay but is regarded as a synonym of Cyclopterus lumpus. description: This species is distinguished from its relatives by

having a compressed body; the gill opening extending below the upper edge of the pectoral fin; tubercles in three distinct rows on the flank; and the first dorsal fin covered by a humped crest of skin in adults. Smaller tubercles may be absent, and prickles on the larger tubercles may be weakly developed, perhaps related to lower temperatures and salinities as in Hudson Bay (hence Cyclopterus lumpus var. hudsonius). The dorsal fin has 6–8 spines and 9–12 soft rays; the anal fin-rays number 9–11, and the pectoral fin-rays 17–24. There are 5–17 tubercles on the crest of the first dorsal fin, 2–7 mandibular tubercles, and 17–27 dorso-medial, 8–21 medial, and 3–8 ventral tubercles. The

overall colour is highly variable, grey, grey blue, blue green to yellow green, or chocolate brown. Some may be darkly blotched or dotted with black. The belly is yellowish to white. Spawning males have a grey-black or red-mauve body and a reddish belly, sides, and fins. Young may be bright green, brown, dark purple red, brown spotted, yellow with silvery dots and stripes, or red and often match the background. They have an eye stripe. The species attains 70.0 cm in length and 9.64 kg in weight.

habitat: This species is found from tidal pools and floating seaweed down to 1,272 m and is common in Canadian waters generally. It may rarely enter fresh water in southern Hudson Bay. Adults are found attached to rocks by their sucker or among seaweed but are also epibenthic and benthopelagic in Canadian waters at depths of 0–80 m over water as deep as 380 m and many kilometers from shore. Lumpfish are mostly seen as adults when breeding in shallow water or as young when living in tidal pools before dispersing. They are sluggish and easily picked up. In the sea their coloration is very close to their environment, and they are difficult to see. They are often solitary, found in deeper water in winter and entering shallower water in summer. They are more dominant in the western Hudson Strait than in the eastern part, and in Ungava Bay and Davis Strait. biology: Its food is crustaceans such as euphausiids and caprellids; comb jellies; jellyfish; arrow worms; polychaete worms; and small fishes such as Atlantic Herring and Sand Lances. They do not feed during the spawning season, and the male does not feed when guarding the eggs. Seals are predators on Lumpfish and will sometimes peel their skin off before eating them. They are also eaten by

Cyclopterus lumpus



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distribution: The species is found in Davis Strait, Cumberland Sound, Hudson Strait, and James, Hudson, and Ungava Bays; on all coasts of Greenland; and to Chesapeake Bay and Bermuda in the western Atlantic Ocean. It is found also in Europe.

Cyclopterus lumpus, disc

Sperm Whales, Killer Whales, and Greenland Sharks. In Hudson Bay they are eaten by Arctic Char. Their growth is rapid, to 5.3 cm, in the first year. Males are about four years old, and females five, when first spawning. Males are much smaller than females. This species lives to about 14 years of age. Spawning occurs inshore after a migration in spring to summer at about 8°C. However, spawning has been observed at temperatures as low as −1°C. A territory is established in shallow water by the male that arrives first on the spawning ground. Females mate indiscriminately with males, but there is a courtship involving quivering, nest cleaning, and fin brushing. Up to 300,000 sticky eggs, 2.2–3.1 mm in diameter, are laid in crevices or among seaweed. Each female may lay several egg masses at intervals of 8–10 days that hatch in 14–70 days depending on temperature. The male guards, fans, and puffs water over the eggs and is very aggressive. Fanning shortly after the eggs are laid disperses the waste ammonia that results from egg adhesion. Oxygen is not in short supply in turbulent shore waters. Spent females retire to deeper water. The eggs may take up to 70 days to hatch in cold northern waters. Puffing water at the egg mass increases in frequency near hatching and may help the larvae to break free. Egg masses are individually variable in colour, from black to brown, red, pink, orange, yellow, green, or purple. The young are pelagic.

importance: An inshore fishery off Newfoundland processed

about 200 t for caviar in 1972, but catches are often in the 2–4 t range for Canada. They are not really utilized in the Arctic, being eaten infrequently, but they have been used as dog food.

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Distribution of Cyclopterus lumpus

sources: Cox (1920); Cox & Anderson (1922); Vladykov (1933a);

Ryder et al. (1964); Davenport (1985); Morin & Dodson (1986); Crawford (1989); Jørgensen et al. (2005); Kudryavtseva & Karamushko (2005); Laidre, Heide-Jørgensen, & Orr (2006).

Eumicrotremus derjugini Popov, 1926

Leatherfin Lumpsucker, petite poule de mer arctique

common names: A local name is Læderfinnet Stenbider (Danish/Greenlandic).

taxonomy: The genus comes from Greek eu (very), mikros (small), and trema (hole). The species is named after the Russian ichthyologist Konstantin Mikhailovich Derjugin (1878–1938), who led the Russian Pacific expedition on RV Gagara. Eumicrotremus spinosus variabilis Jensen, 1944, described in part from Whale Sound, northwest Greenland (77°17' N, 69°59' W), is a synonym.

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Eumicrotremus derjugini

description: This species is separated from its relatives by having the gill opening above the pectoral fin; many large pointed tubercles, each with many spines, on the head, body, and caudal peduncle; supplemental pores above the lateral line; a rounded anterior body; the first dorsal fin-rays covered with skin and not visible; and the pectoral fin base without tubercles. The dorsal fin spines number 6–7, and the soft rays 11–13; the anal fin-rays number 10–13, and the pectoral fin-rays 25–27. The chin and throat are naked or have a few tubercles under the lower jaw in very large fish. The lateral-line pores are developed. There are no barbel-like chin flaps. The tubercles are low, conical, covered with prickles, and distant from each other by at least the diameter of their bases. The body is olive in colour on the flanks and darker on the back. The species reaches 12.7 cm in total length. habitat: The species is found on mud, gravel, or stone usually at

50–280 m, with young in shallow water (5 m in Arctic Canada), at temperatures in the range of −2.0°C (in Hudson Bay) to 4.93°C. It has been caught at Herschel Island at 150–152 m, in Hudson Strait at



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203–217 m, and in Baffin Bay, Davis Strait, and Hudson Strait at 105– 1,038 m. Salinity is usually at or near that of sea water. The juveniles are pelagic.

biology: Its food is hyperiid and gammarid amphipods, mysids,

oikopleura, copepods, caprellids, barnacle nauplii, and polychaete worms. Its food is predominately planktonic organisms, and, along with Arctic Cod, it is the only shallow-water Beaufort Sea species to feed so. This lumpfish is eaten by Thick-billed Murre chicks in Digges Sound, Hudson Bay, and by Atlantic Cod at Killiniq (Port Burwell) in Ungava Bay. Spawning is in late summer to fall. The eggs attain 4.3 mm, perhaps to 5.0 mm, in diameter and are deposited on the bottom.

importance: It is not economically important. distribution: The species is found in Nares Strait, Baffin Bay, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson Bay, Arctic islands, Dease Strait, Amundsen

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Gulf, and the Beaufort Sea. It is found also in the Alaskan Beaufort Sea (but not apparently the Chukchi Sea), the northern Sea of Okhotsk, northwest and northeast Greenland, Labrador, and across northern Eurasia.

tubercles, each with many spines, on the head, the body, and the caudal peduncle; supplemental pores above the lateral line; a rounded anterior body; the first dorsal fin spines not covered by a thickened layer of skin; the pectoral fin base with tubercles in females but not in all males; and the caudal area tubercles larger dorsally. The dorsal fin spines number 5–10, and the soft rays 10–13; the anal fin-rays number 9–13, and the pectoral fin-rays 20–27. The lateral-line pores are developed and number 13–14. The tubercles are large and rough, and their number is thought to increase with age. There are one to four rows of tubercles under the lower jaw in females. Tubercles are found along the base of the first dorsal fin in females, but are absent in most males. Females have an average of 94 total tubercles (range 56–138), while males have 26 total tubercles (range 15–42), the latter based on fish initially identified as E. eggvinii. The colour in young is olive or red, and in adults is darker, usually brown, with some stripes or bands posteriorly and with brown fin-rays. The belly is whitish. The tubercles are sometimes fringed with a dark ring. Some fish may be quite pale in colour and have smooth tubercles. The species reaches 13.7 cm in total length.

habitat: This species lives on stone, among algae (the bladders

Distribution of Eumicrotremus derjugini

sources: Gaston et al. (1981); Gaston et al. (1985); Ponomarenko (1995).

Eumicrotremus spinosus (Fabricius, 1776)

Atlantic Spiny Lumpsucker, petite poule de mer atlantique

common names: Local names are Man-iktoe (Inuktitut) and Nipisarluk (Greenlandic). Another common name is Pimpled Lumpsucker. taxonomy: The species name is the Latin spinosus (spinous).

The subspecies Eumicrotremus spinosus variabilis Jensen, 1944, was described from northwest Greenland near Canadian waters but is regarded as a synonym of Eumicrotremus derjugini (see above). Eumicrotremus eggvinii Koefoed, 1956, described from off the coast of Jan Mayen Island but sometimes recorded from Canadian and Greenland waters, is a synonym based on a male E. spinosus

description: This species is separated from its relatives by having the gill opening above the pectoral fin; many large pointed

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of which it may resemble), and on gravel, sand, or mud bottoms, often at 5–100 m but down to 930 m, with young in shallower waters. It was caught in Hudson Strait at 0–483 m for 145 collections, in Ungava Bay at 158–457 m for 62 collections, and in Davis Strait and Baffin Bay at 164–930 m. Its preferred temperature range is −1.4°C to 5.4°C. It is found in low-salinity waters in Richmond Gulf.

biology: Its food comprises crustaceans such as amphipods,

copepods, and mysids; oikopleura; and fish. This lumpfish is eaten by seals; by Atlantic Cod at Killiniq (Port Burwell), Ungava; and by Thick-billed Murre chicks in Digges Sound, Hudson Bay. It is eaten by Brünnich’s Murres and by Thick-billed Murre chicks (0.89% of diet) at Akpatok Island, Ungava Bay. Arctic Char take this species in Frobisher Bay and at Victoria Island, halibut take it in Davis Strait, and Northern Wolffish in Mould Bay, Prince Patrick Island. Spawning occurs in March and/or early autumn at 20–30 m. The eggs are orange or golden yellow in colour, 4.5 mm in diameter, and deposited on the bottom.

importance: It is not economically important. distribution: The species is found in Nares Strait, Baffin Bay, Davis Strait, the mouth of Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson Bay (sparsely), Foxe Basin, Gulf of Boothia, the Arctic islands, Dease Strait, Amundsen Gulf, and the Beaufort Sea, but not in the Alaskan Beaufort Sea. General reports for James Bay are not confirmed by specimens. It is found also in the North Atlantic Ocean from all coasts of Greenland south to Cape Cod in the west, and in northern Europe.

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Eumicrotremus spinosus

sources: Dresel (1885); Johansen (1927a); Grainger (1953); MacDonald (1954); Tuck & Squires (1955); Tuck (1960); Gaston et al. (1981); Gaston et al. (1985); Jørgensen et al. (2005); Byrkjedal, Rees, & Willassen (2007).

Distribution of Eumicrotremus spinosus



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Family Liparidae Snailfishes, Limaces de mer

Brian W. Coad

at Akpatok Island, Ungava Bay (1.66% of diet). Snailfishes are probably eaten by Bearded Seals at Aujuittuq (Grise Fiord) and possibly at Mittimatalik (Pond Inlet) and Kangiqtugaapik (Clyde River) on Baffin Island. Children in Richmond Gulf play a game where they place a Sea Tadpole on their arm, to which it attaches by the sucker, and are chased by others who are trying to pull it away. It may be caught through the ice in winter when one is fishing for cod.

sources: Burke (1930); Vogt (1977); Able & McAllister (1980);

Able & Irion (1985); Able, Fahay, & Markle (1986); Chernova (1988, 2005a, 2005b, 2008); Able (1990); Andriashev (1991); Vogt & Chernova (2002); Chernova, Stein, & Andriashev (2004); Andriashev & Chernova (2010).

Snailfishes or Sea Snails are known mostly from northern cold and temperate waters and from the Antarctic. They may be worldwide in deep, cold waters. There are about 334 species, including 51 reported from Canada, of which 11 are in the Arctic. The maximum size is about 80 cm although some species are only one-tenth of this length. Snailfishes are easily recognized by their loose skin with jelly-like tissue underneath. This may aid in flotation for those species without a gas bladder. The skin lacks scales and tubercles but may bear prickles in some species. The lateral line is absent. The body is often tadpole shaped. The pelvic fins are modified into a thoracic sucking or adhesive disc in many species. A few, often pelagic, species lack pelvic fins. The dorsal and anal fins are long and often merge with the caudal fin. Snailfishes are difficult to identify, and characters include pore counts and patterns, fin-ray numbers, the gill slit position and size, teeth shape and rows, peritoneum colour, body colour, disc form, and several others. Snailfishes are found in tide pools and offshore in moderately deep waters. Some are found at the incredible depth of 7,588 m, an area known as the hadal zone. Some deep-sea species are pelagic. In certain parts of the world they make up a major part of the ichthyofauna, as much as 85.2% in bays in the western Bering Sea, for example. Their food is usually benthic invertebrates although some deepsea species feed on nekton. The eggs are often few and large, as few as six per female and up to 8.0 mm in diameter. They may be laid in the gill cavities of crabs and scallops. The adhesive sucking disc is used to cling to rocks or algae such as Laminaria fronds in Resolute Bay, Cornwallis Island. In the Canadian Arctic their food is crustaceans and other invertebrates and rarely other fishes. Their eggs are adhesive and sometimes laid on or in other animals. Some species guard their eggs. Liparis species taken in shallow water are a food item for Black Guillemot chicks at Prince Leopold Island, off Somerset Island; for adult Thickbilled Murres at Digges Sound, Hudson Bay; and for murre chicks

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Careproctus kidoi

Knudsen and Møller, 2008 Kido’s Snailfish, limace de Kido

common names: A local name is Kidos Ringbug (Danish/

Greenlandic).

taxonomy: The genus comes from the Greek kara (head) and proktos (anus). The species is named after the liparid ichthyologist Kaoru Kido, Hokkaido University, Japan, who first recognized that this could be a new species. description: This species is distinguished from its relatives by

having a small adhesive disc (similar to eye diameter); a short lower pectoral fin lobe (shorter than head length); a slender body (depth usually less than head length); usually fewer pectoral fin-rays (26 or less, rather than 25 or more); and generally a brown to black overall colour internally and externally. The dorsal fin-rays number 54–60; the anal fin-rays number 50–54; and the total pectoral fin-rays number 21–26, the upper lobe pectoral fin-rays being 11–14 and lower lobe rays 5–7, with 3–8 in the notch between the upper and lower lobes. The teeth are conical, pointed, and simple. The pyloric caeca number 3–12 on the left side of the abdominal cavity. The overall colour is light to dark brown with a darker black band on the trunk and posteriorly on the body. Freshly caught fish are covered by a gelatinous layer that is lost after preservation. The eyes are blue. The skin is transparent. The stomach is dusky to dark brown, and the peritoneum and gill cavity are black. The species attains 9.5 cm in standard length.

habitat: This species is caught at depths of 952 m to 1,487 m and

temperatures of −0.09°C to 0.50°C, apparently replacing C. reinhardti below about 700–800 m. Non-closing trawls were used to capture this species, so the vertical distribution is uncertain.

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Careproctus kidoi

biology: A mature female measuring 89 mm in standard length

had 7 large eggs of up to 7.2 mm in diameter, along with 20 smaller eggs.

importance: It is not economically important. distribution: The species is found in Davis Strait, northwest and southwest Greenland, and off northern Labrador.

Careproctus longipinnis Burke, 1912

Longfin Snailfish, limace à longues nageoires

common names: None. taxonomy: The species name comes from the Latin longus (long) and pinna (fin). This species may be a synonym of Careproctus reinhardti. description: This snailfish is separated from its relatives by having a small adhesive disc (slightly smaller than eye diameter); a disc that is round rather than pear shaped as in C. reinhardti; one pair of nostrils (i.e., a single nostril on each side of the head, a nearby pore not to be confused with a nostril); lower pectoral fin lobe rays longer than head; the pectoral fin not inserted low (as opposed to uppermost ray level being opposite the oblique mouth in C. reinhardti); the gill slit short, reaching to the fourth pectoral fin-ray (reaching to lower pectoral lobe in C. reinhardti); a deep body, equal to or greater than head length; a horizontal rather than an oblique mouth; lack of silvery belly pigmentation; and a high anal fin-ray count. The dorsal fin-rays number 51–55, the anal fin-rays 44–49, and the pectoral fin-rays 30–32. The gill slit is above the pectoral fin. The overall colour is reddish, yellowish, or white. The peritoneum and mouth cavity are pale. The species reaches 27.0 cm in total length. habitat: It is found on sand or mud bottoms at 340–1,322 m. biology: The food of this epibenthic species includes amphipods

Distribution of Careproctus kidoi

and other crustaceans. A female of 19.8 cm standard length had unripe eggs about 1 mm in diameter.

source: Knudsen & Møller (2008).



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Careproctus longipinnis

importance: It is not economically important.

Careproctus reinhardti

distribution: The species has been found as two records at Reso-

Sea Tadpole, petite limace de mer

lution Island in the eastern Hudson Strait (Hudon, 1990a, 1999), and south to Nova Scotia. It is found also in the northeastern Atlantic Ocean. The Canadian Arctic literature records need confirmation by specimens (and note also that there is possible confusion or synonymy with the following species).

(Krøyer, 1862)

common names: A local name is Reinhardts Ringbug (Danish/

Greenlandic). Other common names are Jellyfish, Longfin Seasnail, Longfin Snailfish, and limace à longues nageoires.

taxonomy: The species is named after the zoologist Johannes Christofer Hagemann Reinhardt (1776–1845), the first professor of zoology at the University of Copenhagen, who described new species from Greenland. This name has been widely applied and may well conceal more than one species, yet to be determined. description: This snailfish is separated from its relatives by having a small (slightly smaller than eye diameter) deep-cupped and pear-shaped adhesive disc; one pair of nostrils (i.e., a single nostril on each side of the head, a nearby pore not to be confused with a nostril); slender body (depth generally less than head length); a high anal fin-ray count; and 16–30 pyloric caeca. Other characters are listed under C. longipinnis. It is named Jellyfish for the thick gelatinous sheath encasing most of the body except the pectoral and caudal fins. The dorsal fin-rays number 50–58, the anal fin-rays 41–58, and the pectoral fin-rays 25–34. The overall colour is reddish, pinkish, or whitish, most colour being that of the visible internal organs. The stomach area appears iridescent. The peritoneum is pale. The species reaches 30.0 cm in total length. Distribution of Careproctus longipinnis

sources: Hudon (1990a, 1999).

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habitat: This species occurs epibenthically at 48–384 m in the

Gulf of St Lawrence and down to 1,284 m over mud bottoms, perhaps to 1,840 m or more, elsewhere. In Hudson Strait it is caught at

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Careproctus reinhardti

Careproctus reinhardti



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260–360 m and in Davis Strait and southern Baffin Bay at 151–1,482 m and 0.4°C–3.0°C. Deeper records in Davis Strait and Baffin Bay may have been C. kidoi. It is found in temperatures near or below 0°C in Richmond Gulf and at −1.7°C to 4.0°C elsewhere. Salinity is at or near that of sea water.

biology: Its food includes small pelagic crustaceans such as

hyperiids; benthic crustaceans; and rarely small fishes. This species is eaten by Roughhead Grenadiers in the Davis Strait. Females are mature at sizes larger than 12 cm. The maximum age in Europe is seven years. Spawning probably occurs in spring and summer in the Gulf of St Lawrence. Eggs are up to 4.5 mm in diameter and number about 300 in each female. Eggs and larvae are probably demersal, and juveniles pelagic.

importance: It is not economically important. distribution: The species is found from northern Baffin Island, Baffin Bay, Davis Strait, Hudson Strait, Ungava Bay, eastern Hudson Bay (rarely), and all coasts of Greenland south to the Gulf of St Lawrence. General reports for James Bay are not confirmed by specimens. It is found also in the western Arctic in Bathurst Inlet and Amundsen Gulf (Arctic Laboratories Ltd. & LGL Limited, 1987) and the Beaufort Sea (ROM 51611). There are no records for Alaska. It is found elsewhere in the Arctic waters of the northern hemisphere.

Liparis atlanticus

(Jordan and Evermann, 1898) Atlantic Seasnail, limace atlantique

common names: Another common name is Atlantic Snailfish. taxonomy: The genus comes from the Greek liparos (sleekskinned, shiny with oil). The species name is the Latin atlanticus (of the Atlantic).

description: This snailfish is distinguished from its relatives by

having a large adhesive disc; two pairs of nostrils (two nostrils on each side of the head); a dorsal fin with an anterior notch in males (at the fifth or sixth ray) and 30–35 rays; and anal fin-rays numbering 25–29, pectoral fin-rays 25–31, pyloric caeca 19–45, and total vertebrae 38–42. The lower pectoral lobe has finger-like rays. The teeth are trilobed and in bands. Males have thumb-tack-shaped prickles on the body and elongated dorsal fin-rays. The colour varies with the habitat, from black, grey, and olive to reddish brown. The fins are often barred with white, blue, or pink. There may be light bars on the flank. The caudal fin is faintly barred. The peritoneum is pale with faint brown dots. The species reaches 14.4 cm in total length.

habitat: This snailfish is common in weedy tide pools, attached

tenaciously by the disc to the underside of rocks or to kelp, but it may descend to 250–415 m in Ungava Bay. It may be found over hard sand, clay, or sand-mud bottoms. Its favoured temperatures are 7.0°C–14.6°C; the 2°C August surface-temperature isotherm is the northern limit, and 20°C the southern limit. Adults move into shallow water in October, retreating to deeper water only in very cold weather.

biology: Its food is taken mostly in the early morning and even-

ing and is principally amphipods and polychaete worms. Winter Skates and Sea Ravens are known to eat this species. Males tend to be larger than females. Most fish are mature at 6–7 cm and two years of age. Males clean a nest site by rubbing it with their snout and driving away conspecifics. Spawning occurs from March to June in Atlantic Canada, judging by females with large eggs. A large female has about 700 mature eggs that are laid in small egg masses and then gathered into a single, large, attached mass by the male. Egg masses may be attached to algal stems by the male. The male fans and guards the egg mass. Fertilized eggs are up to 1.4 mm in diameter. Larvae are planktonic but may be more abundant near the bottom.

importance: It is not economically important. Distribution of Careproctus reinhardti

sources: Edwards (1961); Finley & Evans (1983); Arctic Labora-

tories Ltd. & LGL Limited (1987); Hudon (1990a); Nielsen et al. (1992); Jørgensen (1996); Treble et al. (2000).

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distribution: The species is found from Ungava Bay along the Labrador coast to the estuary and Gulf of St Lawrence, to Newfoundland, and south to the Bay of Fundy and on to New Jersey. A record from near Qausuittuq (Resolute), Cornwallis Island, is not confirmed by material.

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Liparis atlanticus

Distribution of Liparis atlanticus

sources: See the family sources and the bibliography.



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Liparis fabricii Krøyer, 1847

Gelatinous Seasnail, limace gélatineuse

common names: A local name is Fabricius Ringbug (Danish/ Greenlandic). Another common name is Gelatinous Snailfish.

taxonomy: The species is named after Otto Fabricius (1744–1822),

a Danish naturalist and missionary who studied the fishes of Greenland and wrote Fauna Groenlandica. Ophidium parrii Ross, 1826, described from Baffin Bay, between west Greenland and Baffin Island, and Prince Regent’s Inlet, off northern Canada, is a senior synonym. The name has been ruled invalid and placed on the Official Index of Rejected and Invalid Names in Zoology. Liparis koefoedi Parr, 1932, described from Spitsbergen, is a synonym, but the name has often been used in the Canadian Arctic and is sometimes recognized as distinct. Further work is needed to reconfirm this distinction. Records of Liparis major Gill, 1864, described from Arctic Seas, are also probably this species.

description: This snailfish is distinguished from its relatives by

having a large adhesive disc; two pairs of nostrils (one pair on each side of the head); dorsal fin without a notch; dorsal fin-rays numbering 41–54; anal fin-rays numbering 35–42 (usually 37–40); pectoral fin-rays numbering 32–40; a dark peritoneum; and posterior teeth that are usually simple with only anterior teeth obviously trilobed. The lower pectoral fin lobe rays are finger-like. The gill slit extends down to the pectoral fin-rays, which are 5–11 usually, perhaps to 15. The pyloric caeca number 18–34, and total vertebrae 48–53. Males develop skin prickles. The colour varies, becoming darker shades of brown or black with age. The side of the head and the mid-flank may

bear pink tinges. The belly is silvery in young. The dorsal, anal, and pectoral fins are dark brown to black. Young fish are semi-transparent with 6–8 dark bands on the posterior part of the body, extending onto the dorsal and anal fins. The caudal fin has 2–3 bars. The mouth and gill cavities, and the stomach, are dark. The peritoneum has spots in young that become so dense in adults that the peritoneum is black. The species attains 19.4 cm in total length.

habitat: This snailfish is found usually on mud, mud and sand,

or shell and detritus bottoms, and sometimes pelagically, as shallow as 6 m and as deep as 1,880 m. Very deep collections may be fish trapped in shallower depths as deep-set nets are hauled to the surface. In Arctic Canada it is found at temperatures of −1.80°C to 0.56°C and rarely as high as 11.00°C, and elsewhere down to −1.91°C. In Davis Strait and southern Baffin Bay it was found at 422 m to 1,487 m and −0.08°C to 3.80°C for 62 collections, but other collections there have taken it as shallow as 100 m. In Hudson Strait it is found at 146–409 m based on 42 collections. Off southeastern Baffin Island, young of the year were the most abundant species caught in surface and mid-water trawls outside protected bays, along with tentatively identified Kelp Snailfish. At 40–50 mm in length they descend to the bottom. In NAFO Subarea 0A it dominated in tows at depths greater than 600 m, declining as catches of Gaidropsarus ensis increased. At Kuujjuarapik in southeastern Hudson Bay, larvae are found at temperatures up to 10.8°C. In Resolute Bay, Cornwallis Island, this species is associated with seaweeds (Laminaria and Agarum) or is taken from the water column, but never more than 1 m above the bottom or away from seaweeds. This species has a biological anti-freeze that lowers the freezing point of body fluids below that of sea water (−1.9°C). It has been reported as swimming among ice and being immobile in ice crevices, apparently hiding from predators. It may be caught under pack ice over depths of 2 km, so the deeper records may be erroneous, resulting from midwater pelagic captures.

Careproctus kidoi Liparis fabricii

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Liparis fabricii

biology: Its food is crustaceans such as pelagic hyperiid amphipods, mysids, euphausiids, gammarids, and gastropods; polychaete worms; and larvaceans. Pelagic fish feed on planktonic copepods and ostracods. The main diet items in Resolute Bay, Cornwallis Island, were copepods and amphipods. Calanoid copepods dominate off southeastern Baffin Island. Larval and post-larval fish feed on copepods such as Acartia clausi, Calanus glacialis, C. hyperboreus, Cyclops sp., Euchaeta glacialis, Eurytemora herdmani, E. raboti, Jaschnovia tolli, Limnocalanus macrurus, Metridia longa, Microcalanus pygmaeus, Microsetella norvegica, Oithona similis, Oncaea borealis, and Pseudocalanus minutus; amphipods such as Acanthostepheia behringiensis, Apherusa glacialis, Onesimus glacialis, and Parathemistio abyssorum; mysids such as Mysis litoralis; pteropods such as Limacina helicina; larvaceans such as Fritillaria borealis and Oikopleura vanhoeffeni; diatoms such as Coscinodiscus sp.; and, in the Beaufort Sea, decapods, hyperiids, and podocop ostracods. It is eaten by Atlantic Cod at Killiniq (Port Burwell), Ungava; by Arctic Terns at Alert; and by Thick-billed Murres at the Nuvuk Islands, Digges Sound, Hudson Bay; it may be eaten by Arctic Char at Wakeham Bay, Ungava. It is eaten by Brünnich’s Murres at Akpatok Island, Ungava Bay, and is fed to chicks by Thick-billed Murres



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at Coats Island, Nunavut, and at Prince Leopold Island, off Somerset Island. This snailfish is also eaten by seals and Shorthorn Sculpins. Spawning in Russian Arctic waters may be in September–October, with egg diameter reaching 2.7 mm and egg numbers 735. Young, at 12–31 mm in length, have been collected in late July and August near Baffin Island and Labrador, and at 15–25 mm from Brevoort Island, southeast of Baffin Island, in early August, suggesting an earlier spawning. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae occur in May, suggesting a winter spawning.

importance: It is not economically important. distribution: The species is found at the northern tip of Ellesmere Island (Alert), in Nares Strait, Baffin Bay, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, Hudson Bay (rarely), Foxe Basin, Lancaster Sound, Gulf of Boothia, the Arctic islands, Dease Strait, Bathurst Inlet, Amundsen Gulf, and the Beaufort Sea, including the Alaskan Beaufort Sea. It has been reported under ice at 81°10' N, 158°00' E. Populations in Siberia and Alaska are not as frequently encountered as in Canada. It is found also on all coasts of Greenland, off Labrador, on the Grand Banks, and in the Gulf of St Lawrence.

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common names: A local name is Pukkelrygget Ringbug (Danish/Greenlandic). Other common names are Dusky Snailfish and Polka-dot Seasnail. taxonomy: The species name is the Latin gibbus (hump).

Liparis cyclostigma Gilbert, 1895, described from Bristol Bay, Alaska, is a synonym but has been used in the Canadian Arctic and is sometimes recognized as distinct. Liparis liparis bathyarcticus Parr, 1931, formerly considered a synonym of L. gibbus, is now recognized as a distinct species. There are no confirmed records from Arctic Canada for this circumpolar species although it is known from Labrador, west Greenland, and the Alaskan Bering Sea.

description: This snailfish is distinguished from its relatives

Distribution of Liparis fabricii

sources: Tuck & Squires (1955); Andriashev & McAllister (1978);

den Beste & McCart (1978); Tsinovsky & Mel’nikov (1980); Gaston & Nettleship (1981); Astaf ’yeva, Antonov, & Petrov (1983); Green (1983); Gaston (1987); Gaston et al. (1987); Chernova (1988); Nielsen et al. (1992); Ponton et al. (1993); Woehrmann (1997); Wöhrmann (1997); Chambers & Dick (2007); Chernova (2008).

Liparis gibbus Bean, 1881

Variegated Snailfish, limace marbrée

by having a large adhesive disc; two pairs of nostrils (one pair on each side of the head); the dorsal fin without a notch; dorsal fin-rays numbering 37–46 (usually 41–44); anal fin-rays numbering 31–38 (usually 33–36); pectoral fin-rays numbering 35–45, usually 38–42, with gill opening opposite rays 8–16; the peritoneum being light in colour; posterior teeth with a central lobe that is longer than the lateral lobes; and anterior teeth with three equal lobes. The pyloric caeca number 15–45, and the total vertebrae 45–49. Males have prickles. The overall colour varies and may be uniform reddish brown to reddish pink either with yellowish tints or alternately striped light and dark on the head and body. Some fish are finely spotted. The fins have black margins and may have bands and patches. The caudal fin may have two to three bands. The peritoneum is generally pale with a few pigment spots. The species reaches 52.4 cm in total length and 1.027 kg in weight.

habitat: The species is found on rock, sand, and mud bottoms and

reportedly in empty scallop shells. It is usually deeper than 100 m but can be as shallow as 25 m, and is found at 27 m and as deep as 981 m in the eastern Canadian Arctic. In Hudson Strait it is caught at 260–362 m, and in Ungava Bay at 158–415 m. Variegated Snailfish are found offshore and possibly pelagically. In Europe, juvenile fish up to 4.3 cm are pelagic at 35–200 m. Its preferred temperatures are as low as −1.85°C.

biology: Its food is benthic and pelagic crustaceans including

amphipods, mysids, hyperiids, caprellids, copepods, cumaceans, isopods, and crabs and prawns; as well as gastropods, nematodes, polychaete worms, and fish. Some food associated with plants is taken above the bottom. This snailfish is eaten by Greenland Halibut and Atlantic Cod at Killiniq in Ungava Bay. The maximum age in Europe is about six years. Reproduction may occur in spring and summer in the St Lawrence River estuary but from July to September or perhaps during winter in the Arctic.

importance: It is not economically important.

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distribution: The species is found near Aujuittuq (Grise Fiord), in Baffin Bay, Davis Strait, Frobisher Bay, Hudson Strait, Ungava Bay, sparsely in Hudson and James Bays, and rarely in the Gulf of

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Liparis gibbus

Boothia and the Arctic islands including Resolute Bay, Cornwallis Island, Melville Sound, Amundsen Gulf, and the Beaufort Sea. It is found also in the Alaskan Beaufort Sea, Bering and Chukchi Seas, northwest, southwest, and northeast Greenland, and south to northern Nova Scotia in the Atlantic, and to British Columbia and Kamchatka in the Pacific. It occurs also in Arctic Eurasia.

sources: Gillis & Allard (1984); Falk-Petersen, Frivoll, Gulliksen, Haug, & Vader (1988); D.B. Stewart et al. (1991); Atkinson & Percy (1992); Chernova (2008).

Distribution of Liparis gibbus



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Liparis tunicatus Reinhardt, 1836

Kelp Snailfish, limace des laminaires

common names: Local names are Nee-fitz-shak, Nipi-sak, and

Nipishah (Inuktitut); and Grønlandsk Ringbug (Danish/Greenlandic). Other common names are Bartail Snailfish, Greenland Snailfish, and limace de Groenland.

taxonomy: The species name is the Latin tunicatus (bearing a cloak). Liparis herschelinus Scofield in Jordan and Evermann, 1898, described from Herschel Island, Arctic Ocean, is a synonym although it is sometimes recognized as distinct. The distinction is unclear, and these fishes require more detailed study. Liparis arctica Gill, 1864, from Greenland, is also a synonym. description: This snailfish is distinguished from its relatives by having a large adhesive disc; two pairs of nostrils (one pair on each side of the head); the dorsal fin without a notch; posterior teeth with three about-equal lobes; a light-coloured peritoneum; dorsal

Liparis tunicatus

Liparis tunicatus

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fin-rays numbering 36–45 (usually 41–43); anal fin-rays numbering 27–38 (usually 34–36); pectoral fin-rays numbering 31–40, usually 32–37, with a gill opening above this fin or opposite rays 0–7 (usually 4–6); pyloric caeca numbering 10–50; and total vertebrae numbering 44–50. Males have prickles on the tip of the head from the nostrils back to the anterior dorsal fin. The overall colour varies with habitat and is usually grey brown or reddish brown to black. Some may be lightly striped or speckled and mottled on the flank. The dorsal and anal fins may be orange reddish with 6–8 bars. The caudal fin has 1–3 bars. The peritoneum is pale with numerous small dots dorsally. The species reaches about 20.0 cm in total length.

importance: It is not economically important. distribution: The species is found at the northern tip of Elles-

mere Island, Nares Strait, northern Baffin Bay, with a large gap before southern Davis Strait, Frobisher Bay, Hudson Strait, Ungava Bay, northern Hudson Bay, and, after a gap, southern James Bay, Foxe Basin, Gulf of Boothia, through the Arctic islands, Melville Sound, Dolphin and Union Strait, Amundsen Gulf, Prince of Wales Strait, and the Beaufort Sea. It is found also in the Alaskan Beaufort Sea, Chukchi Sea, northern Bering Sea, northwest, southwest, and southeast Greenland, and the Labrador coast. General reports for Cumberland Sound are not confirmed by specimens.

habitat: The species is usually found attached to kelp fronds by

the adhesive disc and curled up in shallow shore waters including tidal pools, and even under sea ice. Its bottom types are pebbles, stones or boulders, mud, or sand. In the Alaskan Beaufort Sea, however, they are one of the commonest pelagic species after Arctic Cod. Juveniles there are associated with jellyfish medusa, which may afford some protection from predators. In Resolute Bay, Cornwallis Island, this species is associated with seaweeds (Laminaria and Agarum) or caught from the water column but never more than 1 m above the bottom or away from seaweeds. Young of the year are found in the water column, and even adults will swim up into surface waters at night in Ungava Bay. They have been collected from under stones in Ungava Bay. They are rarely collected below 50 m but can be as deep as 620 m. Fifteen collections from Ungava Bay had a depth range of 210–620 m, while nine collections in Hudson Strait were at 311–388 m. Its preferred temperatures vary between −1.8°C and 4.1°C, and the southern limit agrees with the 6°C August surface isotherm.

biology: Its food is principally amphipods, mysids, cumaceans,

and copepods, with some other crustaceans in Jones Sound. At Cape Hatt, northern Baffin Island, the pelagic pteropod Limacina helicina is the major part of the prey biomass at 46%. The main diet items in Resolute Bay, Cornwallis Island, are copepods and amphipods. Open-water fish eat a common littoral amphipod (Ischyrocerus anguipes) and copepods. Fish at the ice edge do not take this amphipod, although it is present, but take epibenthic cumaceans, copepods, mysids, and lysianassid amphipods. Fish smaller than 100 mm in total length eat primarily cyclopoid and harpacticoid copepods, while larger fish eat amphipods, mainly Onesimus glacialis, with a more varied diet including polychaetes, squid, and decapods in Wellington Channel and Resolute Passage. Larval and post-larval fish feed on copepods such as Calanus glacialis, Jaschnovia tolli, Metridia longa, and Pseudocalanus minutus; amphipods such as Apherusa glacialis and Gammarus sp.; and cumaceans in the Beaufort Sea. It is eaten by Brünnich’s Murres at Akpatok Island, Ungava Bay, and by cod in Ungava Bay generally. This species is eaten by seals in the Canadian Arctic, probably at Cumberland Sound, and by Arctic Char generally. Spawning probably occurs in winter and spring.



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Distribution of Liparis tunicatus

sources: Hildebrand (1948); Dunbar & Hildebrand (1952); Tuck

& Squires (1955); Bain, Thomson, Foy, & Griffiths (1977); Green (1983); Byers & Prach (1988); Thorsteinson et al. (1991); Nielsen et al. (1992); Chernova (2008).

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Paraliparis bathybius (Collett, 1879)

Black Seasnail, limace noire

common names: A local name is Arktisk Dybhavsringbug (Danish/Greenlandic).

is potential for overlap with related species). The lower pectoral fin lobe is made up of 3–4 separate rays. The pectoral fin is above the level of the eye centre. The gill opening is above the level of the pectoral fin. The body is black with a violet tinge. The gill, mouth, and peritoneum are black, but the stomach and pyloric caeca are pale. The tail is dusky to black. The species reaches 25.3 cm in standard length.

habitat: This snailfish is bathydemersal, found in cold waters on

aros (sleek-skinned, shiny with oil). The species name comes from the Greek bathys (deep) and bios (life). It was originally spelled bathybii, but most authors give the spelling above.

or above the bottom at 600–4,009 m, usually below 1,000 m, perhaps as shallow as 20 m. In the northern Davis Strait it has been recorded at 1,173–1,600 m and −0.08°C to 0.38°C, and in southern Baffin Bay and Davis Strait generally at 545–1,483.5 m and −0.1°C to 0.9ºC. It may be found in groups.

description: This snailfish is separated from its relatives by the

biology: Its food includes pelagic amphipods, benthic snails, and

taxonomy: The genus comes from the Greek para (near) and lip-

lack of an adhesive disc and therefore of pelvic fins and by having one pair of nostrils, upper-lobe pectoral fin-rays numbering about 18–20, and a black body. The mouth is horizontal, and well-developed teeth are in broad bands. The dorsal fin-rays number 56–66, the anal fin-rays 48–55, and the pectoral fin-rays about 18–20 (as few specimens are available, there

crustaceans such as mysids. Spawning occurs in the summer and possibly at other times. Females have up to 434 bright-yellow eggs that are up to 5.0 mm in diameter and account for about 27% of body weight.

importance: It is not economically important.

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distribution: The species is found in Baffin Bay and Davis Strait, northwest and northeast Greenland, and in the North Atlantic Ocean including northern Europe.

Paraliparis copei

Goode and Bean, 1896 Blacksnout Snailfish, limace à museau noir

Paraliparis copei

common names: A local name is Copes Dybhavsringbug (Danish/Greenlandic). Another common name is Blacksnout Seasnail. taxonomy: The species is named after Edward Drinker Cope, an American ichthyologist, from whom Copeia, the journal of the American Society of Ichthyologists and Herpetologists, derives its name. description: This snailfish is separated from its relatives by the

Distribution of Paraliparis bathybius

sources: Jensen (1950a); Tsinovsky (1980); Jørgensen et al. (2005).

lack of an adhesive disc and therefore of pelvic fins and by having one pair of nostrils, weak teeth in a single row, and the lower snout and mouth brown to black in colour, strongly contrasting with surrounding tissues. The dorsal fin-rays number 59–68, the anal fin-rays 53–60, and the upper pectoral fin lobe rays 13–17 (total 20–22). There are 8–9 caudal fin-rays and 6 pyloric caeca. The body mostly lacks pigment and is a translucent or milky white. The abdomen, gill cavity, chin, lower snout, and lips are all black. The peritoneum is dark, and the stomach and pyloric caeca pale. The species reaches 23.2 cm in standard length.

habitat: This species occurs at 210–500 m in the Gulf of St Lawrence within 2 m of the bottom, and down to 1,976 m elsewhere. It has been caught in Davis Strait at 216–1,078 m, at 988–1,463 m and 3.28°C, and at 1,287.5–1,451.0 m and 0.20ºC–3.50ºC.

Paraliparis copei



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biology: Its food is apparently pelagic comb jellies, cnidarians,

and possibly salps. Spawning may occur in spring and summer in Canadian Atlantic waters, but in the United States mature males and females are caught almost year round. The eggs are up to 3.5 mm in diameter and number up to 88.

Paraliparis garmani

importance: It is not economically important.

common names: A local name is Garmans Dybhavsringbug

distribution: The species is found in Davis Strait off south-

ern Baffin Island, eastern Hudson Strait and Ungava Bay, from southwest and southeast Greenland south to Cape Hatteras, and throughout Atlantic Canada. It is found also at the Azores and off South Africa.

Burke, 1912

Pouty Snailfish, limace pote

(Danish/Greenlandic).

taxonomy: The species is presumably named after the American Samuel Walton Garman (1843–1927), the first curator of ichthyology at the Museum of Comparative Zoology, University of Harvard, who studied liparids. description: This snailfish is separated from its relatives by the

lack of an adhesive disc and therefore of pelvic fins, by having one pair of nostrils, teeth in bands, an oblique mouth, eight caudal finrays, and the snout and mouth light in colour, similar to that of the surrounding tissues. The dorsal fin-rays number 54–60, the anal fin-rays 49–53, and the upper pectoral fin lobe rays 13–14 (total 18–22). There are 6 pyloric caeca. The body is a dusky white to a light pale brown with scattered black pigment spots. The peritoneum is black, and the stomach and pyloric caeca pale. The species reaches 14.2 cm in standard length.

habitat: This epibenthic species has been caught in the depth

range of 226–987 m, and at 546–987 m and 3.7ºC–4.6ºC in Davis Strait. They have been observed from submersibles, swimming 1–3 m above the sea floor and angled upward at 15°–30° to the horizontal. Their swimming is by rhythmic fin movements.

biology: Its food is pelagic crustaceans, but amphipods that are

Distribution of Paraliparis copei

scavenging dead fish lying on the sea bottom may also be eaten. Females with up to 317 eggs greater than 2 mm in diameter have been caught in January and September off the U.S. coast. The maximum egg size is 3.5 mm in diameter.

sources: See the family sources and the bibliography.

importance: It is not economically important.

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distribution: The species is found in Davis Strait off southern Baffin Island, the mouth of Frobisher Bay, eastern Hudson Strait, and southwest and southeast Greenland, south to Cape Hatteras, and is assumed to occur along Canada’s Atlantic coast.

Rhodichthys regina Collett, 1879

Threadfin Snailfish, limace à filaments

common names: A local name is Rød Dybhavsringbug (Danish/ Greenlandic). Another common name is Threadfin Seasnail. taxonomy: The genus comes from the Greek rhodeos (rose) and ichthys (fish). The species name is the Latin regina (queen). It may be placed in the genus Paraliparis by some authors.

Distribution of Paraliparis garmani

description: This species is separated from other Arctic Snailfishes by the lack of an adhesive disc and pelvic fins, and the lower pectoral fin lobe comprising three rays bound together as a filament that is longer than the head. The filament may split into two or three branches near its tip. The gap between the upper and lower lobes of the pectoral fin has four to five rudimentary rays. The teeth are in bands. There is one pair of nostrils (not two as often stated). The gill opening is wide, extending from above the pectoral fin almost to the lower lobe. The dorsal finrays number 56–60, the anal fin-rays 52–57, and the pectoral fin-rays 16–21. The pyloric caeca number 9–10. The body is pink to bright red, and the fin margins are black. The peritoneum is dark, and the stomach, mouth, and pyloric caeca are pale. The gill cavity is greyish. The species reaches 31.0 cm in standard length.

source: Karrer (1973).

habitat: This species is pelagic above or near soft mud bottoms

at depths usually below 1,500 m, although its depth range is 1,080– 2,365 m and a single capture in the Beaufort Sea was at 998 m and in Baffin Bay at 731 m. At these depths in northern waters the

Rhodichthys regina



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Rhodichthys regina

temperature is below 0°C and salinity is about 35‰. In the Davis Strait and Baffin Bay it has been collected at 1,124 m to 1,880 m and bottom temperatures of −0.20°C to 0.74°C. They have been observed in Arctic waters in a head-down position over the bottom, with the lower pectoral fin-rays erect and pointing downwards, or touching the bottom. This may be associated with taste organs on the pectoral fin-rays.

biology: Its food items are bathypelagic and planktonic crus-

taceans. Spawning may occur through much of the year as ripe females have been caught from early summer to December. The eggs number 70 or more and are up to 7.0 mm in diameter.

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, the Beaufort Sea, northwest and northeast Greenland, and on both sides of the North Atlantic Ocean.

Distribution of Rhodichthys regina

source: Knudsen, Møller, & Gravlund (2007).

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Caristius fasciatus (Borodin, 1930)

Family Caristiidae

Banded Manefish, cariste barré

Manefishes, Caristes

Brian W. Coad

Caristius fasciatus

Manefishes or Veilfishes are found worldwide in the open ocean. There are about 15 species in the family, including four species on the Atlantic, eastern Arctic, and Pacific coasts of Canada and one in Arctic Canada. The maximum length is at least 35 cm. They have a deep body with a very steep head profile, a long dorsal fin that originates on the head and folds into a fleshy sheath, 6–7 branchiostegal rays, 15 branched caudal fin-rays, and a pelvic fin with 1 spine and 5 elongate soft rays. There are no anal fin spines. The anal fin has a groove, but this extends forward to the base of the pelvic fins and so can accommodate the latter also. The scales are cycloid. The skeleton is weak, and captured fishes are often damaged. The overall colour is light brown with barring and other markings. These are rare fishes of the open ocean where they are epipelagic to bathypelagic (100 m to 2,000 m). They are often found in association with siphonophores. They are oviparous, and the eggs are pelagic. Some species are known from only a few specimens.

sources: Stevenson & Kenaley (2011, 2013).

common names: The common name is coined here from the

Latin name.

taxonomy: The genus comes from the Greek charizomai (to please, help) and thence charistia (pardon, grant a favour), a festival. The species name is the Latin fasciatus (banded). Caristius groenlandicus Jensen, 1941, described from Davis Strait, west Greenland, 62°53' N, 54°15' W, is a synonym. description: This species is distinguished by its high, long dorsal

fin originating on the head above a steep forehead and by the scale sheaths for the dorsal, anal, and pelvic fins. The dorsal fin-rays number 31–37, usually 32–34; the anal fin-rays 18–23, usually 20–21; and the pectoral fin-rays 16–19. The lower, principal caudal fin-ray is serrated. The scales are small and deciduous. The posterior nostril is a slit, and the anterior nostril is circular. The jaws have one row of canine teeth along the sides and two to five rows at the front. The suborbital bone series is not expanded, the space between the orbit and the mouth is narrow, palatine teeth are present, and the upper jaw is long, reaching the posterior level of the orbit. The gill rakers are long, flattened, and overlay each other, numbering 17–20. The overall colour is a pinkish brown, and the fins are black. The snout and jaws are white. The flanks in young bear four black bands, the first from the nape to the gill opening, the second in the middle of the body, the third continued on the dorsal and anal fins, and the fourth around the caudal peduncle. The eye is golden. The peritoneum is black, and since the body wall is thin, this is visible externally. The species reaches at least 30.0 cm in total length.

habitat: This is a bathypelagic species. A Canadian Arctic record

was captured at 918 m and 3.5°C, but fish that are presumably this species have been captured down to 1,660 m in Davis Strait and as shallow as 146 m in Atlantic Canada.



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Caristius fasciatus, adult and young

biology: These Manefishes are quite rare, and their biology is poorly known. Their food includes squid and probably fishes and crustaceans.

importance: It is not economically important. distribution: This Manefish is found from Baffin Bay and Davis Strait south to the waters off Nova Scotia. It may be also in the eastern and South Atlantic Ocean.

Distribution of Caristius fasciatus

sources: Hartel & Triant (1998); Britz & Hartel (2012); Kukuev, Parin, & Trunov (2013).

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Family Zoarcidae Eelpouts, Lycodes

Peter Rask Møller

The family is found from the Arctic to the Antarctic in cold to temperate waters. There are more than 300 species in the family, including 63 species in Canada and at least 6 genera and circa 32 species in the Arctic, making it the most speciose family in the Canadian Arctic. The taxonomy of many of the species is particularly difficult, and several of the species treated here (Gymnelus spp.) need to be studied in more detail before their validity can be confirmed. Several new zoarcid species have been described in recent years from the Canadian Arctic area, and others are being described. This gives an indication of how little is known about the distribution and systematics of these fishes in the enormous Arctic Ocean. Information on their biology is generally very scarce as well. Some are Arctic endemics, whereas others reach the northern limit of a large Atlantic distribution in the southern Canadian Arctic. They are more or less elongated fishes, with a 1:1 ratio of vertebrae and dorsal and anal fin-rays. The mouth is terminal, subterminal, or inferior. The first anal and dorsal fin-rays are segmented and/or divided bilaterally. The caudal fin is small and confluent with the dorsal and anal fins. The pelvic fins are either reduced or absent; if present, they are placed before the pectoral fins. The lateral-line and head-pore

systems are species specific and form important characters for the identification of species. The lateral lines consist of free neuromasts appearing as tiny white dots, smaller than scales. The colour pattern often varies with size and locality and should be used for identification together with other characters. The scales are very small, embedded, or absent. Most morphometric characters change with growth and should be used with caution. Sexual dimorphism is seen in several body proportions, with males being larger, for example, in the length and width of the head. The teeth vary from blunt to sharp, probably reflecting the different diets of the species. The gas bladder is absent, and all but one of the Canadian species (Melanostigma atlanticum) are bottom dwellers. The eggs are few and large and are known to be guarded by the parents in some species. Many species eat small infaunal or epibenthic invertebrates, whereas others feed on larger shrimps and fish. They occur from the shallowest rock pools to the deep sea. Only the Ocean Pout (Zoarces americanus) is of direct commercial importance, though not in Arctic Canada, but several species occur in high densities and are important participants in the Arctic ecosystem.

sources: Anderson (1984, 1994).

Gymnelus barsukovi Chernova, 1999

Barsukov’s Pout, unernak de Barsukov

common names: None. taxonomy: The genus comes from the Greek gymnos (naked) and enchelys (eel). The species is named after the Russian ichthyologist Vladimir V. Barsukov (1922–89), author of The Fauna of the USSR: The Family Anarhichadidae. The original description is based on material from the northeast tip of Ungava Bay and from the Bering, Chukchi (holotype), East

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Siberian, and Laptev Seas. Most of the specimens were previously referred to Gymnelus viridis. The differences between the two species should be studied further because the CO1 bar-code DNA sequence is similar in the two. G. barsukovi has been treated as a junior synonym of G. viridis.

description: The body is elongated, and the distance from

the snout to the anal fin is 35%–39% of standard length. The head is small (14%–16% of standard length). Vertebrae number 20–23 + 74–78 = 95–101; dorsal fin-rays 93–99; and anal fin-rays 74–78. The predorsal distance is 16%–20% of standard length with no free dorsal-fin pterygiophores. The pectoral fin is small (7%–10% of standard length), with 12–14 fin-rays. Scales and pelvic fins are absent. The mouth is terminal. Head pores are moderate sized with three occipital pores. The colour is usually uniformly dark brown. Females and juveniles sometimes have vague irregular light spots, not arranged in any apparent pattern. The anal fin is black in males. Eye spots (ocelli) may be present in the dorsal fin. The peritoneum, mouth, and gill cavities are light. The species reaches 24.3 cm in total length (males) and 18.7 cm in total length (females).

habitat: In the Canadian Arctic (northeastern Ungava Bay, Fox

Harbour Head) it is known from 0–1 m. The overall depth range covers tide pools to 51 m. It is often found beneath stones and boulders.

Distribution of Gymnelus barsukovi

biology: Specimens less than 16.0 cm in length have been

burg & Anderson (2015).

reported to be immature.

sources: Chernova (1999b); Mecklenburg et al. (2011); Mecklen-

importance: It is not economically important. distribution: Found in northern Ungava Bay as a single record at 60°25'18" N, 64°51'18" W (CMNFI 1986-0592B), this Arctic shallow-water species is also known from the Bering, Chukchi, East Siberian, and Laptev Seas.

Gymnelus bilabrus Andriashev, 1937

Twolip Pout, unernak à deux lèvres

common names: None. taxonomy: The species name comes from the Latin bi (two) and labrum (lip), referring to the lip structure, which is free at the sides but attached to the snout on top.

Gymnelus bilabrus

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The original description is based on three specimens from the northern Bering Sea between Cape Chukotsky (Chukchi) and St Lawrence Island, 64°09.5' N, 171°58' W. The species was regarded as synonymous with Gymnelus viridis by authors but was re-established, mainly on the basis of coloration and head pore counts. The differences between the two species should be studied further, as the CO1 bar-code DNA sequence is similar in the two.

description: The body is elongated, and the distance from

the snout to the anal fin is 34%–39% of standard length. The head is small (15%–17% of standard length). Vertebrae number 20–22 + 74–78 = 93–99; dorsal fin-rays 90–96; and anal fin-rays 72–79. The predorsal distance is 16%–20% of standard length with 0–1 free dorsal-fin pterygiophores. The pectoral fin is relatively small (8%–10% of standard length), with 12–15 fin-rays. Scales and pelvic fins are absent. The mouth is terminal. The head pores are moderate sized with three occipital pores. It has 8–11 dark, wide bands with a light cellular pattern on a light background. A dark band on the predorsal area is also present in addition to dark spots on the head, the snout, and the interorbital region. The anal fin is black in males. One eyespot (ocellus) may be present in the anterior part of the dorsal fin. The peritoneum, mouth, and gill cavities are light. The species reaches 19.1 cm in total length (females) and 18.6 cm in total length (males).

habitat: In the Alaskan Beaufort Sea it is known from one record

at 48 m. The overall depth range is from 40 m to 48 m at temperatures from −1.1ºC to 0.7ºC.

?

Distribution of Gymnelus bilabrus

sources: Andriashev (1937); Chernova (1998a); Mecklenburg et al. (2011); Mecklenburg & Anderson (2015).

biology: Unknown. importance: It is not economically important.

Gymnelus knipowitschi

distribution: It is found in the northern Bering, Beaufort, and

Knipowitsch’s Pout, unernak de Knipowitsch

possibly Chukchi Seas. A record at 70°35' N, 146°35' W (CMNFI 1974-0284; Chernova, 1988a) is near the Canadian border in the Beaufort Sea, and the species is included as a possible Canadian one.

Chernova, 1999

common names: None. taxonomy: The species is named after the Russian ichthyologist Nikolai Mikhailovich Knipowitsch (1862–1929).

Gymnelus knipowitschi



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The original description is based on specimens from the Beaufort Sea and off Novaya Zemlya, Barents Sea. Specimens from Arctic seas, identified as Gymnelus hemifasciatus Andriashev, 1937, were described as a new species, G. knipowitschi Chernova, 1999. Thus, G. hemifasciatus is found in the Bering Sea only and not in the Canadian Arctic area. G. knipowitschi may be a junior synonym of G. hemifasciatus as the CO1 bar-code DNA sequence is similar in the two. The difference between the two named species should be documented further with more characters in future studies.

description: The body is elongated, and the distance from the

snout to the anal fin is 35%–44% of standard length. The head is relatively long (16%–20% of standard length). Vertebrae number 18–21 + 68–75 = 88–94; dorsal fin-rays 86–92; and anal fin-rays 71–75. The predorsal distance is 17%–21% of standard length with 1–2 free dorsal-fin pterygiophores. The pectoral fin is relatively small (7%–9% of standard length), with 10–12 fin-rays. The eye is moderate in size, 3.5%–4.8% of standard length; 25%–29% (females) and 23%–28% (males) of head, versus 26%–31% (females) and 19%–25% (males) of head in G. hemifasciatus. Scales and pelvic fins are absent. The mouth is terminal. Head pores are moderate sized with 0–2 occipital pores (3 in G. hemifasciatus). Up to 10–15 dark brownish bands are present. The bands pass below the lateral line in most specimens. They are most distinct at the anterior part of the body. The dorsal part of the head is dark with a light stripe across the predorsal area. The anal fin is black in males. Up to four eyespots (ocelli) may be present in the dorsal fin. The peritoneum, mouth, and gill cavities are light. The species reaches 14.2 cm in total length (males) and 12.6 cm in total length (females).

Distribution of Gymnelus knipowitschi

sources: Andriashev (1937); Anderson (1982); Coad, with

Waszczuk & Labignan (1995); Chernova (1999a); Mecklenburg et al. (2011); Mecklenburg & Anderson (2015).

habitat: In Canadian Arctic records (Beaufort Sea) the species is

found from 6 m to 70 m at temperatures from −1.2ºC to 1.1ºC and salinities from 27.6‰ to 32.1‰. Its overall depth range is from 6 m to 175 m (mostly below 100 m) at temperatures from −1.8ºC to 1.3ºC. It is mainly recorded from sandy and muddy bottoms.

biology: Females with ripe eggs occur from about 9.0 cm standard length. Females carry up to 26 eggs, 4.0 mm in diameter. Spawning probably takes place in autumn in the Barents Sea. The species feeds on bivalves, polychaetes, and gammarid amphipods. importance: It is not economically important. distribution: It is found in Dease Strait, Amundsen Gulf, and Beaufort Sea. An Arctic shallow-water species, it occurs in a semi-circumpolar pattern from the Barents Sea to the Canadian Beaufort Sea, in the Bering Sea, and in the northern Gulf of Alaska in the Pacific Ocean. A record from off west Greenland seems to be an error. Anderson (1982) and Coad, with Waszczuk and Labignan (1995) referred all these to G. hemifasciatus.

Gymnelus retrodorsalis Le Danois, 1913

Aurora Pout, unernak aurore

common names: A local name is Kortfinnet Fiskedoktor (Danish/Greenlandic). Another common name is Aurora Unernak. taxonomy: The species name comes from the Latin retro (back, backward) and dorsalis (pertaining to the back) and refers to the posterior origin of the dorsal fin. The species was originally described from the Denmark Strait (65°52' N, 29°20' W). Most of the specimens from the Barents and Kara Seas described as a new species, Gymnelus andersoni Chernova, 1998, were until then referred to Gymnelus retrodorsalis. The separation of the two species was mainly based on head-pore counts and needs further documentation from additional characters. description: The body is elongated, and the distance from

the snout to the anal fin is 34%–38% of standard length. The head is small (13%–18% of standard length). Vertebrae number 18–21 + 72–80 = 91–100; dorsal fin-rays 82–90; and anal fin-rays

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Gymnelus retrodorsalis

74–81. The most characteristic character is the posterior displacement of the origin of the dorsal fin, the predorsal distance being 23%–29% of standard length, with 4–7 free dorsal-fin pterygiophores. The pectoral fin is small (7%–10% of standard length), with 9–12 fin-rays. Scales and pelvic fins are absent. The mouth is terminal. Head pores are moderate in size with 0–2 occipital pores. Its colour is light to translucent brown with 0–15 light bars. The abdomen and the ventral part of the head are usually light. The anal fin is black in males. Up to five black eye spots (ocelli) with white margins are present in the dorsal fin. The peritoneum, mouth, and gill cavities are light. The species reaches 18.6 cm in standard length.

habitat: This is an Arctic-shelf, upper-slope species. In the Can-

adian Arctic it is known rarely from 7 m and mostly from 55–300 m, at temperatures from −1.3ºC to −0.1ºC, and at high salinities. It has been caught in 5–9 m in northeastern Greenland, down to 481 m off Jan Mayen Island, and down to 581 m in Davis Strait. Its overall temperature range is from −1.8ºC to 4.0ºC. It lives on muddy or sandy bottoms, often associated with shell rubble or gravel.



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biology: It is eaten by Thick-billed Murre chicks at Coats Island,

Hudson Bay; by Black Guillemot chicks at Prince Leopold Island, Lancaster Sound; and by kittiwakes (presumably Black-legged Kittiwakes) in Baffin Bay. Ripening gonads were observed in females of 9.0–10.0 cm standard length. These have about 14–28 eggs, 2.9– 3.1 mm in diameter. Eggs up to 4.5 mm have been reported, and egg numbers may be as low as five. Spawning appears to occur in late summer or early fall. A specimen of 12.2 cm standard length was aged to five years.

importance: It is not economically important. distribution: The species is found in Davis Strait, Frobisher Bay, Ungava Bay, Hudson Strait, northern Hudson Bay, near Ikpiarjuk (Arctic Bay) on northern Baffin Island, in Nansen Sound on Ellesmere Island, and off southern Prince of Wales Island. It is also found south to off Newfoundland; off all the coasts of Greenland, Iceland, Jan Mayen Island, and Svalbard; and in the Barents Sea. According to Chernova (1998b), records from western Arctic Canada and Kara Sea belong to different species.

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The taxonomy of G. viridis is not easy to evaluate. G. bilabrus Andriashev, 1937, has been placed in the synonymy of G. viridis and later re-established with a narrower diagnosis for G. viridis. The species G. knipowitschi, G. esipovi, G. taeniatus, G. barsukovi, and G. platycephalus were described partly from specimens that had been referred to G. viridis until then. The separation of these species from G. viridis was mainly based on head-pore counts and needs further documentation from additional characters. It cannot be elucidated whether some of the above-mentioned species, or even new undescribed species, are present among the Canadian Arctic G. viridis material. Gymnelus viridis var. unimaculatus Richardson, 1855, described from Northumberland Sound is an infra-subspecific taxon, is not available for nomenclatural purposes, and has not been used in North America.

description: The body is elongated, and the distance from

Distribution of Gymnelus retrodorsalis

sources: McAllister, Anderson, & Hunter (1981); Chernova (1998b); Anderson & Fedorov (2004).

Gymnelus viridis (Fabricius, 1780)

Fish Doctor, unernak caméléon

the snout to the anal fin is 31%–39% of standard length. The head is small (12%–18% of standard length). Vertebrae number 17–21 + 70–84 = 89–105; dorsal fin-rays 83–102; and anal fin-rays 72–85. The origin of the dorsal fin is located above the pectoral fin, and the predorsal distance is 14%–22% of standard length, with 0–3 free dorsal-fin pterygiophores. The pectoral fin is small (6%–11% of standard length), with 9–14 fin-rays. The eye is small, 2.1%–3.4% of standard length. Scales are absent. The head pores are moderate in size with 0–2 occipital pores. The colour is variable, usually green to brown with 0–17 light bars. The abdomen and the ventral part of the head are usually light. The anal fin is black in males. Up to six black ocelli with white margins are present in the dorsal fin. The peritoneum, mouth, and gill cavities are light. The species reaches 25.6 cm in standard length.

habitat: An Arctic-shelf species, it is known in the Canadian Arc-

tic from intertidal waters to 110 m, mostly to about 50 m (but to 256 m in Frobisher Bay). At Cape Hatt in northern Baffin Island it was the second most common species after Shorthorn Sculpins, at 23% of subtidal specimens. Temperatures range from −1.4ºC to 4.3ºC; and down to −1.9ºC elsewhere in its distribution area. It lives on muddy bottoms, often associated with brown kelp, and is known to burrow under rocks at Cape Hatt.

biology: The species feeds on amphipods, polychaetes, bivalves,

Gymnelus viridis

common names: Local names are Coogjannernak, Koupjhaunohuk, and Kugsaunak (Inuktitut); and Unernak and Grøn Fiskedoktor (Danish/Greenlandic). Another common name is Green Ocean Pout. taxonomy: The species name is the Latin viridis (green) and refers to the green colour described by Otto Fabricius. It is usually spelled viridis; viride is the original but is seldom used. The original description is based on specimens from southwest Greenland waters.

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mysids, and other infaunal invertebrates. At Cape Hatt the pelagic pteropod Limacina helicina was the major part of the prey biomass at 53%. It is probably eaten by Atlantic Cod in Ungava Bay and is eaten by Thick-billed Murre chicks at Coats Island, northern Hudson Bay. Its lifespan at Cape Hatt is 11 years, with maturity at 7–8 years of age and at lengths of 15.5 cm for females and 17.0 cm for males. Ripening gonads have been found in females of 11.0–12.0 cm standard length. Females have up to 106 eggs, 4.0–4.5 mm in diameter. Spawning appears to occur in late summer or early fall, perhaps in August at Cape Hatt, although there is evidence of mature females in May in the central Canadian Arctic at Brentford Bay, Boothia Peninsula. Spawning may occur under rocks; a ripe male and female have been caught together at Cape Hatt in this type of protected habitat.

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Gymnelus viridis

importance: It is not economically important. distribution: The species is found in Nares Strait, Baffin Bay,

Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, northern Hudson Bay, Foxe Basin, Gulf of Boothia, between the Boothia Peninsula and King William Island, in Queen Maud Gulf, Dease Strait, Bathurst Inlet, the Canadian Arctic islands, as far north as Alert at the tip of northern Ellesmere Island, in Amundsen Gulf and the Beaufort Sea, from off all coasts of Greenland, and in the Chukchi, Bering, East Siberian, Laptev, and Barents Seas. It is also found in Canada south to the Saguenay Fjord but not in southern Hudson Bay and James Bay. This wide distribution was recently abridged by Chernova (1998a), claiming that G. viridis is restricted to Greenland waters only and that records from elsewhere belong to other species. This is a topic for future study.

sources: McAllister et al. (1981); Anderson (1982); Fabijan (1983);

Gillis & Allard (1986); Chernova (1998a, 1998b, 1999a, 1999b); Braune et al. (2014).

Distribution of Gymnelus viridis



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Lycenchelys kolthoffi

biology: Its feeding biology has not been studied in detail, but

Greenlandic).

specimens from the Kara Sea contained small bivalves, Yoldiella frigida. Mature specimens are from 14.0 cm total length in west Greenland waters. About 51% of the females and 55% of the males above this length had ripening gonads. Six females (18.0–19.0 cm) caught in Davis Strait in July had 23–27 yellow-green eggs, with diameters from 4.5 mm to 6.0 mm. These observations indicate an autumn spawning.

taxonomy: The genus comes from the Greek lykos (wolf) and

importance: It is not economically important.

Jensen, 1904

Checkered Wolf Eel, lycode quadrillée

common names: A local name is Kolthoffs Porebrosme (Danish/

enchelys (eel). The species is named after the Swedish zoologist Gustaf Isak Kolthoff (1845–1913), a member of several Arctic expeditions. The species was described from Greenland.

description: The body is elongated, eel-like, and the distance

distribution: The species is found in Nares Strait, Baffin Bay, Davis Strait, Ungava Bay, Hudson Strait, and Amundsen Gulf, as well as on all coasts of Greenland to the Kara Sea.

from the snout to the anal fin is 28%–32% of standard length. The head is small (13%–15% of standard length). Among the Canadian Arctic Lycenchelys, L. kolthoffi has the lowest number of vertebrae, 20–23 + 96–100 = 116–122; of dorsal fin-rays, 114–124; and of anal fin-rays, 96–110. The anterior dorsal-fin pterygiophore is associated with vertebrae numbers 5–6. The last precaudal vertebra is associated with dorsal fin-rays numbers 15–16. The pectoral fins are relatively large (9.3%–11.6% of standard length) with 14–15 finrays. The squamation in L. kolthoffi differs from other species in the area by the naked abdomen and by the relatively few (10–13) vertical scale rows between the anal fin origin and the dorsal fin base. The lateral-line system has ventral and medio-lateral lines. A short row of neuromasts in front of the dorsal fin is also present. The head pores large. The coloration is very characteristic, with a darkand-light striped dorsal fin. The body and head are light with dark reticulations dorsally and dark spots on the ventral side. The coloration of the dorsal fin is a very useful character to separate L. kolthoffi from all other Lycenchelys spp. in the Atlantic and Arctic Oceans. The dark spots on the abdomen are often seen in Lycenchelys sarsii, but the dorsal fin of this species is always uniformly brownish. The peritoneum is black. The species reaches 21.0 cm in total length. Distribution of Lycenchelys kolthoffi

habitat: It is an Arctic shelf-to-slope species. Canadian Arctic specimens from Ungava Bay and Hudson Strait were caught in depths of 244–360 m, in Baffin Bay at 672–930 m, and in the Beaufort Sea at 350 m. Off west Greenland it is known from depths of 202–672 m and temperatures of 1.7ºC to 4.1ºC. Elsewhere in its distribution area it is found from depths of 202–930 m and temperatures of −0.9ºC to 3.0ºC. It lives on sandy and muddy bottoms with stones.

sources: Jensen (1904); Møller (1995); Møller & Jørgensen (2000).

Lycenchelys kolthoffi

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Lycenchelys muraena (Collett, 1878)

Moray Wolf Eel, lycode murène

common names: A local name is Muræne-porebrosme (Danish/

Greenlandic).

taxonomy: The species name comes from the Greek myraina

(moray) after the superficial resemblance to moray eels. The species was originally described from 325 km west-southwest of Bodø, Norway.

description: The body is elongated, eel-like, and the distance

from the snout to the anal fin is 26%–29% of standard length. The head is very small (11%–13% of standard length). Vertebrae number 23 + 98–102 = 121–125; dorsal fin-rays 109; and anal fin-rays 100. The anterior dorsal-fin pterygiophore is associated with vertebra number 6, and the last precaudal vertebra is associated with dorsal fin-rays numbers 17–18. The predorsal length is 15.1%–18.2% of standard length. The pectoral fins are relatively small (7.5%–9.1% of standard length) with 14–15 fin-rays. The squamation is dense, covering the abdomen, and with 14–19 vertical scale rows between the anal fin origin and the dorsal fin base. The lateral-line system has a well-developed ventral line. A medio-lateral row and a dorsal row of separate neuromasts are also present, as is a short row in front of the

dorsal fin. The head pores are large. The colour is light to dark brown dorsally and lighter ventrally. The dorsal fin is dark, and the anal fin is light. The pectoral fins are light, with a dark dorsal margin. The coloration of the pectoral fin makes it a very useful character to separate L. muraena from all other Lycenchelys spp. in the Atlantic and Arctic Oceans. The peritoneum is black. The species reaches 21.0 cm in total length.

habitat: An Arctic deep-water species with few records from the

Canadian Arctic it is found at depths of 551–1,337 m at temperatures of 0.3ºC–0.5ºC; off Baffin Island it has been found at 1.5ºC at 676 m and as shallow as 239 m. Off west Greenland it occurs at depths of 534–1,271 m and temperatures of 1.5ºC–3.4ºC. Elsewhere in its distribution area it is known at depths of 467–1,371 m and temperatures of −0.7ºC to 3.4ºC. It lives on muddy bottoms.

biology: Its feeding biology has not been studied in detail, but

specimens from the Norwegian Sea contained small crustaceans, Podocerus assimilis and Astacilla granulata. A female of 20.3 cm total length, caught in September in the Denmark Strait, had 21 light-yellow eggs, 8.0 mm in diameter.

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, Hudson Strait, and northwest, southeast, and northeast Greenland to the Kara Sea.

Lycenchelys muraena



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description: The body is elongated, eel-like, and the distance

from the snout to the anal fin is 26%–34% of standard length. The head is small (11.0%–15.5% of standard length). Among the Canadian Arctic Lycenchelys spp., Lycenchelys paxillus has the highest number of vertebrae, 23–27 + 102–112 = 127–137; of dorsal fin-rays, 112–127; and of anal fin-rays, 106–117. The anterior dorsal-fin pterygiophore is associated with vertebrae numbers 8–11. The last precaudal vertebra is associated with dorsal fin-rays numbers 15–18. The pectoral fins are relatively small (6.5%–9.1% of standard length) with 13–17 fin-rays. The eye is relatively large (2.4%–3.7% of standard length). The squamation is dense, covering the abdomen, predorsal area, and dorsal and anal fins. The distance from the snout to the anterior scales is 10.0%–15.9% of standard length, and there are 20–28 scale rows between the anal fin origin and the dorsal fin. The lateral-line system consists of ventral, medio-lateral, dorsal, and predorsal lines. The head pores are large, there being 7 (rarely 6) suborbital head pores, versus 6 (rarely 7) in L. sarsii. The coloration is generally uniformly light to dark brown when the fish is fresh or recently preserved. Most juveniles and some adults are slightly lighter on the ventral side. The peritoneum is black. The species reaches 36.0 cm in total length, which is considerably larger than other Canadian Arctic Lycenchelys spp. Distribution of Lycenchelys muraena

sources: Jensen (1904); Collett (1905); Møller & Jørgensen (2000); Treble et al. (2000).

Lycenchelys paxillus (Goode and Bean, 1879)

habitat: It is a boreal shelf-to-slope species. Canadian Arctic specimens from Davis Strait were caught at 658 m, 1.6ºC. In the eastern part of the strait it occurs at depths of 424–1,093 m and temperatures of 0.9ºC–5.5ºC, most commonly at depths of 600–800 m and temperatures of 3ºC–5ºC. Elsewhere it is known from off Nova Scotia to Virginia at depths of 46–1,525 m and temperatures down to −0.9ºC. It lives on muddy bottoms. biology: In Norfolk Canyon molluscs were found to be the most

Greenlandic).

important food (bivalves, Nuculanidae; gastropods, Scaphandridae, Retusidae, Buccinidae; scaphopods). The cumacean Diastylis sp. was the most important crustacean. All stomachs contained sediment, indicating that L. paxillus searches for infaunal food items. Mature specimens were from 18.0 cm total length. Females have up to 85 eggs, up to 4.1 mm in diameter.

taxonomy: The species name is the Latin paxillus (a peg).

importance: It is not economically important.

Common Wolf Eel, lycode commune

common names: A local name is Stor Porebrosme (Danish/

The original description is based on a specimen caught between the Le Have and Sable Island banks (42°48' N, 63°07' W). Lycenchelys ingolfianus Jensen, 1902, described from the Davis Strait, is a junior synonym. Lycenchelys paxillus can easily be confused with L. sarsii.

distribution: The species is found in Davis Strait, southwest Greenland, and southwards to off Virginia. It is endemic to the northwestern Atlantic Ocean.

Lycenchelys paxillus

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Lycenchelys paxillus

Lycenchelys sarsii (Collett, 1871)

Theologian Eelpout, lycode de Sars

common names: A local name is Sars Porebrosme (Danish/

Greenlandic). Another common name is Sars’ Wolf Eel.

taxonomy: The species is named after the Norwegian zoologist and theologian Michael Sars (1805–69). The species is easily confused with the related L. paxillus. It was originally described from Utne, Hardanger Fjord, southwestern coast of Norway. description: The body is elongated, eel-like, and the distance

Distribution of Lycenchelys paxillus

sources: Jensen (1904); Wenner (1978); Hudon (1990); Møller (1999); Møller & Jørgensen (2000).



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from the snout to the anal fin is 28%–33% of standard length. The head is small (13%–16% of standard length). Vertebrae number 21–25 + 97–102 = 121–126; dorsal fin-rays 106–112; and anal fin-rays 101–105. The anterior dorsal-fin pterygiophore is associated with vertebrae numbers 9–12. The last precaudal vertebra is associated with dorsal fin-rays numbers 12–14. The pectoral fins are relatively small (7.9%–10.7% of standard length) with 15–16 fin-rays. Squamation is dense, covering the abdomen and bases of the dorsal and anal fins. The predorsal area is scaled. The distance from the snout to the anterior scales is 13.9%–16.7% of standard length, with 18–21 scales between the anal fin origin and the dorsal fin. The lateral-line

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Lycenchelys sarsii

system consists of ventral, medio-lateral, dorsal, and predorsal lines. The head pores are large, with 6 (rarely 7) suborbital head pores, versus 7 (rarely 6) in L. paxillus. The coloration is generally dark brown dorsally and light on the ventral side, sometimes with irregular brown spots on the abdomen and along the anal fin. The pectoral fin is light anteriorly, dark posteriorly. The peritoneum is black. The species reaches 23.0 cm in total length, which is considerably smaller than L. paxillus.

habitat: It is a boreal shelf species, with records from the Canadian Arctic (Davis Strait) caught in depths of 350–460 m, at temperatures around 4ºC. In the eastern Davis Strait it is caught between 60º15' N and 68º53' N at depths of 107–572 m and temperatures of 2.2ºC –5.6ºC. It is, however, rare in depths below 200 m and temperatures below 3ºC. A cruise record from the Canadian Davis Strait was at 755 m. In the northeastern Atlantic it is known from depths of 150–600 m and temperatures of 0ºC–6ºC. It lives on muddy bottoms.

biology: In the Barents Sea it is known to feed on polychaetes,

amphipods, isopods, cumaceans, copepods, and small bivalves (Yoldiella and Propeamissium). Mature specimens are from 11.0 cm total length off west Greenland. About 60% of the females and 70% of the males above this length had ripening gonads. Six females (14.0–16.0 cm) caught in July in Davis Strait had 12–18 eggs, with diameters of 3.5–4.5 mm.

Distribution of Lycenchelys sarsii

sources: Møller & Jørgensen (2000); Treble et al. (2000).

importance: It is not economically important. distribution: The species is found in Davis Strait, off southwest Greenland; the Grand Banks off Newfoundland; north of Iceland; and the Norwegian fiords from Skagerrak to Kola Bay.

Lycodes adolfi

Nielsen and Fosså, 1993 Adolf’s Eelpout, lycode d’Adolf

common names: A local name is Adolfs Ålebrosme (Danish/

Greenlandic).

taxonomy: The genus comes from the Greek lycodes (wolfish) or

lykos (wolf) and the suffix -oides (similar to). The species is named after Adolf Severin Jensen (1866–1953), a naturalist and ichthyologist

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Lycodes adolfi

of the Copenhagen Zoological Museum who conducted much work on the fishes of Greenland. The original description is based on specimens from off east Greenland and Baffin Bay. It is probably closely related to Lycodes frigidus. Scaleless juveniles of less than 10.0 cm of the two species are difficult to separate, but L. adolfi differs by having fewer pectoral finrays (16–19 versus 19–23) and fewer precaudal vertebrae (19–21 versus 21–24).

distribution: It is found in Baffin Bay, Davis Strait, Ungava Bay, the Beaufort Sea, northwest and northeast Greenland, the Greenland Sea, Jan Mayen Island, the eastern Norwegian Sea, and north of Spitsbergen in the Arctic Ocean. It may also occur in the deep central Arctic Ocean.

description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 38%–44% of standard length. The head is relatively small (20.7%–23.7% of standard length). Vertebrae number 19–21 + 77–84 = 97–104; dorsal fin-rays 89–98; and anal fin-rays 79–85. The pectoral fins are rounded and of moderate length (12%– 18% of standard length), with 16–19 fin-rays. Squamation is reduced, covering the posterior part of the tail only. This kind of squamation is also found in Lycodes seminudus, but L. adolfi differs from that species by a longer tail and a ventral lateral line. No medio-lateral neuromasts are present, but rows of widely spaced neuromasts are found dorsally and predorsally. Coloration is always uniformly brown, without crossbars. The head, peritoneum, and fins are black. The skin is thin and fragile. A small species, it has a maximum total length of about 24.0 cm.

habitat: It is an Arctic deep-sea species, with Canadian Arctic

records from Baffin Bay caught at 1,183–1,880 m and temperatures of −0.08ºC to 0.5ºC. In the southeastern part of the bay it occurs in depths of 420–1,436 m and temperatures of 0.1ºC–3.7ºC. An Ungava Bay record from cruise data was at 274 m. In the Beaufort Sea it is found from 500 m to 1001 m. In the east Greenland and Iceland area it has been caught at depths of 386–1,380 m and temperatures between −1.1ºC and −0.2ºC. It is most common in depths of more than 1,000 m and goes as deep as 2,150 m. It lives on muddy bottoms.

Distribution of Lycodes adolfi

sources: Nielsen & Fosså (1993); Møller & Jørgensen (2000);

Treble et al. (2000); Byrkjedal, Brattegard, & Møller (2009); Byrkjedal, Langhelle, Wenneck, & Wienerroither (2011); Mecklenburg et al. (2014).

biology: The species feeds on crustaceans, for example, amphipods of the Family Amphilochidae. A female of 20.4 cm standard length, caught in the Baffin Bay in October, contained 19 yellow eggs, 6.0 mm in diameter. importance: It is not economically important.



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Lycodes esmarkii Collett, 1875

Greater Eelpout, grande lycode

common names: A local name is Esmarks Ålebrosme (Danish/Greenlandic). Other common names are Esmark’s Eelpout and lycode d’Esmark. taxonomy: The species is named after the Norwegian zoologist

and Oslo University professor Lauritz Martin Esmark (1806–84). It was originally described from Varanger Fjord, northern Norway. Lycodes vachonii Vladykov and Tremblay, 1936, described from the St Lawrence River estuary, is a junior synonym. It is probably a close relative of Lycodes eudipleurostictus and L. terraenovae.

description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 36.5%–44.5% of standard length. The head is relatively small (20.7%–24.9% of standard length). Vertebrae number 23–25 + 90–94 = 114–117; dorsal fin-rays 105–108; and anal fin-rays 90–94. The pectoral fins are rounded and of moderate length (10.9%–14.6% of standard length), with 21–23 fin-rays. Squamation is dense and covers the predorsal area, the abdomen, the dorsal and anal fins, and the pectoral fin base. The lateral-line system consists of a well-developed ventral line, in addition to medio-lateral, dorsal, and predorsal rows of widely spaced neuromasts. Coloration is very characteristic, grey to black dorsally, with 4–9 white marks. The marks change from more or less straight bars

in juveniles to Y-shaped or quadrate-shaped marks in adults. A straight bar, or two white spots, is present above the opercular lobes. The head and the ventral parts of the body are lighter, greyish. The pectoral fins are dark dorsally and light ventrally. The peritoneum and the oral cavity are black. This species lacks pyloric caeca, which is unique. All other Canadian Arctic Lycodes have two. It reaches 75.0 cm in total length, but fishes of more than 45.0 cm are rare.

habitat: It is an Atlantic boreal, shelf, and slope species, with

Canadian Arctic records from the Davis Strait caught at depths of 146–619 m and at temperatures of 1.5ºC–4.4ºC. Records from collections in Ungava Bay were at 240–425 m, and from Hudson Strait at 274–431 m. Another collection from Ungava Bay was as deep as 950 m. In the eastern part of Davis Strait it occurs in depths of 310–1,090 m and temperatures of 1.0ºC–5.3ºC. The overall depth range is 143–1,090 m, and temperatures range from −0.4ºC to 5.6ºC. It lives on muddy bottoms.

biology: The species feeds almost exclusively on ophiurids. In the

southern Davis Strait, ripening gonads were observed in specimens above 35.0 cm (females) and 38.0 cm (males). A 36.0 cm female had 216 eggs, but up to 1,200 eggs, 6.0 mm in diameter, have been reported. In Icelandic waters ripe specimens were reported in June.

importance: It is not economically important. distribution: It is found in Davis Strait; central and eastern

Hudson Strait and Ungava Bay; northwest, southwest, and southeast Greenland; southward to off Virginia in the western Atlantic;

Lycodes esmarkii

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and from east Greenland to the southern Barents Sea in the eastern Atlantic. Cruise records from Baffin Bay (not mapped) are probably in error and need confirmation by voucher specimens deposited in a museum.

Lycodes eudipleurostictus Jensen, 1902

Doubleline Eelpout, lycode à deux lignes

common names: A local name is Dobbeltliniet Ålebrosme (Danish/Greenlandic).

taxonomy: The species name comes from the Greek eu (well), di (in two), pleura (side), and stiktos (punctured) and refers to the well-developed medio-lateral and ventro-lateral lateral lines. The original description is based on specimens from the Norwegian Sea and west Greenland waters. It is suggested to be a close relative of Lycodes esmarkii and L. terraenovae. description: It is a “long-tailed” Lycodes, and the distance from

Distribution of Lycodes esmarkii

sources: Jensen (1904); Joensen & Tåning (1970); Markle & Sedberry (1978); Hudon (1990); Valtysson (1995); Møller (2000a); Møller & Jørgensen (2000); Jørgensen et al. (2005).

the snout to the anal fin is 39.1%–44.9% of standard length. The head is moderate in size (20.4%–24.3% of standard length). L. eudipleurostictus has a high number of vertebrae, 20–23 + 84–91 = 105–112; of dorsal fin-rays, 98–106; and of anal fin-rays, 86–92. The pectoral fins are usually emarginate, of moderate length (12.3%–15.6% of standard length), with 19–23 fin-rays. Squamation is dense even in juveniles (from ca. 8.0 cm in standard length), with 15–34 scales in a vertical row between the anal fin origin and the dorsal fin (increasing with size). The predorsal area, the abdomen, and the dorsal and anal fins are almost completely scaled in adults. The pectoral fin base is always naked (scaled in L. esmarkii and L. terraenovae). The lateral-line system is characteristic in having well-developed medio-lateral and ventral lines in addition to dorsal and predorsal rows of widely spaced neuromasts. The palatine and premaxillary teeth rows are of equal length in L. eudipleurostictus, which is different from the closely related L. esmarkii where the premaxillary row is 1.6–1.8 times as great as the palatine row. Coloration is extremely variable: light greyish to dark brown. Juveniles and most adults have 5–13 light bars on the body and the dorsal fin, and only occasionally a straight light band on the predorsal area connecting the opercular lobes. Juveniles have a blue belly. The pectoral fin is usually dark with a light margin. The peritoneum is black. A uniformly black variety is known from a few specimens. The species reaches 45.0 cm in total length (males) and 40.0 cm in total length (females).

Lycodes eudipleurostictus



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habitat: It is an Arctic deep-water species, with Canadian Arctic specimens from Baffin Bay caught in depths of 441–1,288 m and temperatures of 0.2ºC–4.0ºC. Cruise data from Baffin Bay to Hudson Strait gave a depth range of 169–1,366 m. In the Beaufort Sea it is known from five specimens caught at a depth range of 343–464 m. The overall depth range is 25–1,287.5 m (but rarely shallower than 250 m), and temperatures range from −1.1ºC to 4.9ºC. biology: Small specimens eat mainly endobenthic prey, but the

amount of epibenthic food items increase with size. Its food includes crustaceans, polychaetes, ophiurids, and priapulids. Large males are often caught on long lines in west Greenland fiords. In eastern Baffin Bay, ripening gonads were observed in specimens above 23.0 cm in length (females) and 29.0 cm in length (males). The largest females have up to 300 orange eggs, up to 8.0 mm in diameter.

importance: It is not economically important. distribution: The species is found from Nares Strait, Baffin Bay,

Davis Strait, Hudson Strait, Ungava Bay, the Alaskan Beaufort Sea, recently confirmed in the Canadian Beaufort Sea, and off all the coasts of Greenland to the northern Kara Sea, north of Severnaya Zemlya.

Lycodes frigidus Collett, 1879

Glacial Eelpout, lycode glaciale

common names: A local name is Småskællet Ålebrosme (Danish/Greenlandic). Other common names are Abyssal Eelpout and Coldwater Eelpout. taxonomy: The species name is the Latin frigidus (frozen, cold). It

was originally described from Bear Island and Spitsbergen. Earlier reports on this species may have been confused with the newly described Lycodes adolfi. The two species have a different squamation, and young scaleless specimens of L. frigidus can usually be identified by the higher number of pectoral fin-rays (19– 23 versus 16–19) and of precaudal vertebrae (21–24 versus 19–21). The Glacial Eelpout has also been confused with the Atlantic deepsea zoarcid Lycodes terraenovae in early literature. The figure in Coad, with Waszczuk and Labignan (1995) is a mistake, showing a specimen of the genus Pachycara.

description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 40%–50% of standard length. The head is relatively long (22.3%–25.9% of standard length), especially in large specimens. Vertebrae number 21–24 + 79–85 = 101–107; dorsal fin-rays 92–98; and anal fin-rays 79–85. The pectoral fins are large (12.1%–17.7% of standard length) with 19–23 fin-rays. Squamation is unique with the smallest scales in the genus. Individuals of less than 11.0 cm standard length are generally without scales. Specimens above 15.0 cm have more than 35 scales in a vertical row between the anal fin origin and the dorsal fin. The number of rows increases with the size of the fish, up to 75 in the largest specimens. The dorsal and anal fins are almost completely scaled in adults. The predorsal area, the abdomen, and the pectoral fin base are scaled in adults. The lateral-line system consists of a well-developed ventral line in addition to predorsal and dorsal rows of wide-spread neuromasts. Medio-lateral neuromasts are absent. The skin is relatively thin and fragile. The coloration is uniformly greyish to brown or black. Juveniles are generally lighter than adults. The peritoneum is black. The species reaches 69.0 cm in total length and 1.986 kg in weight.

habitat: It is an Arctic deep-sea species, the only Canadian Arctic

Distribution of Lycodes eudipleurostictus

sources: McAllister et al. (1981); Valtysson (1995); Møller & Jørgensen (2000); Treble et al. (2000); Jørgensen et al. (2005).

specimen from the central Arctic Ocean being caught at a depth of 2,075 m and a temperature of −0.37ºC. It descends to 2,505 m in the central Arctic Basin. Northeastern Atlantic specimens are reported at depths of 491–3,000 m, at temperatures below 0ºC, and west of Svalbard at depths to 3,576 m. It is rare in depths less than 1,000 m and lives on muddy bottoms.

biology: It feeds on crustaceans (Mesidothea megalura, Eurycope cornuta, Astacilla granulata, Pasiphaea tarda, Phoxus crenulatus, Padocerus assimilis, Themisto libelula, Hymenodora glacialis), ophiuroids, cephalopods, molluscs, sipunculids, and fishes. The age

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Lycodes frigidus

range west of Svalbard was 5–33 years. Growth is correlated with temperature – the higher the mean annual temperature, the higher the growth rate – but maximum age decreases with increased temperature. The length gain per year is very low, at 0.63–2.4 cm. A 50.0 cm female caught in August north of the Faroe Islands contained 500 eggs, 7.0 mm in diameter.

sources: Jensen (1904); Nizovtsev, Ponomarenko, & Shevelev

(1976); Prouse & McAllister (1986); Coad, with Waszczuk & Labignan (1995); Møller (1996); Hildebrandt, Bergmann, & Knust (2011).

importance: It is not economically important. distribution: It is found northwest of Ellesmere Island in the Canadian Arctic, and elsewhere in the Norwegian, Greenland, Laptev, East Siberian, and Chukchi Seas. A record at 85º48'27" N, 110º43'39" W, is the northernmost collection for the Canadian Arctic. All Glacial Eelpouts reported from Baffin Bay have later been re-identified to other species.

Lycodes jugoricus Knipowitsch, 1906

Shulupaoluk, lycode plume

common names: None. taxonomy: The species is named after the type locality, Jugorsky Shar Strait between the eastern Barents Sea and the Kara Sea. description: It is a “short-tailed” Lycodes, and the distance from

the snout to the anal fin is 43%–52% of standard length. The body is relatively robust with a moderate-sized head (20.7%–25.3% of standard length). Vertebrae number 24–25 + 75–77 = 99–102; dorsal fin-rays 90–105; and anal fin-rays 74–84. The pectoral fins are long (13.4%–17.8% of standard length) with 16–19 fin-rays. Scales are absent. The lateral line consists of medio-lateral, dorsal, and predorsal rows of neuromasts. The head pores are small. Dentition is very characteristic, the teeth being very robust and blond. The teeth on the vomer and the palatines form an almost uninterrupted roof of teeth in the mouth, probably an adaptation to bivalve food. The coloration changes with the size. Juveniles and medium-sized adults have 7–13 light bars on the body and the dorsal fin and a straight light band on the predorsal area connecting the opercular lobes. The dark areas in between the bars are triangular. Large specimens have an almost uniform coloration, with a few reticulations. The peritoneum is light. The species reaches 38.5 cm in total length (males) and 47.5 cm in total length (females). Distribution of Lycodes frigidus



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Lycodes jugoricus

habitat: It is an Arctic shallow-water species. Specimens from the

Canadian Arctic (Beaufort Sea) were caught at depths of 3–37 m, rarely to 46 m, at temperatures from −1.4ºC to 5.0ºC, elsewhere to 90 m and 8.0ºC. This is a brackish-water species found in water with low salinity (12‰–32‰). It is found in the mixed estuarine zone of the Mackenzie River and favours the deeper, relatively warmer, winter waters of Tuktoyaktuk Harbour and the deeper inlets of Liverpool Bay. In the Eurasian Arctic it is reported at depths of 9–15 m, rarely to 90 m. It lives on muddy and sandy bottoms and on stones.

biology: In Tuktoyaktuk Harbour polychaetes predominate in the

diet, with some mysids, isopods, and fish. Russian specimens were reported to feed on amphipods, isopods, bivalves, and polychaetes. Spawning probably occurs in autumn with about 1,280 eggs, 4.5 mm in diameter.

importance: It is not economically important. distribution: The species is found at the southwest Boothia Peninsula (Spence Bay), Dease Strait, and Beaufort Sea, and from the White, Kara, Laptev, and Chukchi Seas, but not apparently in the Alaskan Beaufort Sea.

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Lycodes lavalaei

Vladykov and Tremblay, 1936 Laval Eelpout, lycode de Laval

Lycodes lavalaei

common names: Other common names are Newfoundland Eelpout and lycode de Labrador. “Newfoundland Eelpout” (and “lycode de Labrador”) are used by the American Fisheries Society (see the bibliography), whose list does not include L. terraenovae. The latter species is more appropriately called Newfoundland Eelpout. taxonomy: The species was named as a dedication to the Université Laval in Québec. The original description is based on specimens from the Gulf of St Lawrence (Trois-Pistoles) and Newfoundland. The name is incorrectly spelled lavalei by some authors. Mixed with L. reticulatus in some literature – for example, the figure of L. reticulatus in Scott and Scott (1988, p. 410) – is an adult L. lavalaei. It is suggested to be a close relative of Lycodes raridens Taranetz and Andriashev in Andriashev, 1937, of the North Pacific Ocean. description: It is a “short-tailed” Lycodes, and the distance from

the snout to the anal fin is 42%–54% of standard length. The head is long (20%–27% of standard length) and broad (9%–18% of standard length), especially in large specimens. In the group of “short-tailed” Eelpouts, L. lavalaei has the highest number of vertebrae, 23–27

+ 74–80 = 99–105; of dorsal fin-rays, 93–97; and of anal fin-rays, 74–80. The pectoral fins are large (13.7%–18.7% of standard length) with 18–20 fin-rays. The caudal fin is short, measuring 1.4%–2.6% of standard length, versus 2.9%–4.4% of standard length in L. reticulatus, in specimens greater than 22.5 cm standard length. Squamation is dense even in juveniles (from ca. 8.0 cm standard length), with 21–37 scales in a vertical row between the anal fin origin and the dorsal fin (increasing with the size of the fish). The dorsal and anal fins are almost completely scaled in adults. The predorsal area is scaled in most adults. The belly and the pectoral fin base are always naked. The lateral-line system consists of medio-lateral, dorsal, and predorsal rows of neuromasts. The head pores are relatively large (seen without magnification in adults). The coloration changes with size. Juveniles have 6–9 light bars on the body and the dorsal fin, and a straight light band connecting the opercular lobes. The band on the predorsal area is said to be narrower in L. lavalaei than in L. reticulatus, but individual variation is too large in this character to be useful. The bands become reticulated and eventually disappear (in specimens greater than ca. 38.0 cm), so that the adult colour is brownish to greyish with black and light spots and reticulations. During growth an uninterrupted black band develops near the margin of the dorsal fin, flanked by a narrow light band along the edge of the fin. The uninterrupted dark-and-white stripes at the dorsal fin margin are a very useful character, which separates L. lavalaei from all other Lycodes in the Atlantic and the Arctic. The Arctic Eelpout (L. reticulatus) has a coloration somewhat like L. lavalaei but never develops continuous black-and-white stripes at the dorsal fin margin. The peritoneum is light. The species reaches 59.0 cm in total length (males) and 51.0 cm in total length (females).

habitat: It is a sub-Arctic species generally, with Arctic specimens

from Frobisher Bay and Hudson Strait caught in depths from 57 m to 388 m. Off Labrador it is reported at depths from 24 m to 128 m, at temperatures of −1.2ºC to 2.5ºC. In the Gulf of St Lawrence it occurs from 25 m to 200 m, at temperatures of 0.3ºC to 3.1ºC, and on the banks off Newfoundland from 80 m to at least 300 m. The overall depth range is 24–535 m. It lives on muddy and sandy bottoms and over gravel.

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biology: Its food consists of both bottom invertebrates and fishes

such as other Eelpouts. A 53.5 cm specimen was reported to have eaten a 21.0 cm Lycodes sp. Two females (31.3 cm and 46.8 cm in standard length), caught at the Newfoundland banks in April and December, contained about 300 and 1,800 eggs, 6.0 mm and 3.5 mm in diameter. These observations indicate a spring spawning, which is not in accordance with literature suggesting an autumn spawning.

Lycodes luetkenii Collett, 1880

Pink Eelpout, lycode rose

importance: It is not economically important.

common names: A local name is Lütkens Ålebrosme (Danish/Greenlandic). Other common names are Lütken’s Eelpout and lycode de Lütken.

distribution: The species is found in Frobisher Bay, Ungava Bay,

taxonomy: The species is named after the Danish ichthyolo-

and Hudson Strait and southwards along the coasts of Labrador and Newfoundland (including the Grand Banks) into the Gulf of St Lawrence, at least to the Saguenay Fjord. It is found mainly in the western North Atlantic, with a couple of records from the eastern North Atlantic (Jan Mayen and Faroe Islands) needing confirmation. This species does not occur in Greenland waters.

gist Christian Frederik Lütken (1827–1901) who studied the fishes of Greenland. It was originally described from 115 km west of Norskøerne, Spitsbergen. This species is often misidentified as the Arctic Eelpout (L. reticulatus) and is therefore rarely reported.

description: It is a “short-tailed” Lycodes, and the distance from

the snout to the anal fin is 46%–55% of standard length. The head is long (24%–29% of standard length). The vertebrae number 22–24 + 71–75 = 93–98; dorsal fin-rays 89–93; and anal fin-rays 70–73. The pectoral fins are very large (15%–20% of standard length) and have the highest number of fin-rays (22–24) among the short-tailed Lycodes. Squamation is dense with 22–29 scale rows in a vertical row between the anal fin origin and the dorsal fin. The fins, predorsal area, and abdomen are naked. The lateral-line system consists of medio-lateral, dorsal, and predorsal rows of neuromasts. The head pores are relatively large, especially in young specimens. Fresh specimens have pink pectoral fins and a more or less pink coloration on the body. The pink colour, however, disappears on preservation, leaving a grey-brown ground colour. The abdomen, the peritoneum, and the ventral parts of the head are light. All specimens have 6–8 irregular light bars dorsally on the body continuing onto the dorsal fin. The dorsal fin is dark between the bars. A light horseshoe-shaped bar on the nape is also present. The species reaches 55.0 cm in total length.

habitat: Canadian Arctic specimens (Baffin Bay) were caught in Distribution of Lycodes lavalaei

sources: Vladykov & Tremblay (1936); Backus (1957); Whitehead, Bauchot, Hureau, Nielsen, & Tortonese (1984–6); Hudon (1990); Møller (1996).

depths of 552 m and 635 m and temperatures of 1.2ºC–1.5ºC, and in southern Davis Strait at 422–1,463 m. In Ungava Bay it has been recorded as shallow as 365 m. Off west Greenland it was reported at depths of 259–553 m, at temperatures of 1.5ºC–3.4ºC. The depth range of the species is 112–1,463 m, and the temperature range is from −1.0ºC to 3.4ºC. It is rarely caught in trawls, indicating that it might prefer hard-bottom localities.

biology: Its food consists of both bottom invertebrates and fishes.

The type-specimen was reported to have eaten a Cottunculus sp. A 46.2 cm female, caught off west Greenland in August, contained about 1,000 orange eggs, 5.0 mm in diameter.

importance: It is not economically important.

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Lycodes luetkenii

distribution: The species is found in Baffin Bay, Davis Strait, Cumberland Sound (but no accurate locality), Hudson Strait, Ungava Bay, northwest, southwest, and northeast Greenland, Denmark Strait, and the Norwegian, northern Barents, and Kara Seas.

Lycodes marisalbi Knipowitsch, 1906

White Sea Eelpout, lycode de la mer Blanche

common names: None. taxonomy: The species name comes from the Latin maris (sea)

and albus (white), named after the type locality, the White Sea, Russia. The taxonomy of this species is complicated. It has been regarded as a subspecies of Lycodes pallidus since 1954, and it was thought to occur in the White Sea only. Recently it was discovered that the specimens from the western Canadian Arctic, hitherto identified to L. pallidus, belong to L. marisalbi. Furthermore, it has been mixed up with the Archer Eelpout (Lycodes sagittarius), which has the same type of lateral-line system as L. marisalbi.

description: It is a “long-tailed” Lycodes, and the distance from

Distribution of Lycodes luetkenii

sources: Collett (1880); Møller & Petersen (1997); Møller & Jørgensen (2000); Treble et al. (2000).



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the snout to the anal fin is 38.6%–44.9% of standard length. The head is relatively small (17.9%–24.7% of standard length). The vertebrae number 19–22 + 78–85 = 98–106; dorsal fin-rays 88–96; and anal fin-rays 76–83. The pectoral fins are rounded and of moderate length (10.8%–15.7% of standard length), with 16–18 fin-rays. Squamation on the body is weak, there being 9–21 scales in a vertical row from the anal fin origin to the dorsal fin base. The predorsal area, the abdomen, the dorsal and anal fins, and the pectoral fin base are naked. The lateral-line system consists of a ventro-medio-lateral line, with the bend well behind the anal fin origin. A short predorsal row and a dorso-lateral row of widely spaced neuromasts are also present. The coloration is light to dark brown with 6–11 bars dorsally on the body and the dorsal fin. Uniformly brown specimens are not uncommon (about 30%) in the Beaufort Sea population but are not seen in the White Sea. A straight white band connects the opercular lobes in some specimens. Scales do not appear as white spots.

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Lycodes marisalbi

The peritoneum, oral chamber, and branchial chamber are light, speckled with tiny dark spots. The species reaches 14.4 cm in standard length (females) and 21.3 cm in standard length (males).

habitat: It is an Arctic shallow-water species, with Canadian Arctic records from the Beaufort Sea at depths from 3 m to 335 m and temperatures from −1.8ºC to 2.5ºC, usually below 0ºC. In the White Sea it occurs in depths from 51 m to 326 m and a temperature of −1.4ºC to −0.4ºC. It lives on muddy and sandy bottoms in low-saline water, usually 28‰–32‰, influenced by Arctic rivers (e.g., Dvina and Mackenzie Rivers). biology: Ripening gonads were observed in two females caught in

Dease Strait in August. The smaller (11.9 cm in standard length) had 32 orange eggs, 2.0 mm in diameter, and the larger (13.8 cm in standard length) had 38 eggs, 2.5 mm in diameter.

importance: It is not economically important. distribution: It is found in Dease Strait, Bathurst Inlet, Prince Patrick Island (Mould Bay), Amundsen Gulf, the Beaufort Sea, and the White Sea. It may also be present off major Siberian rivers. Distribution of Lycodes marisalbi

sources: Knipowitsch (1906); McAllister (1962); McAllister et al. (1981); Møller (2000b).

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biology: Its food is unknown, but several large males were caught

Lycodes mcallisteri

on long lines baited with fish. Reproduction is unknown because all known specimens were unripe.

Møller, 2001

McAllister’s Eelpout, lycode de McAllister

importance: It is not economically important.

common names: A local name is McAllisters Ålebrosme

(Danish/Greenlandic).

taxonomy: The species is named after the Canadian ichthyologist Don E. McAllister (see the dedication herein). It was originally described from western Baffin Bay, Hudson Strait, and Cumberland Sound.

distribution: The species is found in Baffin Bay, Davis Strait, Cumberland Sound, and Hudson Strait. It is probably endemic to eastern Arctic Canada and southwest and northwest Greenland.

description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 42.9%–45.8% of standard length. The head is relatively long (22.7%–26.8% of standard length). The vertebrae number 20–21 + 80–84 = 100–104; dorsal fin-rays 94–98; and anal fin-rays 80–84. The pectoral fins are rounded, without emargination, and relatively large (12.9%–17.0% of standard length), with 22–23 finrays. Other characteristic features are the two equally long rows of teeth on each palatine and a broad isthmus (greater than 5% of standard length). Squamation on the body is dense, with 28–34 scales in a vertical row from the anal fin origin to the dorsal fin base. The predorsal area, the abdomen, and the pectoral fin base are scaled. The dorsal, anal, and caudal fins are naked. The lateral-line system consists of a ventro-medio-lateral line, with the bend above the anal fin origin. A short predorsal row and a dorso-lateral row of widely spaced neuromasts are also present. The coloration is reddish brown to dark brown with six to ten white bars or patches dorsally on the body and the dorsal fin. The scales appear as white spots. The peritoneum, oral chamber, and branchial chamber are black or brown. The species reaches at least 24.4 cm in standard length (females) and 37.5 cm in standard length (males).

Distribution of Lycodes mcallisteri

sources: Møller (2001); Jørgensen et al. (2005).

habitat: It is caught at depths of 298–1,277.5 m, at temperatures of 0.2ºC–1.1ºC. Cruise data for southern Baffin Bay and Davis Strait in Canadian waters were at 530–1,045 m. It lives on muddy bottoms.

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Lycodes mucosus Richardson, 1855

Saddled Eelpout, lycode à selles

90–94; dorsal fin-rays 84–87; and anal fin-rays 65–68. The pectoral fins are moderate (12.7%–16.5% of standard length), with a low number of fin-rays (16–19). Scales are completely absent in eastern Canadian Arctic specimens but are present posteriorly on the tail in some western Canadian Arctic specimens. The lateral-line system consists of medio-lateral, predorsal, and dorsal rows of neuromasts. The opercular lobe is short, with a very tiny notch on the upper edge. The submental crests are not fused anteriorly. The abdomen and the ventral parts of the head including the cheeks are white. The dorsal parts of the head and the body are dark brown, interrupted by 3–9 irregular light bars. The border between light and dark areas is clearly delineated. The dark areas have a saddle-shaped appearance in some specimens. A light horseshoe-shaped bar on the nape is also present. Light areas may also be present ventrally on the tail and the anal fin. The peritoneum is light. The species reaches 49.0 cm in total length.

Lycodes mucosus

common names: Local names are Slimet Ålebrosme (Danish/

Greenlandic) and Kugrauna and Kuxrauna (Alaska). Other common names are Lightcheek Eelpout and Richardson’s Eelpout.

taxonomy: The species name is the Latin mucosus (slimy). The

original description is based on a specimen from Northumberland Sound, Perry Island, Arctic Canada. Some confusion over the taxonomy of L. mucosus exists, mainly due to the presence or absence of scales in different populations. Regarded as junior synonyms of L. mucosus are Lycodes coccineus Bean, 1881, described from the Bering Strait; Lycodes knipowitschi panthera Schmidt, 1950, described from the Okhotsk Sea; and Lycodes panthera (elevated to species by Matsubara, 1955, a misspelling of Lycodes knipowitschi panthera). The Sea of Okhotsk population was regarded as a separate species, L. knipowitschi Popov, 1931, by some authors.

description: It is a “short-tailed” Lycodes, and the distance from

the snout to the anal fin is 45%–61% of standard length, being longest in large males. The head is long (21%–29% of standard length), especially in large males. The vertebrae number 24–27 + 65–68 =

habitat: This is a shallow-water species, sometimes found in rock

pools. Arctic Canadian specimens were caught in depths from 3 m to 57 m, at temperatures from −1.8ºC to 7.8ºC, and at salinities from 20.0‰ to 31.5‰. Off northwest Greenland it has been reported at depths from 0 m to 180 m. This species is rarely caught in trawls, indicating a preference for shallow hard-bottom localities. It is, however, known to live in burrows in the mud in Resolute Bay, where it has been collected by scuba divers, and under rocks at Cape Hatt, northern Baffin Island.

biology: Its food includes Fourhorn Sculpins and amphipods at

Bernard Harbour. At Cape Hatt, amphipods (Anonyx nugax and Paroediceros lynceus) accounted for 91% of the prey biomass. This fish is eaten by Bearded Seals in the Canadian High Arctic.

importance: It is not economically important. distribution: The species is found in Nares Strait, sparsely on

the coast of Baffin Island, in Hudson Strait, the Foxe Basin, the Arctic islands, Dease Strait, Dolphin and Union Strait, Amundsen Gulf, and the Beaufort Sea, and from northwest Greenland to the Chukchi Sea, Bering Sea, and Sea of Okhotsk.

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Lycodes mucosus

Lycodes paamiuti Møller, 2001

Paamiut Eelpout, lycode de Paamiut

common names: A local name is Paamiuts Ålebrosme (Danish/

Greenlandic).

taxonomy: The species is named after the research vessel RV Paamiut of the Greenland Natural Resources Institute. It was originally described from Davis and Denmark Straits and the Greenland and Norwegian Seas. Specimens of this species have previously been identified as L. pallidus and L. squamiventer, which have the same type of lateral-line system. description: It is a “long-tailed” Lycodes, and the distance from

Distribution of Lycodes mucosus

sources: Johansen (1926); McAllister (1962); Finley & Evans (1983); Toyoshima (1985); Hudon (1990); Anderson (1994); Møller (1996); Møller & Jørgensen (2000).



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the snout to the anal fin is 37.1%–45.6% of standard length. The head is moderate in size (19.7%–24.8% of standard length). The vertebrae number 18–21 + 73–88 = 93–107; dorsal fin-rays 88–101; and anal fin-rays 74–89. An anterior dorsal-fin pterygiophore is associated with vertebrae numbers 4–5 (rarely 6). The pectoral fins are moderate in size (10.3%–15.2% of standard length), with 18–21 finrays. The species differs from L. pallidus by the shorter nostril tubes (0.4%–1.0% of standard length) and larger pupils (2.1%–3.6% of standard length). Squamation on the body is relatively dense, with 10–26 scales in a vertical row from the anal fin origin to the dorsal

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Lycodes paamiuti

fin base. The abdomen is scaled except for a narrow stripe ventrally; it is completely scaled in some specimens. The dorsal and anal fins are naked anteriorly but are approximately half-covered with small scales posteriorly. The caudal fin is scaled. The predorsal area and the pectoral fin base are naked. The lateral-line system consists of a ventral and a “half ” medio-lateral line from above the anal fin origin. A short predorsal row and a dorso-lateral row of widely spaced neuromasts are also present. The colour is light to dark brown, without light crossbars, being darker dorsally and paler on the ventral part. The predorsal area and the base of the dorsal fin are lighter, sometimes almost orange in appearance. The edges of unpaired fins are usually dark grey or black. Some specimens, however, are almost uniformly brownish. The scales appear as white spots. The abdomen is bluish in young and pale specimens. The peritoneum, the oral chamber, the branchial chamber, and the edge of the operculum are dark brown or black. The pectoral fins are brown, sometimes light at the base. The species reaches 22.2 cm in standard length (females) and 24.0 cm in standard length (males).

habitat: It is an Arctic slope species, with Canadian Baffin Bay

records from 519 m to 1,366 m, at temperatures from 0.1ºC to 3.4ºC. The total depth range is from 122 m to 1,450 m, and temperatures range from −1.0ºC to 4.4ºC. It lives on muddy bottoms.

biology: In the Davis Strait and Baffin Bay, ripening gonads were observed in specimens above 14.0 cm in length. In summer to autumn about 78% of the females and 90% of the males above this length were ripening. The ripest females (at 16.0 and 22.0 cm in length) had 21 and 34 yellow eggs, with diameters of 6.0 mm and 4.0 mm, respectively. In the western Davis Strait it occurs in moderate densities, up to 179 specimens per sq km.

importance: It is not economically important. distribution: It is found in Baffin Bay, Davis Strait, eastern Hudson Strait, Ungava Bay, off all coasts of Greenland, in Denmark Strait, and in the Norwegian Sea.

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This is a variable species. Several subspecies have been described: L. pallidus similis Jensen, 1904, from off Jan Mayen Island, Greenland Sea; L. pallidus var. squamiventer Jensen, 1904, now regarded as a separate species, L. squamiventer, from the Beaufort Sea, Greenland, and the Norwegian Sea; and L. pallidus marisalbi Knipowitsch, 1906, from the White Sea, recently restored as a separate species. Specimens from Arctic Canada west of the Boothia Peninsula reported as L. pallidus belong to L. marisalbi. Furthermore, specimens of a recently described species (Lycodes paamiuti) have been identified as L. pallidus in many papers.

description: It is a “long-tailed” Lycodes, and the distance from

Distribution of Lycodes paamiuti

sources: Møller & Jørgensen (2000); Treble et al. (2000); Møller (2001b).

Lycodes pallidus Collett, 1879

Pale Eelpout, lycode pâle

common names: A local name is Bleg Ålebrosme (Danish/

Greenlandic).

taxonomy: The species name is the Latin pallidus (pale) after the

pale colour of the type-specimens. It was originally described from west of Norskøyane, Spitsbergen.

the snout to the anal fin is 38.8%–48.5% of standard length. The head is moderate in size (19.6%–25.6% of standard length). The vertebrae number 18–23 + 73–86 = 93–105; dorsal fin-rays 86–97; and anal finrays 72–86. An anterior dorsal-fin pterygiophore is associated with vertebrae numbers (rarely 5) 6–9. The pectoral fins are variable (9.1%–17.8% of standard length), with 17–21 fin-rays. The species differs from L. paamiuti by the longer nostril tubes (0.8%–2.0% of standard length) and the smaller pupils (1.6%–3.2% of standard length). Squamation on the body is relatively weak, with 10–18 scales in a vertical row from the anal fin origin to the dorsal fin base. The predorsal area, the abdomen, and the pectoral fin base are naked; the dorsal and anal fins are naked or have a few scales posteriorly in large specimens. The lateral-line system consists of a ventral and a “half ” medio-lateral line from above the anal fin origin. A short predorsal row and a dorso-lateral row of widely spaced neuromasts are also present. The colour is variable: uniformly brownish or with 3–12 light bars on the dorsal fin. Occasionally there is a straight light band on the predorsal area. The scales may appear as lighter spots. The underside of the head is usually lighter than the body and the dorsal part of the head. The peritoneum is brown but appears bluish through the body in young and pale specimens. The oral and branchial chambers are murky spotted or dark. The pectoral fins are brown, sometimes light at the base. The species reaches 19.9 cm in standard length (females) and 28.7 cm in standard length (males).

habitat: Canadian Arctic records indicate this species at depths of

6–1,482 m, at temperatures from −1.6ºC to 0.7ºC. The overall depth range is 65–1,750 m, with juveniles as shallow as 6 m. The different populations have different vertical distributions, and the deep records are restricted to the southern areas of the distribution in the

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northeastern Atlantic. This species normally prefers cold water below 0ºC (to −1.8ºC), but in the mouth of Raudfjord, Spitsbergen, it was caught at a high temperature (3.7ºC). It lives on muddy bottoms.

biology: It feeds mainly on endobenthic prey, for example,

polychaetes and small crustaceans. Its burrowing habit results in ingestion of certain amounts of detritus. Small specimens also eat food of pelagic origin, for example, amphipods and copepods. The number of eggs is very dependent on the fish size. A 18.7 cm specimen from Kongsfjorden, Svalbard, had 162 yellow-white eggs, 1.0 mm in diameter; a 16.2 cm female caught off west Greenland in August had 58 eggs, 4.7 mm in diameter; whereas a 13.5 cm specimen from Raudfjord, Svalbard, only had 13 eggs. Lycodes pallidus was reported to be the most abundant zoarcid in the Svalbard archipelago and northern Barents Sea (140–400 m) with densities of up to 6,296 per sq km.

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, Hudson Strait, James Bay, near Ikpiarjuk (Arctic Bay) on northern Baffin Island, and off the Arctic islands. Records from Baffin Bay are based on cruise data, and identities are suspect for more northerly localities. It is also found off northwest and northeast Greenland, south to the Gulf of St Lawrence, in the Denmark Strait, north of Iceland and the Faroe Islands, in the Svalbard archipelago and the Barents Sea to Kara Sea.

Lycodes polaris (Sabine, 1824)

Canadian Eelpout, lycode polaire

Lycodes polaris

common names: A local name is Polar-ålebrosme (Danish/ Greenlandic). Another common name is Polar Eelpout.

taxonomy: The species name is the Latin polaris (polar). The original description is based on a specimen left by the ebb tide on the shore of North Georgia (the east side of the Boothia Peninsula) at 75° N, Arctic Canada. Lycodes agnostus Jensen, 1902, from the Kara Sea, and L. turneri atlanticus Vladykov and Tremblay, 1936, from the St Lawrence River estuary, are synonyms. The species has been confused with L. turneri Bean, 1879, from the Bering Strait. Lycodes polaris differs by the smaller maximum size (26.0 cm versus 64.0 cm total length), coloration, fewer vertebrae (88–95 versus 93–99), a narrower interorbital space (1.1%–1.4% versus 1.8%–2.4% of standard length) and a shorter isthmus (4.3%–7.3% versus 8.6%–11.5% of standard length). description: It is a “short-tailed” Lycodes, and the distance

Distribution of Lycodes pallidus

from the snout to the anal fin is 45%–55% of standard length, being longest in large males. The head is moderate in size (21%–26% of standard length). The vertebrae number 23–25 + 64–71 = 88–95; dorsal fin-rays 85–89; and anal fin-rays 65–71. The pectoral fins are small (11.1%–14.2% of standard length), with a low number of finrays (15–18). Scales are usually completely absent, but poorly scaled specimens have been reported from the Gulf of St Lawrence and are also rarely found in other populations (Kara Sea, Dease Strait, and off Labrador). The lateral-line system consists of medio-lateral, predorsal, and dorsal rows of neuromasts. The opercular lobe is short, with a very tiny notch on the upper edge. The submental crests are not fused anteriorly. The dorsal part of the body and the head is light to dark brown. A straight white band is present on the predorsal area, and 7–13 short light bands are seen on the back, passing on to the dorsal fin. The outline of the bands is usually darker than the rest of the brown area. In large specimens several light spots may be present in the dark area between the bands. The peritoneum and the ventral parts of the head and the body are uniformly light. The species reaches 26.0 cm in standard length.

sources: McAllister (1962); Hudon (1988, 1990); von Dorrien

habitat: This is a shallow-water species, sometimes even found in

(1993); Valtysson (1995); Møller (2000b, 2001b); Møller & Jørgensen (2000); Treble et al. (2000).

500

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Lycodes polaris

(usually below −1.0ºC), and salinity from 24.0‰ to 32.5‰ (usually below 30.0‰). In the eastern Canadian Arctic it is found down to 300 m, at salinities greater than 34.0‰. Off northwest Greenland it was reported at depths from 47 m to 212 m. The overall depth range of the species is from rarely 0 m, usually 4 m, to 300 m, and at temperatures almost always below 0ºC. It lives on muddy bottoms, where it buries itself tail first.

Eurasian Arctic and one of the most common fish in trawl hauls that are made deeper than 40 m in the Beaufort Sea, though mostly in the Alaskan part.

biology: Its food consists of various bottom amphipods,

polychaetes, clam siphons, brittle stars, and occasionally Arctic Cod. This fish is frequently eaten by Bearded Seals in the Canadian High Arctic. It finds its food in the top layer of the mud, where it engulfs mud and food items, and then ejects the mud with a reverse respiratory current. Its lifespan is more than five years. Spawning may occur in autumn or early winter. A large 22.7 cm female had 187 ripening eggs, 3.0–3.3 mm in diameter. Eggs up to 5.0 mm have been reported.

importance: It is not economically important. distribution: The species is found in Baffin Bay, Davis Strait, Frobisher Bay, Hudson Strait, Ungava Bay, Gulf of Boothia, the Arctic islands, Dease Strait, Bathurst Inlet, Amundsen Gulf, and the Beaufort Sea. The cruise records from deeper in Baffin Bay are suspect and require material for verification. It is almost circumpolar, from northwest Greenland to the eastern Barents Sea. The species has not been recorded from off Iceland, in the Svalbard archipelago, or along the eastern coast of Greenland. It is found in the western North Atlantic southward to the Saguenay River in the Gulf of St Lawrence. The species is one of the most common Lycodes in the



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Distribution of Lycodes polaris

sources: Sabine (1824); Leim & Scott (1966); Finley & Evans (1983); Møller & Jørgensen (2000).

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Lycodes reticulatus Reinhardt, 1835

Arctic Eelpout, lycode arctique

common names: Local names are Akuhaauk (Inuvialuktun, possibly referring to Eelpouts in general) and Sulupavak (Inuktitut); and Netmønstret Ålebrosme (Danish/Greenlandic). taxonomy: The species name comes from the Latin reticularis

(netted), referring to the reticulated colour pattern. The original description is based on two specimens from off west Greenland. Lycodes perspicillum Krøyer, 1845, described from Newfoundland; L. reticulatus hacheyi Vladykov and Tremblay, 1936, described from Hudson Bay; and L. reticulatus laurentianus Vladykov and Tremblay, 1936, described from the St Lawrence River estuary, are junior synonyms of L. reticulatus. Lycodes rossi Malmgren, 1864, might be a synonym of L. reticulatus too as the differences observed by previous authors (coloration, opercular lobe, body depth) are hard to recognize. This topic needs to be studied in more detail.

description: It is a “short-tailed” Lycodes, and the distance from

the snout to the anal fin is 44%–56% of standard length, being longest in large males. The head is moderate in size (22%–27% of standard length). The vertebrae number 22–26 + 66–75 = 90–99; dorsal fin-rays 81–92; and anal fin-rays 67–75. The pectoral fins are large (12%–18% of standard length), with 18–22 rays. Scales are present from a point at 28%–34% of standard length behind the snout. The abdomen and all the fins are naked. The lateral-line system consists of medio-lateral, predorsal, and dorsal rows of neuromasts. The opercular lobe is short. The submental crests are not fused anteriorly. The colour is variable. The dorsal part of the body and head are darker than the ventral part. A straight white band is present on the predorsal area, and 6–10 light bands are seen on the back, passing on to the dorsal fin. In most of the specimens the bands are fused into a reticulated pattern. The degree of reticulation is variable, and the bands stay separated in some specimens. The species reaches 62.0 cm in total length (males) and 53.8 cm in total length (females), or to 76.0 mm in total length generally.

habitat: An Arctic shelf species, in eastern Arctic Canada it is

recorded at depths from 68 m to 930 m, temperatures from −0.3ºC

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Lycodes reticulatus

to 2.0ºC, and salinities around 34‰–35‰. From off Labrador to Nova Scotia it occurs at depths from 51 m to 750 m, at temperatures from −1.4ºC to 3.5ºC. The overall depth range of the species is between 18 m and 930 m and at temperatures between −1.4ºC and 4.5ºC. It lives on muddy bottoms.

biology: It feeds mainly on epibenthic prey. Small specimens feed

mostly on amphipods, isopods, and bivalves, switching to shrimps, euphausiids, and fish with increasing size. In the eastern Davis Strait, ripening gonads were observed in specimens above 20.0 cm in length (females) and 23.0 cm in length (males). The most mature female (27.0 cm) had 325 yellow eggs, 4.0 mm in diameter. A specimen (28.0 cm in length), caught in the Denmark Strait in September 1994, had 208 eggs, 10.0 mm in diameter.

importance: It is not economically important. distribution: The species is found in Nares Strait, near Ikpiarjuk (Arctic Bay), in Baffin Bay, Davis Strait, Hudson Strait, Ungava Bay, Hudson Bay, Foxe Basin, the Arctic islands including the northern tip of Ellesmere Island, in Dease Strait, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and the Beaufort Sea. It is also found off Labrador and Nova Scotia in the western Atlantic and off northwest, southeast, and northeast Greenland, off Iceland, in the northern Barents Sea, off Svalbard, and in the Kara Sea and Laptev Sea. Specimens from the western Canadian Arctic were referred to L. rossi by McAllister et al. (1981), but this has to be confirmed by future studies. It has not been recorded from the Alaskan Beaufort Sea.



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Distribution of Lycodes reticulatus

sources: McAllister et al. (1981); Morosova (1982); Hudon (1990);

Valtysson (1995); Møller (1996); Møller & Jørgensen (2000); Treble et al. (2000); Jørgensen et al. (2005).

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Lycodes sagittarius McAllister, 1975

Archer Eelpout, lycode à arc

common names: None. taxonomy: The species name is the Latin sagittarius (archer), in reference to the arching (ventro-medio-lateral) type of lateral line. It was originally described from the Beaufort Sea, Alaska, and the Kara Sea, Russia. The date given in Catalog of Fishes and FishBase is 1976 (accessed 10 April 2014), but the original bears the date 1975. The taxonomy of this species is complicated. At first it was identified by authors as Lycodes squamiventer Jensen, which was then known from Greenland and Norwegian Seas. It was later described from nine uniformly dark-brown specimens as a separate species, with a relatively high number of vertebrae. Then, 26 specimens with light bars dorsally and fewer vertebrae from the Canadian part of the Beaufort Sea were identified as L. sagittarius and implemented in the diagnosis. Recently the striped specimens were assigned to Lycodes marisalbi, which means that L. sagittarius consists of uniformly brown specimens only. description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 36.7%–42.9% of standard length. The head is relatively small (18.3%–21.8% of standard length). The vertebrae number 19–22 + 77–88 = 96–109; dorsal fin-rays 92–101; and anal fin-rays 78–88. The pectoral fins are diagonal, without emargination, relatively small (9.8%–11.4% of standard length), with 16–18 fin-rays. The pelvic fins are very long (4.1%–5.6% of standard length), thin,

with only two fin-rays (three in all other Arctic Lycodes spp.). Squamation on the body is dense, with 21–27 scales in a vertical row from the anal fin origin to the dorsal fin base. The predorsal area and the abdomen are scaled. The pectoral fin base is naked, and the dorsal and anal fins are naked anteriorly and partly scaled posteriorly. The lateral-line system consists of a ventro-medio-lateral line, with the bend above the anal fin origin. A short predorsal row and a dorso-lateral row of widely spaced neuromasts are also present. The coloration is light to dark brown or with bars on the back and the dorsal fin. The scales appear as white spots. The peritoneum and the oral and branchial chambers are dark brown to black. The species reaches 24.2 cm in standard length (females) and 26.9 cm in standard length and 71.4 g in weight (males).

habitat: It is an Arctic shelf-to-slope species caught at depths

from 334 m to 600 m. The specimens from the Kara Sea were caught in cold water, −0.91ºC, with a high salinity of 34.96‰. In the Beaufort Sea it is found at 325–500 m. It lives on muddy bottoms.

biology: Its food is annelids, bivalves, gastropods, and crustaceans. The few data available suggest late-summer or early-fall spawning. A 23.8 cm standard length female contained 106 eggs, 4.2 mm in diameter. importance: It is not economically important. distribution: The species is found in the Kara Sea, the Alaskan

Beaufort Sea, and apparently in Canadian waters. A previous Canadian record is not recognized here, but there are Canadian records from the Transboundary Survey (which are mapped here; see also the “New Information” section of the introduction). Records from Davis Strait based on cruise data are presumably errors.

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Lycodes seminudus Reinhardt, 1837

Longear Eelpout, lycode à oreilles

common names: A local name is Halvnøgen Ålebrosme

(Danish/Greenlandic).

Distribution of Lycodes sagittarius

taxonomy: The species name comes from the Latin semi (half) and nudus (naked), in reference to the scale distribution. The original description is based on a specimen from west Greenland, at 71° N. A dark form, Lycodes nigricans Jensen, 1952, described from Amerdlok Fjord near Holsteinsborg, Greenland, is considered a junior synonym of L. seminudus. Lycodes nigricans was regarded as a valid species on the basis of the presence of a dorsal branch of the lateral-line system, coloration, and longer, sharper teeth. This is not confirmed, because no major differences were found when typical specimens were compared with L. nigricans type-specimens. All specimens have a weak dorso-lateral line in addition to the more pronounced medio-lateral line, which is easier to see on dark specimens.

sources: McAllister (1975a); McAllister et al. (1981); Leim & Scott

description: It is a “short-tailed” eelpout, and the distance from

(1966); Coad, with Waszczuk & Labignan (1995); Møller (2000b, 2001a).

the snout to the anal fin is 46%–55% of standard length. The body is slender with a relatively long head (23%–30% of standard length), especially in large specimens. The snout has a characteristic concave appearance. The vertebrae number 23–26 + 71–77 = 96–101; dorsal fin-rays 87–95; and anal fin-rays 69–76. The pectoral fins are short (9.5%–13.5% of standard length) with 17–22 fin-rays, the lower

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6–9 rays having very free tips. Squamation is characteristic with scales on the posterior part of the body only, ending in a wedge anteriorly at a distance of 30%–56% of standard length from the snout. There are no scales on the fins and the abdomen. The lateral line consists of medio-lateral, dorsal, and predorsal rows of neuromasts. The head pores are small and are not seen without magnification in adults. The jaw teeth are relatively long and slender. The coloration is extremely variable. Most specimens have 4–12 light bars on the body and the dorsal fin, and a straight light band connects the opercular lobes. Some are, however, uniformly brownish or greyish. This is the only species in the group of short-tailed Eelpouts that has a speckled peritoneum (light in the rest). It reaches 56.0 cm in total length (males) and 40.0 cm in total length (females).

habitat: Specimens from cruises in the eastern Canadian Arctic

were caught in depths from 118 m to more than 1,400 m. Longline records from Eclipse Sound are from 1,000–2,000 m. In the Beaufort Sea it is reported at depths from 50 m to 500 m, at temperatures of −1.6ºC to −1.2ºC, and at salinities from 28‰ to 29‰. In west Greenland waters it is known from depths of 181–1,200 m (most common between 300 m and 600 m) and temperatures of 0.5ºC–4.9ºC. The total depth range for the species is 50–1,400 m. It lives on muddy bottoms.

Distribution of Lycodes seminudus

biology: It feeds mostly on epibenthic prey. Larger food items

sources: McAllister et al. (1981); Valtysson (1995); Møller (1996);

such as fish and shrimps increase in quantity with increasing fish size. Large males are often caught on long lines. Ripening gonads were observed in specimens above 26.0 cm in length (females) and 32.0 cm in length (males) off west Greenland. About 61% of the females and 40% of the males above these lengths were ripening. Two females (34.0 and 39.0 cm) had 135 and 400 yellow eggs, with diameters of 3.7 mm and 4.1 mm, respectively. Up to 300 eggs, 8.2 mm in diameter, have been reported, ripe in June.

importance: It is not economically important. distribution: The species is found in Baffin Bay, Jones Sound, Davis Strait, Hudson Strait, Ungava Bay, Dease Strait, Amundsen Gulf, and the Beaufort Sea; and the Chukchi Borderland, northwest, southwest, and northeast Greenland, off northern Iceland to the Faroe Islands, the Svalbard archipelago, northern Barents Sea, and Kara Sea.

Møller & Jørgensen (2000); Mecklenburg et al. (2014).

Lycodes squamiventer Jensen, 1904

Scalebelly Eelpout, lycode ventre-écaillé

common names: Local names are Bugskællet ålebrosme (Danish/Greenlandic) and Nafnlausi mjóri (Icelandic). taxonomy: The species name refers to the scaled abdomen, separating it from L. pallidus, to which it is supposed to be closely related. The original description is based on specimens from north of Iceland and from the Norwegian Sea. The taxonomy and validity of this species has long been discussed. It was described as a subspecies or variant of L. pallidus, has been confused with L. marisalbi, L. paamiuti, and L. pallidus, but is now considered a valid species. New characters in dentition and the length of the nostril tube were recently discovered, supporting the validity of a separate species. description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 37%–45% of standard length. The head is small (19.2%–23.2% of standard length). The eyes are moderate and circular; the pupil diameter is 1.7%–2.7% of standard length; and the

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Lycodes squamiventer

nostril tubes are long, 1.0%–2.0% of standard length. The vertebrae number 19–21 + 77–84 = 97–103; the dorsal fin-rays 87–96; and the anal fin-rays 77–85. The first dorsal fin pterygiophore is associated with vertebrae 6–8. The pectoral fin has 17–20 rays. The pelvic fins are short, less than eye diameter, 2.4%–3.6% of standard length. The body is covered with small cycloid scales, 19–28 in a vertical row from the anal fin origin to the dorsal fin base. The number of rows increases with fish size, and the nape in front of the dorsal fin and the abdomen are scaled. The lateral-line system has short predorsal and dorso-lateral rows of neuromasts, a medio-lateral row from above the anus to near the caudal fin, and a complete ventral row. The head pores are minute with small tubes. The colour is uniformly brownish or greyish, without crossbars. The abdomen is bluish in young. The peritoneum, the oral and branchial chambers, and the edge of the operculum are darker brown. The nostril tubes are white. The species reaches 25.3 cm in total length.

habitat: It is an Arctic deep-water species. In the Canadian Arctic,

specimens from the Beaufort Sea were most common on the upper slope, and the Canadian Arctic depth range is 198–497 m. The overall depth range is 198–1,808 m, at temperatures of –1.2°C to 0.6°C, almost always below 0°C. It lives on muddy bottoms. This species is usually caught in relatively low numbers, except in the Håkon Mosby mud volcano in the Norwegian Sea where it occurs at very high densities.

biology: At the Håkon Mosby mud volcano it feeds mainly on

benthic invertebrates such as pogonophores, gastropods, amphipods, polychaetes, and copepods. About 60 eggs, 3.5 mm in diameter, have been reported in June. An age of up to 21 years has been reported based on otolith reads.

Distribution of Lycodes squamiventer

sources: Andriashev (1986); Milkov, Vogt, et al. (1999); Møller

(2001b); Gebruk, Krylova, Lein, et al. (2003); Soltwedel, Jaeckisch, Ritter, et al. (2009); Hildebrandt, Bergmann, & Knust (2011); Wienerroither, Johannesen, Dolgov, et al. (2011); Mecklenburg, Mecklenburg, Sheiko, & Steinke (2016); Majewski, Atchison, MacPhee, et al. (2017).

importance: It is not economically important. distribution: The species has been found in both the Alaskan

and the Canadian portions of the Beaufort Sea, and in the Norwegian Sea. Specimens previously reported from Davis Strait were probably all Lycodes paamiuti.



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Lycodes terraenovae Collett, 1896

Newfoundland Eelpout, lycode de Terre-Neuve

common names:  A local name is Atlantisk Ålebrosme (Danish/Greenlandic). Other common names are Atlantic Eelpout and lycode atlantique. “Newfoundland Eelpout” is used by the American Fisheries Society (see the bibliography) for L. lavalaei because L. terraenovae does not occur in their list. The names used here seem more appropriate. taxonomy: The species name refers to the type locality and dis-

tribution area. The original description is based on specimens from the Newfoundland Banks. The taxonomy and validity of this species has been questioned by several ichthyologists and was cleared up only recently. The names L. agulhensis Andriashev, 1959, described from South Africa; L. atlanticus Jensen, 1902, described from off Virginia; L. atratus Vladykov and Tremblay, 1936, described from the St Lawrence estuary; and L. brunneus Fowler, 1944, described from north of the Bahamas, are regarded as junior synonyms. It is suggested to be a close relative of Lycodes esmarkii and L. eudipleurostictus.

description: It is a “long-tailed” Lycodes, and the distance

from the snout to the anal fin is 33.5%–45.9% of standard length. The head is small (17.7%–24.0% of standard length). The number of fin-rays and vertebrae declines considerably with latitude. The vertebrae number 20–23 + 84–102 = 105–124; dorsal fin-rays 100–118; and anal fin-rays 86–104. The specimens from the Davis Strait reach the highest counts in the genus. The pectoral fins are rounded and of moderate length (9.4%–13.8% of standard length), with 20–23 fin-rays. Squamation is dense and covers the predorsal area, the abdomen, the dorsal and anal fins, and the pectoral fin base. The lateral-line system consists of a well-developed ventral line, in addition to a medio-lateral row of widely spaced neuromasts. A dorsal row is also present, but the predorsal row

is missing. The northern populations of this species are characterized by the few palatine teeth (2–6), which is less than those of all other Lycodes in the region. Related species such as L. esmarkii, L. eudipleurostictus, and L. vahlii have 5–13, 10–18, and 8–22 teeth on each palatine, respectively. The coloration is uniformly light to dark brown, never with bars or reticulations. The opercular membrane, the peritoneum, and the fins are black. Juvenile specimens (less than 15.0 cm in length) are greyish blue to light brown, with a darker head, abdomen, and fins. The species reaches 51.0 cm in total length.

habitat: It is an Atlantic deep-water species. A single Canadian

Arctic specimen from southern Davis Strait was caught in a depth of 840–870 m. The overall depth range is 150–2,600 m, at temperatures of 2.0ºC–5.0ºC. It lives on muddy bottoms. Although in low densities in the northern part of its distribution area, it was found to be one of the most numerous species on the continental slope off Cape Hatteras.

biology: It feeds on endobenthic (infaunal) prey such as sponge

remains, polychaetes, shelled molluscs, pycnogonids, crustaceans, and brittle stars. Large amounts of sediment in the stomachs indicate that L. terraenovae gulps down sediment along with the food. One specimen was reported to have ingested a piece of coal. In the southern Davis Strait, ripening gonads were observed in specimens above 35.0 cm in length (females) and 38.0 cm in length (males). A large 40.0 cm total length female, caught in September, had 320 orange eggs, about 4.0 mm in diameter.

importance: It is not economically important. distribution: The species has been found from southern Davis Strait as a single record at 64°03' N, 58°42' W (ZMB 22642), with many cruise records in the Canadian Baffin Bay, Davis Strait, and Hudson Strait being regarded as errors and probably L. paamiuti. It is also found off southwest and southeast Greenland, southward to off Florida in the western Atlantic, and from off Ireland to off South Africa in the eastern Atlantic.

Lycodes terraenovae

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Lycodes vahlii Reinhardt, 1831

Checker Eelpout, lycode à carreaux

Lycodes vahlii

common names: A local name is Vahls Ålebrosme (Danish/

Greenlandic). Other common names are Vahl’s Eelpout and lycode de Vahl.

taxonomy: The species is named after the collector Martin Hen-

Distribution of Lycodes terraenovae

sources: Andriashev (1959); Karrer (1973); Sedberry & Musick (1978); Wenner (1987); Hecker (1994); Møller (1997); Møller (2000a); Møller & Jørgensen (2000).

driksen Vahl (1749–1804), a Danish naturalist who sent several different specimens of fish to zoologist J.C.H. Reinhardt. It was originally described from a specimen removed from the stomach of a Greenland Shark, caught off Qaqortoq/Julianehåb, west Greenland. Until recently two subspecies were recognized: L. vahlii vahlii Reinhardt, 1831, described from the northwestern Atlantic; and L. vahlii gracilis Sars, 1866, described from the northeastern Atlantic. Lycodes gracilis was restored on the basis of several characters, for example, meristics, colour, and teeth. Lycodes zoarchus Goode and Bean, 1896, described from off Nova Scotia, and L. vahlii maculatus Vladykov and Tremblay, 1936, described from the St Lawrence River estuary, are synonyms of L. vahlii.

Lycodes vahlii



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importance: It is not economically important. distribution: It is found in Davis Strait, Cumberland Sound,

Hudson Strait, Ungava Bay, off Resolution Island, in northwest and southwest Greenland (between 59°32' N and 73°34' N), northern Labrador Sea, off Labrador, in the Gulf of St Lawrence, to the northern Scotian Shelf. Northeastern Atlantic records belong to a different species, L. gracilis Sars, 1867.

Lycodes vahlii

description: It is a “long-tailed” Lycodes, and the distance from

the snout to the anal fin is 35.6%–44.5% of standard length. The head is moderate in size (15.5%–24.0% of standard length). The vertebrae number 23–27 + 89–98 = 114–122; dorsal fin-rays 95–113; and anal fin-rays 92–97. The pectoral fin is relatively small (8.8%–14.6% of standard length), with 18–20 fin-rays. Squamation on the body is relatively dense, with 19–30 scales in a vertical row from the anal fin origin to the dorsal fin base. The teeth are blunt. The predorsal area and the abdomen are scaled. The dorsal and anal fins are naked anteriorly but have small scales posteriorly. The pectoral fin base is naked. The lateral-line system consists of a well-developed ventral line and of a predorsal row and a dorso-lateral row of widely spaced neuromasts. The colour is olive green to dark brown, lighter ventrally, with 5–12 dark bands along the sides of the body and the dorsal fin. The anterior bands are often more pronounced. A few specimens are uniformly brownish without bars. Scales appear as white spots. The abdomen is bluish in young and pale specimens. The peritoneum, oral chamber, and branchial chamber are brownish. The species reaches 40.0 cm in total length (females) and 57.0 cm in total length (males).

Distribution of Lycodes vahlii

sources: Reinhardt (1831); Morosova (1982); Hudon (1990); Møller (2000a); Møller & Jørgensen (2000); Carl (2002).

habitat: It is an Arctic boreal shelf-to-slope species, with Can-

adian Arctic records from Ungava Bay at depths of 322–391 m, and from Davis Strait and Labrador Sea at depths of 202–1,200 m, with temperatures of 4.2ºC–4.4ºC. The total depth range is rarely from 39 m, more usually from 71 m, to 1,200 m, and at temperatures from 1.4ºC to 5.8ºC. It lives on muddy bottoms.

biology: Its food is unknown (only studied for L. gracilis, which

is now considered a separate species). The relatively powerful blunt teeth indicate a molluscan diet. Ripening gonads were observed in specimens above 14.0 cm in length (females) and 17.0 cm in length (males) off west Greenland. In summer and autumn about 15% of the females and 20% of the males above these lengths had ripening gonads. In 12 females (15.0–22.0 cm in total length) the number of eggs varied from 20 to 75, with diameters from 3.0 mm to 7.0 mm. The species is one of the most abundant Eelpouts in west Greenland waters, with densities up to 7,618 specimens per sq km.

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Lycodonus mirabilis Goode and Bean, 1883

Chevron Scutepout, lycaspine à chevrons

common names: A local name is Vestatlantisk Pladebrosme (Danish/Greenlandic). taxonomy: The genus comes from Lycodes with a meaningless change in the end of the name. The species name is the Latin mirabilis (wonderful). It was originally described from the Atlantic Ocean (38°20'08" N, 73°23'20" W). The species has been confused with the eastern North Atlantic relative Lycodonus flagellicauda (Jensen, 1902) in some literature. Lycodonus mirabilis differs by the higher number of vertebrae

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Lycodonus mirabilis

(115–119 versus 96–103), the anterior dorsal bony plates lacking finrays (present in L. flagellicauda), and a shorter head (9%–12% versus 11%–14% of standard length).

description: The body is elongated, rounded, and eel-like, and

the distance from the snout to the anal fin is 22.9%–24.6% of standard length. The head is small (9.9%–11.6% of standard length). The vertebrae number 22–23 + 93–97 = 115–119; dorsal fin-rays 100–105; and anal fin-rays 96–100. There are five branchiostegal rays (six in all other Canadian Arctic zoarcids). The most characteristic feature is the bony plates (scutes) along the dorsal and anal fin bases. However, the anterior 6–12 dorsal bony plates lack fin-rays (all bony plates have fin-rays in L. flagellicauda). The bony plates give the Scutepout a very rigid body compared to its Lycenchelys relatives. The pectoral fins are relatively small (6.2%–7.7% of standard length) with 15–17 fin-rays. Tiny scales cover the sides of the body and the abdomen, 25–26 in a vertical row above the anal fin origin. The dorsal and anal fins, the pectoral fin base, and the predorsal area are naked. The lateral-line system has ventral, medio-lateral, and dorsal rows of neuromasts. The head pores are very large. The coloration is uniformly grey to dark brown, and the abdomen is bluish in juveniles. Scales and pigmentation on the sides are often missing due to contact with trawls, giving a light appearance. The peritoneum and the pectoral, dorsal, and anal fins are uniformly dark brown. The species reaches 31.0 cm in total length.

observed in specimens above 22.0 cm in length in the eastern Davis Strait in September. In nine females (24.0–30.0 cm) the number of orange-red eggs varied from 32 to 88, with diameters from 3.1 mm to 4.8 mm.

importance: It is not economically important. distribution: It is found in southern Baffin Bay and Davis Strait. A northwest Atlantic deep-water, slope species, it is known from northwest and southwest Greenland and south to off North Carolina.

habitat: In the Canadian Arctic (western Davis Strait, 66º08' to

67º47' N) it was caught in depths from 1,282 m to 1,397 m, at temperatures from −0.1ºC to 0.3ºC. Off Baffin Island it has been caught at 964 m and 3.5ºC, and from cruise data in southern Baffin Bay and Davis Strait at 271–771 m. Off west Greenland it is recorded between 63º04' N and 73º12' N, at depths from 424 m to 1,433 m, and at temperatures between 0.5ºC and 4.5ºC. The greatest abundance was observed south of 66º N at temperatures between 3.0ºC and 4.5ºC. Further south it reaches depths down to 2,394 m. It lives on muddy bottoms.

biology: Its feeding biology has only been studied briefly. In

Distribution of Lycodonus mirabilis

sources: Wenner (1978); Møller & Jørgensen (2000); Treble et al. (2000).

Norfolk Canyon it eats bivalves (Nuculanidae), cumaceans, and polychaetes. A lot of sediment is swallowed. Ripening gonads were



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Melanostigma atlanticum Koefoed, 1952

Atlantic Soft Pout, mollasse atlantique

common names: A local name is Blødkvabbe (Danish/ Greenlandic).

taxonomy: The genus comes from the Greek melas (black) and stigma (spot). The species name comes from the Latin atlanticus (of the Atlantic). It was originally described from the North Atlantic Ocean (57°41' N, 11°48' W).

to 75 large orange eggs of up to 3.9 mm in diameter are guarded. This fish is eaten by other fishes, such as redfishes in the Gulf of St Lawrence.

importance: It is not economically important. distribution: It is found in the Davis Strait without a precise locality (the map spot is an estimate). This is an Atlantic deep-water species, known from southwest and southeast Greenland to the Gulf of St Lawrence and south to off Virginia; and from the Faroe-Iceland Ridge to off Mauritania including the Mediterranean in the eastern Atlantic.

description: The body is elongated and gelatinous, and the distance from the snout to the anal fin is 28%–32% of standard length. The head is small (12%–15% of standard length). The vertebrae number 19–20 + 74–77 = 94–96; dorsal fin-rays 87–88; and anal fin-rays 71–72 (higher literature counts include half of the confluent caudal fin). The most characteristic features are the lack of pelvic fins, a pore-like gill opening, and a terminal mouth. Males have fanglike teeth on the jaws and vomer. The pectoral fins are very small (4%–5% of standard length) with 6–8, rarely 9, fin-rays. Scales and a lateral line are absent. The head pores are small. The colour is pale, transparent. The snout, lips, gill pore, anal opening, and posterior part of the tail are black; the operculum and abdomen are brilliant silvery blue. The species reaches 16.0 cm in total length. habitat: This pelagic species has been reported from just south

and east of the Canadian Arctic area (western Davis Strait, between 63º14' N and 65º04' N at depths from 705 m to 1,199 m and temperatures between 3.0ºC and 3.9ºC). Off the Atlantic coast of Canada it is most common between 276 m and 366 m. It has, however, been reported down to 1,853 m.

Distribution of Melanostigma atlanticum

biology: It feeds on copepods, ostracods, and euphausiids.

Males are larger than females at sexual maturity, with means of 13.7 cm and 12.6 cm length, respectively. It reproduces in July to September, in 15.0–32.0 cm deep burrows in the sediment, where up

sources: Koefoed (1952); McAllister & Rees (1964); Wenner

(1978); Markle & Wenner (1979); Silverberg, Edenborn, Ouellet, & Béland (1987); Anderson (1994); Møller & Jørgensen (2000).

Melanostigma atlanticum

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Zoarces americanus

(Bloch and Schneider, 1801) Ocean Pout, loquette d’Amérique

common names: Other common names include Congo Eel, Eelpout, Laughing Jack, Mother-of-eels, Muttonfish, and Poodler. taxonomy: The genus comes from the Greek zoarkes (life-supporting or viviparous). The species is named after America, from where it was first described. Until recently this species was placed in its own genus, Macrozoarces Gill, 1863, but the differences from the other species of Zoarces are now considered too small to warrant generic status of the western Atlantic species. description: This species differs from other Canadian Arctic

zoarcids by the pronounced notch of spiny fin-rays in the posterior part of the dorsal fin. The body is elongate, the tail being longer than the pre-anal region; the pre-anal length is 34%–37% of standard length. The vertebrae number 25–28 + 103–119 = 129–146; dorsal fin-rays 92–103, followed by 16–24 short spines, and then 16–31 soft rays; and anal finrays 105–124. The pectoral fins are rounded and fleshy, with 18–21 fin-rays. Squamation is dense, covering the abdomen and the posterior part of the dorsal and anal fins. There are no scales on the predorsal area and the pectoral fin base. The lateral-line system consists of a well-developed medio-lateral line, a short predorsal row, and a dorsal

row of wide-spread neuromasts. The head is relatively short, with a blond snout. The jaws are strong with large blunt teeth. The palatine and vomerine teeth are missing. The coloration is yellow to brown with grey to green mottling, fading to a lighter belly. The dorsal fin is darker than the anal fin and has a yellow margin. The rear of the dorsal fin and the anal fin can be an orange red. The pectoral fins are red or orange. The peritoneum is light. Males grow larger than females. The species reaches 1.18 m in length and more than 6.0 kg in weight.

habitat: It inhabits all types of bottom but is more numerous on

rocky, hard-bottom localities. It is a shallow-water fish, with a seasonal migration into deeper water in the fall and to shallower water in the spring. The Canadian Arctic specimens from the Hudson Strait were caught in a depth of 313–388 m. The overall depth range is 1–388 m, at temperatures of 0ºC–16.7ºC, but preferred depths of 55–108 m, temperatures of 6ºC–9ºC, and salinities of 32‰–34‰ have been reported for the Scotian Shelf population.

biology: Ocean Pout food consists of a wide variety of inverte-

brates and occasionally small fish. The robust blunt teeth are able to crush hard food items such as sea urchins, brittle stars, sand dollars, crabs, barnacles, mussels, whelks, periwinkles, scallops, sea squirts, and other molluscs. Polychaetes and amphipods are also taken. Laboratory observations on the feeding of the Ocean Pout have shown that food was extracted from mouthfuls of sediment. In 0-group juveniles, epiphytic harpacticoid copepods form the most important prey type. Ocean Pout have been reported from the stomachs of Barndoor Skates, Longhorn Sculpins, Sea Ravens, Atlantic Cod, and Harbour Seals.

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Zoarces americanus

Growth is slow. In the Bay of Fundy a 5-year-old fish measured 31.0 cm, a 12- to 13-year-old measured 55.0–58.0 cm, and a 16- to 18-year-old measured 64.0–68.0 cm. Growth is more rapid further south and may be slower in the Arctic area. The Ocean Pout has antifreeze proteins (type III) in its blood. It is the only egg-laying species in the genus, the three Eurasian species being viviparous. Ocean Pout form pairs in the spring, and copulatory behaviour has been observed, indicating internal fertilization prior to spawning. Spawning occurs in crevices and holes under boulders, where the egg masses are guarded by the female. A female may produce up to 4,200 eggs, the number increasing with the size of the fish. The yellow eggs measure 6.0–9.2 mm in diameter and hatch two to three months after deposition. Spawning occurs in late August in Newfoundland and later further south. Feeding by the female ceases prior to spawning.

importance: There has been no fishery for Ocean Pout in Canadian waters in recent years. Large quantities were caught as food fish during the Second World War, but the commercial interest has been low since then, mainly due to a protozoan parasite that caused lesions. It is still caught off New England, mainly by trawl between December and May. It is included in the New England Fishery Management Council’s Multispecies Fishery Management Plan under the “nonregulated multispecies” category. An industrial fishery by the U.S. fleet from 1964 to 1974 landed 4,700 t on average annually. The catches declined to about 600 t annually during 1975 to 1983 but increased to 1,500 t in 1984–5 due to a small directed fishery in Cape Cod Bay supplying the fresh fillet market. The 1996 catch by the United States was only 51 t, the lowest since 1963. Fisheries biologists assume that the northeastern U.S. population is over-exploited at a low biomass level. The Ocean Pout is extensively used in physiological and genetic studies, especially those related to heart and skeletal muscles and anti-freeze proteins. distribution: It is found in Davis and Hudson Straits and Ungava Bay. Endemic to the western North Atlantic, it is found south to off North Carolina. The Canadian Arctic records represent

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a considerable range extension from the earlier northern limit at Battle Harbour, southern Labrador. They stem from a fisheries survey, where the specimens were discarded. Thus, their presence in Arctic Canada needs to be confirmed by future records, preferably of specimens entering museum collections.

Distribution of Zoarces americanus

sources: Clemens & Clemens (1921); Olsen & Merriman (1946);

Miller & Jørgensen (1973); Margolis & Arthur (1979); Scott (1982); Macdonald (1983); Keats, South, & Steele (1985); Keats & Steele (1993); Mercer, Brown, Clearwater, & Yao (1993); Anderson (1994); Wigley (1998); Fletcher, Goddard, & Wu (1999).

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“Blennies,” a term including members of this family, are eaten by Thick-billed Murre chicks at Akpatok Island and by ciscoes in James Bay estuaries. Pricklebacks in the Grande Baleine estuary of James Bay produce fewer and larger larvae after ice break-up that feed on larger food items, in contrast to the strategy of the numerous small Sand Lance larvae that hatch before ice break-up and feed on small prey.

Family Stichaeidae Pricklebacks, Stichées

Brian W. Coad

sources: Jensen (1944); Makushok (1958); Mecklenburg & Sheiko (2004).

Acantholumpenus mackayi (Gilbert, 1896)

Blackline Prickleback, terrassier à six lignes The Pricklebacks or Shannies are mostly found in the North Pacific Ocean, with a few species in the North Atlantic Ocean and the Arctic Ocean. There are about 76 species, including 23 species in Canada, of which 8 are in Arctic waters. The maximum length is about 76 cm, although most are smaller. The name “Prickleback” is derived from the long dorsal fin covering most of the back, which is composed wholly of spines in most species (22–127 spines, 0–82 soft rays). The anal fin is also long with 1–5 spines at its origin and 20–102 soft rays. The anal fin is longer than the distance from the snout to the anal fin origin; the similar Gunnels (Pholidae) have a shorter anal fin and a longer pre-anal distance. Gunnels are usually larger than Pricklebacks. Both the dorsal fins and the anal fins may be joined to the caudal fin but are usually separate. The pelvic fins are small, thoracic (just in front of the pectoral fin base), and have 1 spine and 2–4 soft rays. Pelvic fins may be absent. There is a long, compressed, and small body and a small mouth. Some species have long and numerous cirri or “tentacles” on the head and long anterior dorsal fin spines, and some species have a dermal crest on the head. The lateral line is often indistinct and can be single, or multiple with complex branching. The scales are very small and cycloid and may be absent. Pricklebacks are common in tide pools and shallow waters where there are algae, and may occur to 250 m or more. They eat small invertebrates.

common names: Other common names are Pighead Prickleback and Spiny Eelblenny.

taxonomy: The genus comes from the Greek akantha (spine or thorn) and Lumpenus (see the taxonomy of Lumpenus fabricii). The species is named after Charles Lesley McKay (1855–83), a young ichthyologist who drowned at the type locality. description: This species is distinguished by its restricted distribution and a series of characters. The snout has a fraenum; finger-like, lower pectoral fin-rays are absent, as are head cirri; a single complete lateral line is present; the dorsal fin is low anteriorly, its height rising gradually; there are 59–76 dorsal fin spines; and there are 2 anal fin spines with 41–48 soft rays. The dorsal and anal fin-rays are longest well posterior to their origins. The pectoral fin-rays number 13–16. The caudal fin is pointed. The pointed snout projects beyond the upper lip. The body is yellowish to yellowish brown or olive. There is a solid black line along the base of the dorsal fin and two dashed lines or blotches on the flank. The upper of these two lines may form a reticulate pattern. The top of the head has a fine reticulate pattern. The pectoral fins are orange. The caudal fin is dark with a light posterior margin. The roof of the mouth

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Acantholumpenus mackayi

is black. The lower flank is unmarked as are the caudal and anal fins. The species attains 70.0 cm in total length.

habitat: Its depth range is from the shoreline to 60 m, perhaps

to 200 m, on mud, sand, and gravel bottoms. Temperatures up to 9.25°C are recorded for this species in Arctic Canada. This Prickleback is benthic and does not school. It does not appear to be abundant in Canadian waters. In Asia it enters brackish water and is found in the mixed estuarine zone of the Mackenzie River. It favours the deeper, relatively warmer, winter waters of Tuktoyaktuk Harbour and the deeper inlets of Liverpool Bay.

biology: This species may live as long as 16 years, but almost

nothing is known of its life history in Canada. In the Tuktoyaktuk region, polychaete and oligochaete worms, amphipods, cumaceans, and copepods dominate in the diet, and fish eggs, fish, pelecypods, gastropods, and isopods are also eaten. Females taken in Tuktoyaktuk Harbour in August were nearly ripe with eggs up to 1.4 mm in diameter. Maturity in Japan is attained at 30–40 cm in length.

importance: The Committee on the Status of Endangered Wildlife in Canada assessed it as “Data Deficient” in May 2003, while the Canadian Species at Risk Act (SARA) lists it as of “Special Concern.” distribution: The species is found in the Canadian Beaufort Sea (but not the Alaskan Beaufort Sea), the southern and eastern Bering Sea of Alaska, and the Sea of Japan to the Sea of Okhotsk.

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Distribution of Acantholumpenus mackayi

sources: Tokuya & Amaoka (1980); Shchetinnikov (1983); Lawrence & Crawford (1987); Houston & McAllister (1990b); Lacho (1991).

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Anisarchus medius (Reinhardt, 1837)

Stout Eelblenny, lompénie naine

common names: Local names are Shalup-pau-gah; and Nordlig Langebarn (Danish/Greenlandic).

taxonomy: The genus comes from the Greek anisos (unequal)

and archos (anus). The species name is the Latin medius (middle). It is placed in the genus Lumpenus by some authors.

description: This species is distinguished by a combination of

fin-ray counts and other characters. There are 58–63 dorsal fin spines, 37–43 soft anal fin-rays with posterior rays longer, and 13–15 (usually 13–14) pectoral fin-rays; a fraenum and finger-like, lower pectoral fin-rays are absent, and there is a single complete lateral line. The upper jaw does not project. Scales cover the body, but the lateral line is indistinct. The dorsal and anal fins are attached to the caudal fin. The overall colour is light brown to yellowish with a few light flank blotches or 12–14 subcircular spots with yellow or cinnamon-coloured dusting. The back has vague cinnamon-coloured spotting. The dorsal fin has blood-red oblique bars. There is an irregular black spot behind the eye. The caudal fin usually has several bars, up to eight in number. The belly and underside of the head are milky white. In life the body is translucent and pinkish with the silvery peritoneum showing through the body wall. The species reaches about 18.0 cm, perhaps 30.0 cm, in total length.

habitat: It is found on soft bottoms of sand, mud, and clay, occasionally on stones or gravel in Arctic Canada. Its depths are 15–143 m in Arctic Canada, somewhat deeper elsewhere at 10–300 m. On the Labrador coast it is found at temperatures as low as −1.34°C and generally not much higher than 5°C. In Arctic Canada it is found from −1.6°C to 3.0°C, and elsewhere it is recorded down to −1.84°C. biology: Its food is crustaceans, polychaete worms, and bivalve

molluscs. Larval and post-larval fish feed on copepods such as Calanus glacialis, Jaschnovia tolli, Limnocalanus macrurus, Microcalanus pygmaeus, Oithona similis, Pseudocalanus minutus, Tisbe furcate; on larvaceans such as Fritillaria borealis and Oikopleura vanhoeffeni; on diatoms such as Coscinodiscus sp.; and on eggs in the Beaufort Sea. Atlantic Cod are known to eat this species, as are, in the Alaskan Beaufort Sea, Arctic Char. In Digges Sound, Hudson Bay, this species is food for Black Guillemot and Thick-billed Murre chicks, and it is eaten by Brünnich’s Murres at Akpatok Island in Ungava Bay. In Frobisher Bay in early August, eggs are up to 2.1 mm in diameter.

importance: It is not economically important. distribution: This species is circumpolar, found across Arctic Canada, particularly at southern Baffin Island, Frobisher Bay, eastern Hudson Strait and Ungava Bay, sparsely in eastern Hudson Bay, and in Bathurst Inlet, Dease Strait, Amundsen Gulf, and the Beaufort Sea. General reports for James Bay are not confirmed by specimens. It is also found from northwest, southwest, and southeast Greenland to as far south perhaps as Cape Breton Island, Nova Scotia, in Atlantic Canada; in the Alaskan Beaufort Sea; the Chukchi and Bering Seas, and the Seas of Okhotsk and Japan.

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Chirolophis ascanii (Walbaum, 1792)

Atlantic Warbonnet, toupet marbré

common names: Another common name is Yarrell’s Blenny. taxonomy: The genus comes from the Greek cheir (hand) and lophos (crest). The species is named after Peter Ascanius (1723– 1803), the Norwegian naturalist and student of Linnaeus. description: This is the only Arctic Prickleback with obvious

cirri. It has 50–54 dorsal fin spines, 35–40 anal fin-rays, and a large cirrus over each eye, with a smaller one in front. There are various other small cirri on the head and the anterior dorsal fin spines. The body is yellowish brown, has bright red-brown lines, and is darkly blotched or banded. A dark ring around the eye continues ventrally as a cheek bar. It attains 25.0 cm in standard length.

habitat: The Canadian Arctic record is of small, young fish. The Distribution of Anisarchus medius

sources: Tuck & Squires (1955); Edwards (1961); McCart (1980);

Gaston, Cairns, Noble, et al. (1981); Gaston, Cairns, Elliot, et al. (1985); Hudon (1990a); Nielsen et al. (1992).

species is found generally in rocky areas among seaweed from 10 m to 400 m, usually at shallower depths to 30 m. It is not intertidal. The fry are pelagic.

biology: Its food is polychaete worms, molluscs, hydroids, sponges, and algae. In Europe, spawning occurs in October and November, and eggs (up to 2.8 mm in diameter) are deposited in small flattened masses on stones.

importance: It is not economically important. distribution: The record from Kimmirut (Lake Harbour), Baffin Island, by Dunbar (1947) was based on seven specimens 12–14 mm long taken in 1939–40 and listed as Chirolophis ?galerita (sic). Post-larvae of 16.5 mm lack cirri, and identification of this species is tentative. This record has not been confirmed by further material. It is found in the Gulf of St Lawrence and off southeastern Newfoundland, as well as in the eastern North Atlantic Ocean.

Chirolophis ascanii

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Eumesogrammus praecisus (Krøyer, 1837)

Fourline Snakeblenny, quatre-lignes atlantique

Eumesogrammus praecisus

common names: A local name is Firliniet Slimfisk (Danish/

Greenlandic).

taxonomy: The genus comes from the Greek eu (well, good),

mesos (middle), and gramme (line), in reference to the middle lateral line being the longest. The species name is the Latin praecisus (exact). Clinus unimaculatus Reinhardt, 1836, described from southwest Greenland, is a synonym.

Distribution of Chirolophis ascanii

sources: Dunbar (1947a); Russell (1976).

description: The relatively deep, compressed body and four lateral lines (one above and two below a complete mid-flank lateral line) distinguish this species from other Arctic Pricklebacks. The dorsal fin has 47–50 spines, and the anal fin has 1–2 spines and 30–35 soft rays, with the last 3 rays spine-like. The large pectoral fins have 16–19 rays. The overall body colour is brown to grey, lighter

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below, and yellowish, with vague broad bars on the flanks. There are three dusky bars across the cheek. A large black spot with white margins is on the anterior part of the dorsal fin between spines 8–11, sometimes with 1–2 similar smaller spots behind or in front. The anal, caudal, and pectoral fins are dark with a white margin. The caudal fin base is pale or yellowish. The species attains 23.0 cm in total length.

sources: Johansen (1927a); Hildebrand (1948); Dunbar & Hildebrand (1952); Tuck & Squires (1955); Gaston, Cairns, Noble, et al. (1981); Cairns (1982, 1987a, 1987b); Gaston, Cairns, Elliot, et al. (1985); Morrison & Gaston (1986); Gaston (1987, 1989a, 1991); Gaston, Elliot, et al. (1987).

habitat: It is a poorly known species found over mud, sand, gravel,

pebbles, stones, and rock and associated with seaweeds, at depths of 18–33 m in Labrador for example, but they have been taken from the tidal zone down to 400 m. Water temperatures may be as low as −1.34°C on the Labrador coast and at least −1.18°C in Arctic Canada.

biology: Its food is probably crustaceans and other small bot-

Leptoclinus maculatus (Fries, 1838)

Daubed Shanny, lompénie tachetée

tom organisms. This species is food for Atlantic Cod and Arctic Cod in Ungava Bay, for Black Guillemots and Thick-billed Murre chicks at Digges Sound, Hudson Bay, for Thick-billed Murre chicks at Coats Island, for Black Guillemots chicks in the Nuvuk Islands and at Pitsulak and Piqiuliit in northeastern Hudson Bay, and for Brünnich’s Murres at Akpatok Island in Ungava Bay. Spawning may occur in late summer in the Arctic as nearly ripe ova and planktonic young have been found in Ungava Bay in late August.

Leptoclinus maculatus

importance: It is not economically important.

common names: A local name is Plettet Langebarn (Danish/

distribution: The species is found in Cumberland Bay, south-

ern Davis Strait, Hudson Strait, Ungava Bay, Hudson Bay (sparsely), Dease Strait, Melville Sound, and Amundsen Gulf. It is found south from northwest and southwest Greenland to the Gulf of St Lawrence in Atlantic Canada, and from the Alaskan Beaufort Sea, Chukchi Sea, Bering Sea, and Sea of Okhotsk.

Greenlandic). Another common name is Spotted Snakeblenny.

taxonomy: The genus comes from the Greek leptos (slender or

thin) and the unrelated tropical genus Clinus from klino (slope) and kline (bed), apparently in reference to the osteology of that genus. The species name is the Latin maculatus (spotted). It is often placed in the genus Lumpenus. The date 1838 is from Catalog of Fishes. Clinus aculeatus Reinhardt, 1837, described from Greenland, is a synonym and cannot be used even though it was of earlier date, because it is on the Official Index of Rejected and Invalid Works in Zoological Nomenclature.

description: This species is distinguished by fin-rays counts (1–2

anal fin spines and 34–44 soft rays; 57–64 dorsal fin spines) and by the lower 5–6 rays of the pectoral fin being extended and finger-like. The pectoral fin-rays number 14–16. Scales cover the body and the sides of the head except the cheeks, but the lateral line, though complete, is not evident. The snout protrudes over the mouth. The overall colour is grey green to yellowish or reddish brown, or even whitish, with irregular and rounded darker blotches on the lower flanks and back, and a silvery belly. There are about 2–5 bars on the caudal fin, about 10–12 oblique bars on the dorsal fin, and about 5 bars on the pectoral fins. The species reaches 23.0 cm in total length.

habitat: It is found on hard, sandy bottoms usually and also on

Distribution of Eumesogrammus praecisus

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mud, gravel, pebble, or rock down to 400 m, but as shallow as 2 m in the subtidal zone, and as deep as 454–607 m in Hudson Strait and to 710 m in Davis Strait. It is found on the Labrador coast at temperatures down to −1.42°C, in Arctic Canada down to −1.6°C, and elsewhere as high as 11.5°C. The mode and type of lipid storage

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Leptoclinus maculatus

in adult tissues is an adaptive strategy shown by this fish for life in high latitudes and cold water, and lipid accumulation and distribution in gonads is transferred to embryos and larvae for their optimal development. The free lower rays of the pectoral fins are used to support the body on the sea bed and perhaps are used in movement.

biology: Its food is polychaete worms and pelagic amphipods. It

is eaten by a wide range of other fish species such as Atlantic Cod in Ungava Bay; by Thick-billed Murre chicks in Digges Sound and at Coats Island, Hudson Bay; by Black Guillemots at Pitsulak and Piqiuliit in northeastern Hudson Bay; and by Brünnich’s Murres at Akpatok Island in Ungava Bay. Arctic Char also prey on this species. Sexual maturity is attained at about 12.5 cm in length. Spawning is believed to occur in winter with up to 970 eggs being produced. Fry are pelagic.

importance: It is not economically important. distribution: The species is found in Jones Sound in the Arctic islands, near Ikpiarjuk (Arctic Bay) on northern Baffin Island, in Baffin Bay, Davis Strait, Frobisher Bay, Hudson Strait, Ungava Bay, northern Hudson Bay, southeastern Hudson Bay and James Bay (rarely), northern Foxe Basin, Dease Strait, Amundsen Gulf, and the Beaufort Sea. It is also found in the Alaskan Beaufort Sea, Chukchi Sea, East Siberian Sea, Kara Sea, the North Pacific Ocean in the Bering Sea and south to Washington State, and in the Seas of Okhotsk and Japan; and from all coasts of Greenland and in the North Atlantic Ocean south to Cape Cod in the west.



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Distribution of Leptoclinus maculatus

sources: Dunbar & Hildebrand (1952); Tuck & Squires (1955); Gaston, Cairns, Noble, et al. (1981); Gaston, Cairns, Elliot, et al. (1985); Cairns (1987a, 1987b); Gaston, Elliot, et al. (1987); Gaston (1989a, 1991); Murzina et al. (2013).

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Lumpenus fabricii Reinhardt, 1836

Slender Eelblenny, lompénie de Fabricius

Lumpenus fabricii

common names: Local names are Fabricius Langebarn (Danish/

Greenlandic); Hutdaun; and Tejernak (Greenlandic). Another common name is lompenie élancée.

taxonomy: The genus comes from the Danish word “lumpen,” the

name for the European Zoarces viviparus with which these fishes were once confounded. The species is named after Otto Fabricius (1744–1822), a Danish naturalist and missionary who studied the fishes of Greenland and authored Fauna Groenlandica. Lumpenus nubilus Richardson, 1855, described from Wellington Sound, Arctic Canada (Wellington Strait in Catalog of Fishes, 11 February 2013, and now apparently called Wellington Channel, lying between Cornwallis and Devon Islands), is a synonym. The species name is incorrectly given as nubilis in some works.

description: This species is distinguished by fin-ray counts

combined with the absence of other characters seen in related species. There are 60–67 dorsal fin spines, usually 63–66; the anal fin has 1 spine and 38–45 soft rays of equal length; there are 13–17 pectoral fin-rays; the snout lacks a fraenum; the lower pectoral fin-rays are not finger-like; and the lateral line is complete. The upper jaw projects. Scales cover the body. The caudal fin is rounded, and the dorsal and anal fins are not joined to it. It has a light-brown or yellowish body with about 12 darker blotches and spots extending onto the lower flank. The head and the pectoral fin are yellowish, and the anal fin is hyaline. There is a dusky spot at the

pectoral fin base. The caudal fin has 3–9 bars. Live specimens are translucent, and the silvery peritoneum is visible through the body wall. The species attains 36.5 cm in total length.

habitat: It is an inshore, demersal species, rarely from intertidal

areas, and is found down to 235 m, sometimes in water as cold as −1.6°C or as warm as 15.6°C. It is found on sandy to rocky bottoms, often in seagrass or algae. In Hudson and James Bays it uses estuaries on a seasonal basis. Salinities as low as 4‰ are tolerated. It dominates in the estuary at Wemindji, James Bay, during the summer, along with Fourhorn Sculpins and juvenile Cisco and Lake Whitefish. At Kuujjuarapik in southeastern Hudson Bay, larvae are more abundant at intermediate to high salinities (15–25+ psu) and at low temperatures (−1.3°C to 0.8°C). It may rest coiled like a snake or swim in small schools just off the bottom. Numbers vary drastically from year to year in the southeastern Beaufort Sea, from 2.9/ha to 64.1/ha, a measure of different year-class strengths. There are an estimated 832,000 Slender Eelblennies in Liverpool Bay, for example, making this species one of the most abundant in the Beaufort Sea.

biology: Its diet items number 86 species in the Beaufort Sea and

the waters of southeastern Baffin Island, with polychaete worms dominating. Other foods are crustaceans such as entomostraca, copepods, mysids, cumaceans, and amphipods; molluscs; clam siphons; priapulids; and fish eggs. Clam siphons are ripped off. Halibut are known to feed on this species. It is eaten by Atlantic Cod and Ringed Seals in Ungava Bay, by Black Guillemots in Digges Sound, Hudson Bay, by Brünnich’s Murres at Akpatok Island in Ungava Bay, and by Thick-billed Murres at Coats Island. On the northwest coast of Hudson Bay and at Wemindji, James Bay, it is food for Ogac, and in the estuary of the La Grande River of James Bay for Brook Trout and Ciscoes. In Hudson Bay it is eaten by Arctic Char. It may live as long as 17 years. Spawning occurs in summer in seaweed in Greenland, but it may be later in the High Arctic as October to November is reported for the Kara Sea, and fish approaching a ripe condition are reported for September in the Beaufort Sea. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae occur from May to June, suggesting a winter spawning. Eggs are large at 3.0 mm diameter, but few, at 490, in each female.

Lumpenus fabricii

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importance: It is not economically important. distribution: It is found in Baffin Bay, Davis Strait, Frobisher

Bay, Hudson Strait, Ungava Bay, Hudson and James Bays, the Arctic islands (sparsely), Dease Strait, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and the Beaufort Sea. It is also found in the Alaskan Beaufort Sea, Chukchi Sea, Bering Sea, the Sea of Japan, and to southeast Alaska. It occurs from southwest Greenland to as far south as Nova Scotia in the western Atlantic Ocean.

Lumpenus lampretaeformis (Walbaum, 1792)

Snakeblenny, lompénie-serpent

Lumpenus lampretaeformis

common names: A local name is Spidshalet Langebarn (Danish/Greenlandic). Other common names are Serpent Blenny and blennie-serpent.

Distribution of Lumpenus fabricii

sources: Vladykov (1933a); Tuck & Squires (1955); McLaren

(1958); Greendale & Hunter (1978); Morin et al. (1980); Hunter (1981); Ochman & Dodson (1982); Gaston, Cairns, Elliot, et al. (1985); Percy et al. (1985); Gaston, Elliot, et al. (1987); Mikhail & Welch (1989); Atkinson & Percy (1991, 1992); Lacho (1991); Morin et al. (1991, 1992); Ponton et al. (1993); D.B. Stewart et al. (1993).

taxonomy: The species name comes from the Latin lampetra (lamprey) and formis (form), presumably in reference to the elongate shape. It is also spelled lumpretaeformis and lampretiformis, with several authorities advocating various spellings based on interpretations of correct spellings and misprints in the original description. The subspecies L. lampretaeformis serpentinus Storer, 1848, described from off Massachusetts Bay, is the American subspecies, but it has not been used in Arctic Canada. Lumpenus lampetraeformis americanus Vladykov, 1935, described from the Gulf of St Lawrence, and Lumpenus lampetraeformis terraenovae Vladykov, 1935, described from off St John’s, Newfoundland, are synonyms. description: This species is distinguished by a combination

of fin-ray counts, the very elongate and slender body, and other

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characters. The dorsal fin spines number 68–85, usually 71–85; there is 1 spine and 47–62 soft rays in the anal fin (fin-ray counts may vary with capture locality); the snout is without a fraenum; the lateral line is complete; and the lower pectoral fin-rays are not prolonged and finger-like. The pectoral fins have 14–16 rays, and the pelvic fin 1 spine and 3 longer rays. The caudal fin ends in a pointed tip. Scales cover the body, and the lateral line is complete but indistinct. The body is pale brown to brown yellow or pinkish with some bluish tinges and relatively faint brown or greenish-yellow blotches or mottling. The belly is greenish yellow to white. The dorsal fin has up to about 18 oblique brown bars. The caudal fin is also barred with up to 8 present. The pelvic fins, anal fin tips, and lower 8–9 pectoral fin-rays are white. The species reaches 49.0 cm in standard length.

habitat: It is generally found over mud or hard bottoms, from the

shallows down to 475 m, but not intertidally. In Davis and Hudson Straits it has been reported at 116–475 m. The temperatures favoured are around 0°C, down to −1.2°C, but warmer waters are tolerated to 14°C. It constructs a protective burrow in soft substrate, taking 12–24 hours to excavate one. The burrow is usually Y-shaped and up to 73 cm long. Tail flexions are used to irrigate the burrow, and anaerobic metabolism is used. It may also be found among rocks. Larvae are pelagic.

biology: Its food is crustaceans, molluscs, polychaete worms,

Distribution of Lumpenus lampretaeformis

sources: Vladykov (1935); Tuck & Squires (1955); Gordon &

Duncan (1979); Nash (1980); Atkinson, Pelster, Bridges, Taylor, & Morris (1987); Hudon (1988); Robins et al. (1991); Eschmeyer (1998).

small starfish, brittle stars, and fishes. Cod and halibut eat this species, as do Brünnich’s Murres at Akpatok Island in Ungava Bay. Arctic Char eat this species in Ungava Bay. Its lifespan is up to nine years, with rapid growth in the first two years. Maturity is at three years and 20 cm in length. Spawning is probably in winter (December–January) and spring, and up to 1,100 demersal eggs may be deposited.

Stichaeus punctatus

importance: It is not economically important.

Arctic Shanny, stichée arctique

(Fabricius, 1780)

distribution: It is found in Davis Strait, Hudson Strait, Ungava Bay, and at a series of localities in James Bay from an unpublished electronic database (but there are no records for Hudson Bay); and from northwest, southwest, and southeast Greenland south to Massachusetts. The James Bay records are not mapped in the absence of voucher specimens and because of the apparently large gap in distribution. Its presence in James Bay needs specimens for confirmation. It is also found in the eastern North Atlantic Ocean.

Stichaeus punctatus

common names: Local names are Akkulliakitsok (Greenlandic) and Plettet Slimfisk (Danish/Greenlandic). taxonomy: The genus comes from the Greek stichao (to set in rows). The species name is the Latin punctatus (spotted).

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Stichaeus punctatus

description: This species is distinguished by the absence of fin-

ger-like lower pectoral fin-rays and of head cirri; by the presence of a single incomplete lateral line ending near mid-body (under spines 19–25); 46–51 dorsal fin spines; 32–38 soft anal fin-rays; and the dorsal fin spots. This species has a relatively deep body like its relative the Fourline Snakeblenny. There are 1–2 anal fin spines followed by variable counts of soft rays (32–35 in the Atlantic, 36–38 in Hudson Bay). The mouth is terminal. There are 15–16 rays in the large pectoral fin, and 1 spine and 4 branched rays in the pelvic fin. The overall colour is brown to scarlet, fading to paler or white on the belly. The flanks are mottled. The cheek and the ventral surface of the head have 6–7 bars. The anal fin is banded with yellow and charcoal, and the pectoral and caudal fins are banded a red brown. The dorsal fin has 4–9 large dark spots posteriorly edged in white, with the last 2 near the rear of the fin. The pelvic fins are yellow. The species reaches 22.0 cm in total length.

habitat: It is found in shallow waters down to 55 m, sometimes as

deep as 100 m, over hard bottoms and sometimes sandy or muddy areas with some rocks, hiding in crevices and under rocks in storms. It was the most abundant benthic fish of seven species at 3–10 m near the Nuvuk Islands in Hudson Bay. Larvae are planktonic and settle to the bottom in Newfoundland waters in mid-July and early August after appearing in the plankton in mid-June. At Kuujjuarapik in southeastern Hudson Bay, larvae are found at a wide range of salinities (> 1 to > 25, dominantly at 1–25 psu) and at temperatures from −1.5°C to 10.8°C. A territory is defended as soon as young settle from the plankton in Newfoundland, but curiously adults are not territorial.



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biology: Its territory is defended by a threat action that involves

erecting the dorsal fin, arching the body, and inclining the head upwards. Shaking the body, gaping, and slow tail-wagging are also involved. Nipping and chasing follow if threats are unsuccessful. The dorsal fin spots darken and the chin bars blanch in a dominant fish, the reverse in a subordinate. Its food at the Nuvuk Islands, Digges Sound, Hudson Bay, includes crustaceans such as amphipods, isopods, copepods, decapods, cumaceans, mysids, and ostracods; pelecypod and gastropod molluscs; polychaete worms; and fish eggs and larvae. Arctic Shannies are eaten by various fishes such as Greenland Halibut and Atlantic Cod and by seabirds such as the Black Guillemot and Thick-billed Murre in Digges Sound, Hudson Bay, where its importance in the diet suggests that it is one of the most abundant fish. Thick-billed Murre chicks at Coats Island have this species delivered to them by their parents. It is also eaten by Black Guillemots in the Nuvuk Islands and at Pitsulak and Piqiuliit in northeastern Hudson Bay. At Wakeham Bay, Ungava, they are eaten by Arctic Cod. On the northwest coast of Hudson Bay they are eaten by Ogac. They may live 16 years. In the Nuvuk Islands of Digges Sound the population is dominated by 2+ and 3+ fish and fish aged 6 and older. Recruitment is variable, and fish of 4–5 years were absent from samples. Spawning occurs from February to March in Newfoundland, and eggs may number up to 2,475 and up to 1.7 mm in diameter. At Kuujjuarapik in southeastern Hudson Bay, yolk sac larvae occurred from May to June, suggesting a spawning time similar to that in Newfoundland given two- to three-months’ incubation at temperatures below 0°C. Eggs are deposited in ovoid masses.

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importance: It is not economically important. distribution: It is found in Davis Strait, Hudson Strait, Ungava

Bay, and Hudson and James Bays. A record at Elu Inlet, Coronation Gulf (CMNFI 1977-1546), and a literature record (Hunter, 1981) in the Amundsen Gulf are the only western Arctic records. It is present in the Alaskan Beaufort Sea, Chukchi Sea, Bering Sea, the Seas of Okhotsk and Japan, and to British Columbia in the eastern Pacific Ocean. It is also found from northwest and southwest Greenland (as far north as 72º47' N in the Greenland waters of Baffin Bay), south to Massachusetts in the western Atlantic Ocean.

Ulvaria subbifurcata (Storer, 1839)

Radiated Shanny, ulvaire deux-lignes

common names: Another common name is Crinkly Dick. taxonomy: The genus comes from Ulva, the sea lettuce, where this fish lives. The species name comes from the Latin subbifurcatus (somewhat forked), in reference to the lateral line. This species was identified aboard ship, and no voucher specimen was kept. Its probable presence on the Labrador coast and its known presence in western Greenland warrant its tentative inclusion here (but see below).

Distribution of Stichaeus punctatus

description: This species has a relatively deep body like the Arctic Shanny and the Fourline Snakeblenny, but it is distinguished by its mid-flank lateral line from head to tail, a short lateral line above, extending to the pectoral fin tip level, and the dorsal fin blotch. The dorsal fin has 43–44 spines; the anal fin has 2 spines followed by 30–31 soft rays; the pectoral fin-rays number 15; and the branched pelvic fin-rays number 3. The upper corner of the gill cover is elongated as a rearward-pointing flap. The overall colour is brownish to gold with a pale brown, yellow, or yellowish-white belly. The flanks have irregular dull-brown bars or blotches. The anterior part of the dorsal fin has an oval blotch between spines 5–6 and 8–10, followed by 4–5 oblique, dusky bars. A black bar runs obliquely back and down from the eye. The caudal fin is barred with 3–4 rows of spots, which may coalesce. Females become a light yellow with brown body marks and a grey-white belly in the week before spawning. During spawning, the female becomes white, while the male is almost black. After spawning, the female is grey but returns to her normal colour within a few days. The species reaches 18.0 cm in total length.

sources: Johansen (1927a); Brown & Green (1976); Farwell,

habitat: This shanny is found on rocky shores among seaweed

Green, & Pepper (1976); Gaston, Cairns, Noble, et al. (1981); Hunter (1981); Cairns (1982, 1987a, 1987b); Gaston, Cairns, Elliot, et al. (1985); Morrison & Gaston (1986); Gaston (1987, 1989a, 1991); Gaston, Elliot, & Noble (1987); Hudon (1988); Mikhail & Welch (1989); Green (1990); Keats, Steele, Green, & Martel (1993); Ponton et al. (1993).

and down to 55 m or more. Canadian Arctic records were at 130– 333 m, presumably being expatriates in open waters. Adults are most active after dark and have a particular home site to which they can return from at least 270 m away, using sight and smell. Usually they remain in an area of less than 3 sq m.

biology: Its food includes various crustaceans, marine worms,

molluscs, echinoderms, small fishes, Sand Lance and Capelin eggs, and tube feet nipped from sea urchins. Worms are favoured by larger fish in Newfoundland. Radiated Shannies are eaten by Atlantic Cod and the Grubby, by other fishes including Skates and dogfish, and even by ducks. Males live about 10 years, and females 8 years, first spawning at 3–5 years. Spawning occurs in spring and early summer, in early May to June at 1.5°C–4.0°C in Newfoundland. The male swims under and around the female, nudging her belly with his snout so that she tilts to one side, and eggs and sperm are released when both fish are tilted. The

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male guards multiple egg masses, nudging and fanning the eggs. Egg masses in Newfoundland have up to 2,706 eggs. Males may spawn up to four times, but females only once, as the mean number of eggs in the ovary is only 1,512. Eggs have a mean diameter of 1.55 mm. Larvae are pelagic.

importance: It is not economically important. distribution: It is found in Davis Strait off southern Baffin Island at 64.525° N, 61.515° W, and off the tip of Labrador at 60.04817° N, 63.1105° W, as single records (cruise records without voucher specimens). It is also found from the northern Labrador coast (five cruise records, without voucher specimens), the Strait of Belle Isle, south to Massachusetts, and in western Greenland. This is a species of relatively shallow waters, and its presence in Davis Strait at depths greater than those recorded is questionable. The specimen may have been mis-identified but could be an adult expatriate (although this condition has not been recorded for this species) or possibly an expatriate larva that developed while drifting. This type of distribution serves as an example of unusual records that should be preserved as voucher specimens in a museum and thus made available for verification.

Distribution of Ulvaria subbifurcata

sources: See the family sources and the bibliography.



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Pholis fasciata

Family Pholidae

(Bloch and Schneider, 1801) Banded Gunnel, sigouine rubanée

Gunnels, Sigouines

Brian W. Coad

Pholis fasciata

common names: Local names are Kugsaunak; Kurksaunak and Quvssaunaq (Greenlandic); and Båndet Tangspræl (Danish/Greenlandic). Another common name is Bering Gunnel. Gunnels are found principally in the North Pacific Ocean with a few species in the North Atlantic Ocean. There are about 15 species, including 8 in Canada, of which 1 is in the Arctic. The maximum length is 46 cm although most species are under 30 cm. These fishes have eel-like, elongate, compressed bodies with 73–100 spines in the long dorsal fin and no soft rays. The anal fin has 1–3 spines and 32–53 soft rays and is about half as long as the dorsal fin. Gunnels have a shorter anal fin than the superficially similar Shanny Family and are usually larger. The dorsal and anal fins are joined to the rounded caudal fin. The pectoral and pelvic fins are small or absent. Even when present, the pelvic fin has only 1 spine and 1 soft ray. Scales are minute and embedded with an absent or incomplete single lateral line. There are no ribs. Gill membranes are attached to each other but are free from the isthmus. The teeth are small and conical. These fishes are common inshore and can often be found under rocks at low tide or amongst seaweed. They are easily caught by hand, albeit a little slippery to hold. Their movement is eel-like. Their food is crustaceans and molluscs. They are often eaten by various sea bird species and are intermediate hosts for parasites that reach their adult form in the sea birds. The eggs are laid in masses on the bottom, and larvae are pelagic.

sources: Jensen (1942a); Yatsu (1981); Mecklenburg (2003a).

taxonomy: The species name comes from the Latin fasciatus

(banded). Gunnellus groenlandicus Reinhardt, 1830, and Centronotus gunelliformis Günther, 1861, both described from Greenland, are synonyms.

description: There are 83–91 dorsal fin spines, 2 anal fin spines

with 41–48 soft rays, and a shorter anal fin (i.e., having a longer preanal distance) than the superficially similar Shanny Family. The pectoral fins are present with 11–13 rays, but the pelvic fins may be present or absent. It is usually a bright red to orange red in overall colour, fading to a yellowish grey in preservative. A dark band over the head, through and below the eye posteriorly, is offset by a light band with a posterior dark margin. The dorsal fin and the upper back have 10–12 triangular light areas, often with darker areas between forming saddles, and are speckled with darker spots. The flanks have light and dark bars alternating, and extending to the belly, and may become scarlet. The anal fin has up to 8 light bars, and the caudal fin is uniformly coloured a light red or orange red. The species attains 30.0 cm in total length.

habitat: It is found intertidally or below the tide level down to 94

m, usually 30–46 m, and usually associated with gravel, rocks, and boulders but also areas of mud. It is not usually found in tide pools. Temperatures up to 10°C and down to −1°C are tolerated, as are salinities as low as 4‰.

biology: Its food is small crustaceans, such as amphipods, and

polychaete worms. It is eaten by various sea birds such as Black Guillemots in Digges Sound, Hudson Bay, by Thick-billed Murre chicks at Coats Island, and by inshore fishes such as Arctic Cod and Sculpins. At Kuujjuarapik in southeastern Hudson Bay, post-larvae, possibly this species, occur in July and possibly August. The eggs are laid in clam shells in west Greenland.

importance: It is not economically important.

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distribution: The species is found in western Hudson Strait including Digges Sound, Hudson and James Bays, Melville Sound, Coronation Gulf, the Arctic islands in Northumberland Sound, and at eastern Axel Heiburg Island. It has been recorded west of Point Barrow in Alaska, a considerable gap until Coronation Gulf in Canada. It is also found in the North Pacific and North Atlantic Oceans, reaching the Seas of Okhotsk and Japan in the Pacific, and southwest Greenland in the Atlantic. The Arctic islands’ records are from literature reports and require confirmation (Richardson, 1855; Zoltai, Boothroyd, & Scotter, 1981).

Distribution of Pholis fasciata

sources: Richardson (1855); Feilden (1887); Green (1970); Zoltai

et al. (1981); Ochman & Dodson (1982); Gaston, Cairns, Elliot, et al. (1985); Gaston, Elliot, et al. (1987); Gaston (1989a); Hudon (1988); Ponton et al. (1993).



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sources: Barsukov (1959); Smidt (1981); Albikovskaya (1982,

Family Anarhichadidae Wolffishes, Poissons-loups

1983); Riget & Messtorff (1988); Gusev & Shevlev (1997); Mecklenburg (2003b); Kulka, Min, & Thompson (2006); Kulka, Hood, & Huntington (2007); Department of Fisheries and Oceans (2011, 2013b); Simpson, Mello, Miri, & Treble (2011).

Brian W. Coad Anarhichas denticulatus Krøyer, 1845

Northern Wolffish, loup à tête large

common names: Local names are Akoak and Akoaksaluk (Old

Wolffishes or Sea Catfishes are found in the North Atlantic, North Pacific, and Arctic Oceans. There are 9 species in the family, including 5 in Canada, of which 4 are in the Arctic. The maximum size is about 2.5 m. These fishes have large heads with strong, protruding canine teeth anteriorly and large molar teeth posteriorly. The body scales are minute and cycloid or absent. There is no lateral line. The dorsal fin contains only flexible spines. The long dorsal and anal fins may be confluent with the tail. The pectoral fins are large and fan shaped, and pelvic fins are absent. Gill openings are only on the flank. Curiously the teeth are lost and replaced annually, probably during the spawning period. Wolffishes apparently fast during spawning and tooth replacement. These fishes are found in cold coastal waters, feeding on molluscs, crustaceans, sea urchins, and starfish. They are usually found on rocky bottoms in shallow water with a territory based on a crevice, hole, or cave. Anarhichas denticulatus is benthopelagic, however. Males and females guard the large eggs that are deposited in a clump by the females. Fertilization is probably internal. They are economically important and are caught by trawlers. The total Canadian catch was as high as 12,000 t in 1971 and 10,200 t in 1981, but they have not been considered in Arctic waters for exploitation. The 1989 catch in Canada was worth about $439,000. Atlantic Wolffishes (Anarhichas lupus) are incidental catches in trawl fisheries for groundfish and have been recorded in commercial catches as, and are sold as, “catfish.” The skin is used to make leather. Their habitat is damaged or destroyed by bottom trawling and by dredging for scallops and clams. Rocks used for shelter and nesting are disturbed, and sediments are stirred up, smothering spawning areas and eggs and damaging fish gills. Wolffishes must be handled very carefully as their strong jaws and teeth can inflict a nasty wound. Wolffishes are aggressive and attack with a hissing sound.

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Woman Fish) (Inuktitut); and Najorpilik, Qeeraasaq, Tungujortoq, and Utoqulaaq (Greenlandic). Other common names are Blue Catfish, Blue Sea Cat, Blue Wolffish, Jelly Cat, Jelly Wolffish, and loup gélatineuse.

taxonomy: The genus is Anarhichas, an old name for Anarhichas lupus, which derives from the Greek anarrichaomai (to climb or scramble up), the allusion not being obvious. The species name is the Latin denticulatus (small-toothed). Anarhichas latifrons Steenstrup and Hallgrimsson in Steenstrup, 1876, described from Iceland; Lycichthys paucidens Gill, 1905, described from Banquereau near Nova Scotia; and Lycichthys fortidens Gill, 1911, described from near Nova Scotia, Banquereau, are all synonyms. description: This species is distinguished by colour (not distinctly spotted or barred), by the teeth on the vomer not extending back as far as the palatine teeth, and by the upper lip being thicker than the lower lip and covered with papillae. The dorsal fin spines number 76–81, the anal fin-rays 45–50, the pectoral fin-rays 19–22, and the vertebrae 78–82. The caudal fin-rays number 18–22, usually 20. The flesh of this species is more jellied (soft and watery) than that of its relatives. The body is yellowish grey, grey brown, dark brown, blue black or violet, and may be darkly spotted. Spots may not be clearly defined and may form a weak bar. However, some fish do have sharp, distinct spots and superficially resemble Spotted Wolffish (Anarhichas minor), although the background is grey in the Northern Wolffish rather than a sharply contrasting white as in the Spotted Wolffish. The species reaches 143.8 cm, possibly 180.0 cm, in total length, and 19.5 kg, and possibly more, in weight. habitat: This wolffish has been found from the surface down to

1,700 m but prefers deeper waters with temperatures below 6°C, especially 1°C–2°C. It is found over silt, sand, and shell bottoms, preferring rocky bottoms during spawning. On the Greenland side of Davis Strait they are mostly found at 1,200–1,600 m, and on the Canadian side in Davis Strait and Baffin Bay at 164–1,429 m. In Ungava Bay four fish were caught at 0–399 m. It does not school. It has a

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to comb jellies, jellyfish, and smaller fishes, such as Grenadiers and Lumpsuckers. At Mould Bay, Prince Patrick Island, they eat Atlantic Spiny Lumpsucker. However, the majority of food items in stomach contents are pelagic fishes, and it is called a “fish specialist.” This species is apparently eaten by Ringed Seals near Iluvilik Island in Prince Albert Sound in the Arctic, by Atlantic Cod and Golden Redfish in the Atlantic, and elsewhere by Greenland Sharks. This species lives at least 12 years and reaches maturity at 5–6 years, with length at 50% maturity being about 54 cm. Spawning probably occurs in Canadian Atlantic waters in late autumn and early winter. Up to 46,500 eggs are laid on the bottom and are up to 8.0 mm in diameter. Young are 25–26 mm on hatching and are pelagic.

importance: The flesh develops a “jelly” condition or watery

flesh, making this species unsuitable for sale. The skin is also unsuitable as leather. The Committee on the Status of Endangered Wildlife in Canada assessed it as “Threatened” in November 2012. By-catches of this species during the Greenland Halibut fishery in NAFO Division 0A were 4 t in both 2009 and 2010, and in Division 0B were 5 t and 16 t in these years.

The teeth on the central vomer do not extend back beyond the lateral palatines

greater depth range than do other species off Newfoundland. Young have an extended pelagic period, and adults probably migrate considerable distances to counteract the drift away from adult spawning grounds. Adults also feed off the bottom more frequently than do the other species.

biology: This species has conical teeth and feeds on foods with

thin shells. Its food includes both bottom and free-swimming invertebrates, from crabs, brittle stars, starfish, sea urchins, and molluscs



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distribution: It is found in Baffin Bay, Davis Strait, eastern Hudson Strait to Ungava Bay; possibly from Amundsen Gulf, western Victoria Island, and Bathurst Inlet; and definitely from Prince Patrick Island (CMNFI 1959-0013). It has not been recorded from the Alaskan Beaufort Sea. It is also found in the North Atlantic Ocean from northwest, southwest, and southeast Greenland south to Cape Cod. The record from near Iluvilik in Prince Albert Sound in the Amundsen Gulf (Smith, 1977) may not be this species, but the Bering Wolffish (Anarhichas orientalis), because it was based solely on a photograph of a damaged specimen. The Bathurst Inlet record is from the literature (Abrahmson, 1963) and could be a misidentification.

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Anarhichas lupus Linnaeus, 1758

Atlantic Wolffish, loup atlantique

common names: A local name is Qeeraaraq (Greenlandic). Other common names are Catfish, Ocean Whitefish, Sea Cat, Striped Wolffish, and loup de mer. taxonomy: The species name is the Latin lupus (wolf). description: This species is distinguished by the colour pattern

Distribution of Anarhichas denticulatus

sources: Hildebrand (1948); Walters (1953c); MacDonald (1954);

Abrahmson (1963); Barsukov (1972); Konstantinov & Podrazhanskaya (1972); Smith (1977); Templeman (1986c); Hudon (1990a); Fruge & Wiswar (1991); V. Berg et al. (1997); Gusev & Shevelev (1997); Jørgensen et al. (2005).

(distinct bars), by having the teeth on the vomer bone in the centre of the roof of the mouth extending back beyond the palatine teeth at the sides, and by fin-ray counts. The dorsal fin spines number 69–79, anal fin-rays 42–48, pectoral fin-rays 18–21, and vertebrae 72–78. The caudal fin-rays number 22–26, usually 24–25. Mean vertebral counts increase from north to south, the reverse of the usual pattern in other fish species with pelagic eggs. The colour is variable from blue grey or slate, to yellowish, olive green, and purplish brown. The flank and the dorsal fin have 8–14 dark brownish and irregular bars, often edged by white. Juveniles lack these bars, while the juvenile Spotted Wolffish (to 20.0 cm) is barred. There is a white blotch on the anterior dorsal fin, although some have a black blotch in this position. A white patch is evident at the base of the pectoral fin. The lateral-line pores on the head are white. White markings are behaviourally controlled. The belly is a dirty white. Spawning fish develop pink, red, or orange lips and mouth cavity. Juveniles tend to

Anarhichas lupus

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m, perhaps to 918 m, in the south. Even in southern waters it may occasionally be found in tide pools. Water temperatures are as low as −1.9°C and up to 18°C, although temperatures below 4°C are preferred. It does not form schools and favours hard bottoms on the continental shelf, especially when spawning. It prefers a rocky substrate when feeding and may live colonially. Individuals can be observed in shelters found on 15°–30° slopes, with 1–5 openings and good water circulation. The shelters are not defended, and the most convenient one is utilized. Individuals tend to stay in one area for many years, but there are small, seasonal, inshore-offshore migrations, probably restricted to sexually mature adults. Observations vary on whether it is active by day or is only nocturnal.

biology: Its food is mainly sea urchins, starfish, and crustaceans

The teeth on the central vomer extend back beyond the lateral palatines

be yellow brown overall. The species reaches about 152.0 cm in total length and 24.0 kg in weight, but such large fish are rare.

habitat: This wolffish is more common in southern waters than its relatives. It is found close to shore in the north, but at 22–366



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with the occasional unwary redfish (Sebastes spp.) and Canadian Plaice, and also Skates. It favours molluscs and is called a “mollusc specialist.” Anterior teeth are used for grasping prey, and the molariform lateral and posterior teeth for crushing. Green sea urchins are 75% of the diet in eastern Newfoundland. Cannibalism of eggs and larvae occurs. Atlantic Cod are known to eat young Wolffishes. Greenland Shark and a variety of other fish eat this species. Adults are mature at 5–10 years of age when 50–60 cm long, although some females are mature at 31 cm, and some males only at 69 cm. Off northern Labrador the smallest mature female was 43 cm. They live about 22 years. Spawning is preceded by an inshore migration in spring, but only takes place in August–September in Newfoundland waters. Courtship behaviour is extended, lasting four to five months before spawning.

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Eggs are laid under and between boulders and hatch in mid-December. Fertilization is internal, copulation taking place during a 3–6 hour period when the male and the female are close together. Eggs are released over the 7–15 hours following as the female lies on her side. Fecundity reaches 37,920 eggs. The eggs are very large, up to 6.8 mm in diameter, and are yellow. Larvae are 20+ mm on hatching and are pelagic until 3–6 cm long. Males do not feed while guarding egg clusters, which are up to 14 cm wide. Males also rotate the egg mass, coat it with skin mucus to prevent infections, and aerate it.

common names: Local names are Qeeraq and Qeerngaq Milagulaar (Greenlandic). Other common names are Leopardfish, Spotted Catfish, and Spotted Sea Cat.

importance: It is not economically important in Arctic Canada,

taxonomy: The species name is the Latin minor (smaller).

but catches of about 6,000 t annually were taken off west Greenland in the 1970s, mostly Spotted Wolffish but including some of this species. It is also a game fish and is often displayed in aquaria. The Committee on the Status of Endangered Wildlife in Canada assessed it as of “Special Concern” in November 2012. Bottom trawling and dredging have probably damaged the habitat of the species, and its slow growth and take in by-catches have led to declines in numbers in the Newfoundland region. By-catches of this species during the Greenland Halibut fishery in NAFO Division 0B were 6 t and 2 t in 2009 and 2010. It is sold fresh or frozen and is eaten baked, boiled, broiled, fried, and steamed.

distribution: It is found in Davis Strait, central and eastern Hudson Strait, and northern Ungava Bay; from northwest, southwest, and southeast Greenland to Labrador; and through maritime Canada to New Jersey. It is also found in Europe.

Anarhichas minor Olafsen, 1772

Spotted Wolffish, loup tacheté

description: This wolffish is distinguished by colour (distinct

spots), by the teeth on the palatine bones at the margins of the roof of the mouth being equal to or extending beyond the vomer teeth in the centre of the mouth, and by the upper lip being covered with irregular folds and not being thicker than the lower lip. The dorsal fin spines number 74–80, the anal fin-rays 44–48, and the vertebrae 76–79. The caudal fin-rays number 20–23, usually 21. The colour is yellowish or pale olive to chocolate brown with small to large distinctive brown-black spots on the head, the upper flank, and the dorsal and caudal fins. The belly may be whitish or dark like the adjacent flank. The caudal fin is dark with a pale or yellowish margin. Young less than 20 cm long have 5–11 dark bands on the body and up to 7 spots on the dorsal fin. The species reaches about 180.0 cm, perhaps 2.0 m, in total length, but larger sizes are disputed, and 23.0 kg in weight.

habitat: This species is found in deeper waters over mud, sand, or

shell bottoms, preferring stony bottoms when spawning. It is found down to 600 m in Atlantic Canada and to 1,046 m elsewhere, at preferred temperatures below 5°C, and as low as −1.32°C elsewhere. Arctic populations elsewhere are known to be as shallow as 25 m, and there are a few records at 148–337 m in Hudson Strait; at 301.5– 556.5 m and 2.2°C–5.6°C in Davis Strait and southern Baffin Bay; and as shallow as 220 m and as deep as 659 m in the eastern Canadian Arctic. It does not school, but there are movements inshore and into fiords during June and July in Greenland. Juveniles spent most of their time in sheltered areas under laboratory conditions, and this may be an important consideration in their conservation.

biology: Its food includes various hard-shelled crustaceans,

Distribution of Anarhichas lupus

sources: Templeman (1984b, 1986b); Keats, South, & Steele (1985,

1986); Hudon (1990a); Nelson & Ross (1992); Nielsen et al. (1992); O’Dea & Haedrich (2003).

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molluscs, and most importantly echinoderms (it is called an “echinoderm specialst”), but also some polychaete worms, algae, and fish, including those discarded from trawlers. Atlantic Cod and redfishes (Sebastes spp.) are eaten off Newfoundland. Atlantic Cod and Greenland Shark are known to eat this wolffish. Spotted Wolffish live to be 21 years old. Females are mature a year earlier than males, at 48–62 cm in some populations and 75–80 cm in others, and all females mature at sizes greater than 92 cm. Males mature at 53–71 cm. Maturity may occur at smaller sizes further north, off Greenland, for example. The age of maturity in the Barents Sea starts at age seven and 53 cm for females and at age nine and 66 cm for males.

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Spawning occurs in summer in the northwest Atlantic generally. Up to 54,600 eggs of 6.5 mm in diameter are laid in clumps on the sea bed. Larvae are 20–24 mm on hatching, are pelagic, and become bottom dwellers at 4–7 cm. This fish has functionally distinct haemoglobin in its blood, evolved to adapt to temperature fluctuations in Arctic waters.

importance: It is not economically important currently, although

the skin was tanned in Greenland beginning in 1938 as a leather for sale in Denmark, but the fishery later switched mostly to fillets. It is also caught as a game fish and in by-catches from trawl and longline fisheries. The Committee on the Status of Endangered Wildlife in Canada assessed it as “Threatened” in November 2012. By-catches of this species during the Greenland Halibut fishery in NAFO Division 0B were 3 t and 1 t in 2009 and 2010, respectively.

The teeth on the central vomer do not extend back beyond the lateral palatines

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distribution: It is found in Baffin Bay, Davis Strait, eastern Hudson Strait, northern Ungava Bay, and northwest, southwest, and southeast Greenland, south to Massachusetts, possibly New Jersey. It is found as far north as Thule in northwest Greenland and also in Europe.

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Anarhichas orientalis Pallas, 1814

Bering Wolffish, loup de Béring

common names: A local name is Aaqaksaaq (Inuktitut). An­other common name is Old Womanfish. taxonomy: The species name is the Latin orientalis (eastern). description: This wolffish is distinguished by colour (no bars),

Distribution of Anarhichas minor

sources: Hildebrand (1948); Templeman (1986a); Hudon (1990a);

Gusev & Shevelev (1997); Treble et al. (2000); Verde et al. (2002); Jørgensen et al. (2005); Verde & di Prisco (2006); Lachance, Dutil, Larocque, & Daigle (2010).

the teeth on the vomer bone in the centre of the roof of the mouth extending back beyond the palatine teeth at the sides, and by fin-ray counts. The dorsal fin spines number 80–88, the anal fin-rays 50–55, and the pectoral fin-rays 20–22. The caudal fin is deeper and more rounded than in other species, and the upper lip is wrinkled rather than papillate. The overall colour is a dark brown, reddish brown, or black with marbling and blotching. Young have spots on the head, 3–5 interrupted flank stripes on a yellowish-white background, 2 lightbrown stripes on the dorsal fin, an indistinct light-brown stripe near the anal fin margin, and a similar one on the pectoral fin. The species reaches 124.0 cm in total length and 19.5 kg in weight.

habitat: It is known to live in very shallow water among algae-covered stones or over gravel and sand. Its maximum depth is probably not more than 100 m. It moves offshore when ice forms along the coast. Salinities of 19‰–25‰ and temperatures of 4°C–13°C are reported. biology: The species is known only from Bathurst Inlet, Nunavut

(and Camden Bay of the Alaskan Beaufort Sea near the Canadian border). Its lifespan is at least 17 years. Crabs and mussels (e.g., Mytilus edulis in Bathurst Inlet) are important foods. Spawning may occur in spring and summer, and egg diameters are up to 4.5 mm. The larvae are pelagic.

Anarhichas orientalis



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The teeth on the central vomer extend back beyond the lateral palatines

Anarhichas orientalis

importance: The Committee on the Status of Endangered Wildlife in Canada assessed it as “Data Deficient” in November 2002, and the Canadian Species at Risk Act lists it as of “Special Concern.” distribution: The species is found rarely in Bathurst Inlet and western Coppermine Gulf in Canada (four records: CMNFI 19660075, 1967-0719, 1971-0108, and Hiscock and Grant, 2004), the northwest Pacific Ocean in the Seas of Okhotsk and Japan, in the Alaskan Beaufort and Chukchi Seas, and in southeast Alaska, but not on the British Columbia coast.

Distribution of Anarhichas orientalis

sources: Hunter (1981); Houston & McAllister (1990a); Fruge & Wiswar (1991); Hiscock & Grant (2004).

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Family Chiasmodontidae Black Swallowers, Grands avaleurs

Brian W. Coad

the first with 7–8 flexible spines and the second with 0–1 flexible spine and 18–29 segmented rays. Scales may be absent or present, small, and rough. There are 6–7 branchiostegal rays. These are deep-sea, mesopelagic, or bathypelagic fishes that often catch and eat fishes larger than themselves. Some species have an unusual juvenile stage called a gargaropteron with greatly elongated paired fin-rays and a long snout. Early specimens of Black Swallowers became known to science when they floated to the surface after trying unsuccessfully to swallow very large prey. They are not economically important. A recent revision of the genus indicates that there is only one species in Arctic waters and that records of C. bolangeri Osório, 1909, and C. niger Johnson, 1864, are C. harteli.

sources: Melo (2009); Prokofiev & Kukuev (2009); Prokofiev (2010).

Family Chiasmodontidae is found in the temperate to tropical waters of the Atlantic, Pacific, and Indian Oceans with about 15 species, including 5 in Canada, of which 1 enters the Arctic. The maximum size is 26.2 cm in standard length. Black Swallowers have a very distensible mouth and stomach and an overall black colour that gives them their name. The upper jaw bones are long, slender, and joined at the rear. The anterior tip of the premaxilla is wide dorsally and diverges laterally. The halves of the lower jaw can separate to enlarge the mouth. The right and left halves of the pelvic girdle are separate from each other and from the pectoral fins. Even the internal supporting skeleton of the anal fin is not attached to the body muscles, allowing the stomach to distend further. The teeth in both jaws are long and sharp. Either no gill rakers are present, or gill teeth are fused to bony plates. The dorsal head surface is rugose and pitted with sensory pores. There are two dorsal fins,

Chiasmodon harteli Melo, 2009

Hartel’s Swallower, avaleur d’Hartel

common names: A local name is Hartels Slughalsfisk (Danish/ Greenlandic).

taxonomy: The genus comes from the Greek chiasma (a cross in the form of the letter X) and odonos (tooth), referring to the two anterior canine teeth that cross when depressed. The species is named after Karsten Hartel of the Museum of Comparative Zoology at Harvard University.

Chiasmodon harteli



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Chiasmodon harteli

The distinction of this species from the taxon C. niger Johnson, 1864, recognized as a wide-spread species by some authors, is disputed.

description: The distensible mouth and stomach and the colour

are characteristic. The illustration shows a fish containing a large prey item. The skin is naked in fish larger than 56.5 mm in standard length. The dorsal fin has 10–13 spines and 23–27 soft rays preceded by 3–4 unbranched rays. The anal fin has 21–28 rays preceded by 3–5 unbranched rays. The pectoral fin has 12–14 rays, and the pelvic fin has 1 spine and 5–6 soft rays. Lateral-line pores number 89–95, supraorbital pores 6, and total vertebrae 47–48. There is a small and stout spine at the lower corner of the preopercle. The overall colour is black to brownish. The species attains 19.8 cm in standard length.

habitat: This is a mesopelagic to bathypelagic species. Specimens

from Canadian waters have been caught in excess of 1,000 m and floating at the surface. In Davis Strait catches have been made at 459–1,457.5 m and 1.6ºC–4.1ºC.

biology: Its biology is poorly known. It probably undergoes a

daily vertical migration to feed on planktophagous and other piscivorous fishes. Spawning may occur in late autumn–winter.

importance: It is not economically important. distribution: It is found in Davis Strait, southwest and southeast Greenland, the eastern and western North Atlantic Ocean, and probably all oceans.

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Distribution of Chiasmodon harteli

sources: Johnson & Keene, in Smith & Heemstra (1986); Jørgensen et al. (2005); Więcaszek, Melo, Szulc, & Sobecka (2011).

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Family Ammodytidae Sand Lances, Lançons

Brian W. Coad

Sand Lances, Launces, or Sand Eels are found in the Atlantic, Indian, Pacific, and Arctic Oceans. There are about 27 species, including 3 in Canada, of which 2 are in the Arctic. The maximum size is 35 cm in standard length. These fishes are characterized by their elongate shape; one long dorsal fin with 40–69 soft rays, which can fold back into a groove; a shorter anal fin; the absence of pelvic fins in Canadian species; a lateral line running just below the dorsal fin; minute, cycloid scales that seem to form diagonal plates; the absence of teeth; a fleshy ridge running low along each side; diagonal plicae or skin folds; and a protruding lower jaw. The caudal fin is forked. There is no gas bladder. The overall colour is silvery, and the back is bluish to greenish. The taxonomy of some Sand Lances has been confused, and various species names and distributions are advocated by different authors. More extensive material and molecular analyses are required to resolve some of the issues involving these morphologically similar fishes in Arctic waters. Ammodytes americanus DeKay, 1842, is an inshore species found along the Atlantic coast of North America from Virginia as far north as northern Labrador. Some authors have it extending into Hudson Bay. Ammodytes marinus Raitt, 1934, is recorded from the eastern North Atlantic Ocean and from Greenland, including its western shores. This species may be found in the Canadian Arctic, or it may be A. americanus. Ammodytes hexapterus may then be restricted to the northern Pacific Ocean and the western Arctic, with an uncertain eastern boundary. Limited molecular data on Hudson Bay samples identify them as A. hexapterus, and this name is retained here with reservations. These fishes often bury themselves in sandy shallows, with only the head protruding, and may be left on a beach after the tide goes out. They dive in head first, aided by the pointed lower jaw. Burrowing provides protection from predators, and in Arctic waters they may hide in cavities in ice. Normal swimming is with an eel-like motion. The sharp lower jaw may perforate the gut of a predator like the Atlantic Cod, and the Sand Lance then becomes encysted in the body wall. Sand Lances occur in very large schools (up to 1,800 million fish),



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and these are of immense importance as food for other fishes, particularly commercial species, for whales, seals, and seabirds such as Thick-billed Murres in the eastern Arctic. Recent studies have shown an increase in Sand Lances in the diet of Thick-billed Murres associated with a general warming of Hudson Bay waters owing to climate change. Sand Lances and Capelin replace Arctic Cod as an intermediate carnivore in southern Hudson Bay and James Bay. Larvae dominate the ichthyoplankton community in coastal waters of Hudson Bay and are likely to be severely affected by reduction in freshwater flow, and thus by delayed ice break-up, after hydro-electric developments in this region. Blooms of ice microalgae in Hudson Bay facilitate reproduction of copepods, which are the main prey of larval Sand Lances, and any variation in timing can be critical. Sand Lance larvae account for 33% of the total fish larvae off Kuujjuarapik in southeastern Hudson Bay, where adults form a large proportion of the diet of Ringed Seals. The highest densities of larvae were found in cold waters with a salinity exceeding 25 psu. Sand Lance larvae in the Grande Baleine estuary of James Bay are small and numerous, hatch before ice break-up, and feed on small prey, a strategy in contrast to that of Sculpins and Pricklebacks, which produce fewer and larger larvae after ice break-up that in turn feed on larger food items. A strong abundance of larval and juvenile lances in the Beaufort Sea in 2011 could affect survival, by competition, of larval Boreogadus saida and, when the cod are adult, predators of the cod. Lances can be used as bait by anglers; they are fished commercially not for table food but for fish-meal and oil. Over 660,000 t were caught in the western Atlantic in 1982.

sources: Jensen (1941a); Richards, Perlmutter, & McAneny

(1963); Richards (1965); Reay (1970); Dalley & Winters (1987); Winters & Dalley (1988); Winters (1989); Nizinski, Collette, & Washington (1990); Ida, Sirimontaporn, & Monkolprasit (1994); Mitchell, McCarthy, & Verspoor (1998).

Ammodytes dubius Reinhardt, 1837

Northern Sand Lance, lançon du nord

common names: Local names are Amajak (Inuktitut) and Putorugtoq (Greenlandic). Other common names are Offshore Sand Lance and Sand Eel.

taxonomy: The genus comes from the Greek ammos (sand) and dyo or dytes (to dive, diving). The species name is the Latin dubius (doubtful). Ammodytes dubius hudsonius Vladykov, 1933, is described from James and Hudson Bays and Hudson Strait. If diagnosable, this would be a distinct species, Ammodytes hudsonius.

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Ammodytes dubius

description: This species is distinguished from its only Arctic relative, the Pacific Sand Lance, by counts of several characters. Overlaps occur in these counts, which are for the whole species range, although means are for northern fish from Labrador, and some fish may be difficult to assign to species. There are 55–69 dorsal fin-rays (usually 61–67, with a mean of 62.4), 26–36 anal fin-rays (usually 31–34, with a mean of 32.1), and 64–78 vertebrae (usually 71–76, with a mean of 71.9), and the number of oblique, lateral plicae or skin folds is 124–147 (usually 128–138, with a mean of 131.8). Canadian offshore populations of large Sand Lances with slender bodies and high vertebral counts are the Northern Sand Lance. The sides are silvery with an iridescent green or blue lustre, sometimes absent, and the belly is white. The back is bronze, blue green, or olive. In preservative a light stripe between the dark back and the dark flank stripe is usually absent or at least less developed compared to A. hexapterus. The species reaches 37.2 cm in length, possibly larger. habitat: This is an offshore, bottom-living species usually found

at 15–108 m and 1°C–11°C (on the Scotian Shelf) and down to −1.7°C in west Greenland over sand or fine gravel, into which they burrow. Rocky or muddy areas are generally avoided. In Hudson Bay it uses estuaries on a seasonal basis and can be found at 0–31.5 m. Larvae in Rupert Bay and nearby James Bay have been caught mostly at temperatures of 9°C–20°C and salinities of 1‰–17‰ from June to August. This species may hibernate over winter in shallow waters in the north in concentrations of 3–6 per sq m.

biology: Its food is mainly copepod crustaceans that undergo a

migration towards the surface at night, and this Sand Lance probably follows the copepods to feed. Copepods are strained by filter

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feeding from the water, using long gill rakers, but larger food items like polychaete worms are picked selectively. Sand Lances are food for a wide variety of fishes such as Atlantic Salmon in the Koksoak River estuary, Ungava Bay; Atlantic Cod at Killiniq, Ungava Bay; Arctic Char in southeast Ungava Bay and in the Belcher Islands; and Ogac at Wemindji, James Bay. As well, whales and sea birds (such as Thick-billed Murres in Digges Sound, Hudson Bay; at Cape Hay, Bylot Island; and at Coats Island, Nunavut) eat Sand Lances. Sand Lances are the most important food for Canadian Plaice. Atlantic Puffin chicks are fed this Sand Lance at Coburg Island, Nunavut. Its lifespan may be about 9–10 years, and maturity is reached at 1–2 years. Males and females grow at about the same rate, but growth rates vary at different localities. Spawning begins in Canadian waters in late November and peaks in December–January, but extends to May–June in Fortune Bay, Newfoundland. However, larvae identified as this species in Rupert and James Bays suggest spawning in June and July. At Kuujjuarapik in southeastern Hudson Bay, Sand Lance that may be this species appeared as yolk sac larvae from April to June. The eggs are up to 1.23 mm in diameter, with fecundity estimates of up to 22,904 eggs. The young are planktonic.

importance: It is not economically important. distribution: It is found at Nirjutiqavvik (Coburg Island), southern Baffin Island, Hudson Strait, Ungava Bay, Hudson and James Bays including the Belcher Islands, Dease Strait, Amundsen Gulf, and the Beaufort Sea. It is also found from northwest and southwest Greenland, along the Labrador coast, and south to Virginia and North Carolina, and in Europe.

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Ammodytes hexapterus Pallas, 1814

Pacific Sand Lance, lançon gourdeau

Ammodytes hexapterus

common names: Another common name is Stout Sand Lance. taxonomy: The species name comes from the Greek hex (six) and pteros (wings or fins), the meaning of which is unclear. description: This species is distinguished from its only Arctic

Distribution of Ammodytes dubius

sources: Simard & Legendre (1977); Petersen (1978); Morin et

al. (1980); Richards (1982); Gillis & Allard (1984); Robitaille, Côté, Shooner, & Hayeur (1984); Gaston, Cairns, Elliot, et al. (1985); Gaston (1987); Gaston, Elliot, & Noble (1987); Adams, Power, & Barton (1989); Morin et al. (1991); Ponton et al. (1993); Robards et al. (2000); Falardeau, Robert, & Fortier (2012).

relative, the Northern Sand Lance, by counts of several characters. Overlaps occur in these counts, which are for the whole species range, although the means are for northern fish from Labrador, and some fish may be difficult to assign to species. There are 51–63 dorsal fin-rays (usually 55–60, with a mean of 57.9), 23–34 anal fin-rays (usually 26–31, with a mean of 29.7), and 60–75 vertebrae (usually 63–70, with a mean of 67.2), and there are 106–126 (with a mean of 117.6) plicae or oblique, lateral skin folds. The pectoral fin-rays number 11–16. Canadian small inshore Sand Lances with deep bodies and low vertebral counts are the Pacific Sand Lance. The overall colour is silvery to silver blue on the sides, although it is less silvery than A. dubius, with a white belly. The sides may have a steel-blue iridescent stripe. The back may be grey, olive, brown, blue green, or metallic blue. In preservative this species is dark on the back and has a dark stripe on the flank, separated by a light stripe. The species reaches 30.0 cm in total length.

Ammodytes hexapterus



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Ammodytes hexapterus

habitat: Unlike its relative, this species can be found inshore in

shallow waters but also frequents offshore banks. Larger fish are found offshore. They may be found intertidally down to perhaps 275 m, but usually to 100 m. Temperatures may reach 14°C. In Hudson and James Bays it uses estuaries on a seasonal basis. In the Nelson River estuary of Hudson Bay it is found at depths of 3–6 m and salinities of 15‰–20‰. Pacific Sand Lances occur in large schools containing millions of fish but are also found as individuals buried in sand, even between high tide and low tide with the head occasionally protruding from the sand. It may aestivate buried in sand. It is the most numerous fish in the low-salinity Richmond Gulf, found in both schools and buried in sand at the water’s edge. Large numbers are found there dead, apparently stranded as the water recedes. It is common but not abundant in the Canadian Beaufort Sea.

biology: Its food is principally copepod crustaceans taken near

the bottom or in the water column, both by day and by night. Other crustaceans (such as mysids, ostracods, amphipods, cumaceans, isopods, and small crabs), arrow worms, snails, and polychaete worms are also eaten. Pacific Sand Lances are eaten by a wide variety of

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fishes, by whales, and by sea birds such as Thick-billed Murres in Digges Sound, Hudson Bay, and at Coats Island, Nunavut. They are eaten by Brünnich’s Murres at Akpatok Island in Ungava Bay. In coastal areas of the La Grande River, James Bay, this species is eaten by Brook Trout, and in the estuary by Lake Sturgeon, Ciscoes, and Lake Whitefishes. In the Eastmain River of James Bay it is eaten by Brook Trout and Walleye. In Richmond Gulf it is eaten by Fourhorn Sculpins and Ogac and is the most frequently occurring food item in Brook Trout stomachs. In Wakeham Bay, Ungava, it is eaten by Arctic Cod. On the Yukon coast it has been recorded from Arctic Cisco stomachs. Near the Nuvuk Islands, Nunavut, this species is fed by parents to Black Guillemot chicks and at Akpatok Island to Thick-billed Murre chicks (1.33% of diet). Concentrations of predators build up as the Sand Lance schools burrow into sand for the night. This species lives up to 12 years and may mature at the end of the first year of life. Spawning occurs inshore from October to January. At Kuujjuarapik in southeastern Hudson Bay, Sand Lances that may be this species appeared as yolk sac larvae from April to June. In the Nelson

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River estuary small larvae (5–8 mm) can be found in August, suggesting a later hatching. Up to 5,196 demersal, adhesive eggs are laid, which is perhaps a low estimate, with a maximum diameter of 1.1 mm.

importance: It is not economically important in the Arctic but

elsewhere has been used dried and salted or frozen, eaten fried, and used as fish-meal.

distribution: It is found in Davis and Hudson Straits, Ungava

Family Trichiuridae Cutlassfishes, Sabres de mer

Brian W. Coad

Bay, Hudson and James Bays, Gulf of Boothia, Melville Sound, Amundsen Gulf, and the Beaufort Sea, and from the Labrador coast south through Atlantic Canada to Virginia. It is also found in the Alaskan Beaufort Sea, Chukchi Sea, and Bering Sea, and south to Japan and California, including the coast of British Columbia.

Distribution of Ammodytes hexapterus

sources: Johansen (1927a); Tuck & Squires (1955); Edwards

(1961); Kidd et al. (1975); Greendale & Hunter (1978); Dutil & Power (1980); Morin et al. (1980); Richards (1982); Gaston, Cairns, Elliot, et al. (1985); Bond & Erickson (1987, 1993); Gaston (1987); Gaston, Elliot, et al. (1987); Baker et al. (1993); Ponton et al. (1993).

Cutlassfishes, Scabbardfishes, Hairtails, or Frostfishes are found in the Arctic, Atlantic, Indian, and Pacific Oceans. There are 32 species, including 10 species on Canadian coasts, of which 1 is in Arctic waters. The maximum size is about 1.5 m. They have a very long and very compressed body, ribbon-like in shape. There are fang-like teeth in a large head. The snout is pointed, and the lower jaw projects. The dorsal fin is extremely long and contains anterior spines usually separated by a notch from the posterior soft rays, but the spines and soft rays are difficult to distinguish. The anal fin is low and poorly developed compared to the dorsal fin. It is preceded by both a small and a large spine. The caudal fin is small and forked, or absent, in which case the body tapers to a point. The pelvic fin is reduced to a scale-like spine and 0–2 soft rays or is absent. The gill cover is splintered. There may be as many as 192 vertebrae. There are no scales. There is only one nostril on each side. The gill rakers are spiny. The overall colour is silvery to black with an iridescent sheen. These large fishes live mostly in a deep-water habitat and may be washed ashore or found in the stomachs of marine mammals and large fishes. As a result, the specimens are often damaged and difficult to identify. They may descend to 2,000 m or rise to 10–20 m. Cutlassfishes are benthopelagic, fast-swimming, voracious predators on smaller fishes and on shrimps and squids. They spawn year round in warmer waters and produce pelagic eggs and larvae. Some species are economically important in the eastern Atlantic Ocean but not in Canada.

sources: Tucker (1956); Parin & Bekker (1972); Nakamura & Parin (1993); Parin (1995).



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Aphanopus carbo Lowe, 1839

Black Scabbardfish, aphanope charbon

common names: A local name is Sort Sabelfisk (Danish/Green-

landic). Another common name is sabre noir.

taxonomy: The genus comes from the Greek aphanes (hid-

den, invisible) and pous (foot). The species name is the Latin carbo (charcoal). Aphanopus minor Collett, 1887, described from the east coast of Greenland, is a synonym.

variously given as 12–14 years, or as exceeding 40 years, with maturity attained at 80–85 cm. Spawning occurs year round, and the eggs are pelagic.

importance: It is a valuable food fish in some parts of the world, is often taken as a by-catch, and is sometimes fished specifically, as at Madeira, by traditional hook-and-line gear.

distribution: This species is found worldwide, including the southern Davis Strait as two records off southern Baffin Island from Karrer (1973) at 62º44' N, 60º54' W, and from cruise data at 68.3119º N, 59.6468º W; southwest and southeast Greenland; and on the Atlantic and Pacific coasts of Canada.

description: This species is distinguished by the ribbon-like

shape (a highly compressed body) and by the second enlarged anal fin spine being dagger shaped. The dorsal fin spines number 34–42, soft rays 50–57, the fin partly divided by a deep notch; the anal fin has 2 spines and 42–55 soft rays, the spines being separate from the rest of the fin; and the pectoral finrays number about 12. There is a pelvic fin composed of a single spine in young fish, but this is lost in adults. Live fish are coppery to coppery black, with an iridescent sheen, but this fades to black when they are dead. The oral and gill cavities are black. Fishes caught by trawlers often have their delicate skin scraped off and appear white. The oral and gill cavities are black. The species reaches 145.0 cm in total length and probably larger.

habitat: It is usually found swimming above the sea bed (benthopelagic) but may approach the surface at night. They have been caught at 640–661 m in Davis Strait and reported elsewhere at 146– 1,700 m. Young fish are mesopelagic. biology: Its food includes squids, fishes, and crustaceans such as

shrimps. Black Scabbardfishes chase squid, and a modification of the gas bladder may allow rapid pursuit into shallow water without expansion of the gas bladder gases. The gas bladder has a thick wall and is enclosed in a cage of ribs and muscle masses. The lifespan is

Distribution of Aphanopus carbo

sources: Templeman & Squires (1963); Karrer (1973); Biscoito et al. (2011); FAO (2011).

Aphanopus carbo

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Peprilus triacanthus

Family Stromateidae

(Peck, 1804)

Butterfish, stromatée à fossettes

Butterfishes, Stromatées

Brian W. Coad

Peprilus triacanthus

Butterfishes are found in tropical to temperate seas worldwide. There are about 13 species, including 2 species in Canada, 1 of which occurs in the Canadian Arctic. The maximum size exceeds 122 cm. The body is very deep, oval, and compressed. Adults lack pelvic fins, but the young of some species have pelvic fins. The dorsal fin is continuous and similar in size and shape to the anal fin, with the anterior end higher. Some species have 5–10 small, blade-like spines in front of the dorsal fin. The pectoral fins are elongate and pointed. The scales are minute, easily detached, and cycloid. Scales cover the dorsal and anal fin bases. The eye is surrounded by fatty tissue, forming an adipose eyelid. These fishes have toothed, sac-like outgrowths of the gut. If the gill cover is lifted, the sacs may be seen behind the last gill arch. The overall colour is silvery with grey to blue or green on the back and upper flank; some have flank mottling and spots. These are pelagic fishes often found in large schools near the coast. Young fish are found under floating debris and seaweed or associated with jellyfish and can drift with currents. Butterfishes secrete large amounts of mucus from an extensive canal system that may protect against jellyfish stings by inhibiting discharge of stinging cells or by counteracting the toxins produced. Some species are economically important.

common names: Other common names are American

description: This species is distinguished by the sac-like outgrowths of the gut, the short forward-pointing spine in front of the dorsal fin, and general body form. The dorsal fin has 2–4 (usually 3) spines and 40–48 soft rays; the anal fin has 2–3 (usually 3) spines and 37–44 soft rays; and pectoral finrays number 17–22. Lateral-line scales number 96–105. There are no pelvic fins in the adult. The spine in front of the dorsal fin may be concealed by skin. There are 17–25 large pores below the anterior half of the dorsal fin. The teeth at the front of the upper jaw have three small cusps. The colour varies from bluish to greenish to dark grey on the back and upper flank. The lower flanks and belly are silvery, the flanks usually having irregular dark spots dorsally that fade after death. The species reaches 30.5 cm in total length.

sources: Haedrich (1967); Horn (1970).

habitat: The Butterfish is found inshore over sandy or mud

Butterfish, Atlantic Butterfish, Dollar Fish, Pumpkin Scad, and Sheepshead.

taxonomy: The genus comes from peprilos (a fish from Thrace). The species name comes from the Greek treis and the Latin tri(three) and akantha (spine or thorn).

bottoms during May to September in northern Atlantic waters, retreating offshore in winter down to 420 m. It can live in brackish water and is tolerant of a wide range of salinity and temperature. The young are found with jellyfish, such as Cyanea and Chrysaora, in weeds or swimming alone. A single Canadian Arctic record of a young specimen was taken in Davis Strait over a bottom depth of 976 m.

biology: The young feed on jellyfish but also on other fishes,

squid, arrow worms, crustaceans, and worms. Adults eat urochordates and thecosome molluscs (Clione), as well as cnidarians, comb jellies, arrow worms, and crustaceans. Spawning occurs from



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Peprilus triacanthus

May to July in Atlantic waters, and the eggs measure 0.7–0.8 mm in diameter. Most are mature at two years, and they live up to six years. Many economically important fishes feed on Butterfish in more southern waters.

importance: It is economically important in the northwestern

Atlantic Ocean, and over 460 million fish or 19,454 t were caught in 1973, but it is not important in Canadian waters. It may be eaten fresh or smoked and is exported frozen to Japan.

distribution: A single record from Davis Strait at 62.7667° N, 63.2667º W (ARC 8704258), is presumably an expatriate specimen. It is found from eastern Newfoundland and the Gulf of St Lawrence south to the Gulf of Mexico.

Distribution of Peprilus triacanthus

sources: See the family sources and the bibliography.

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Family Pleuronectidae Righteye Flounders, Plies

Noel R. Alfonso

Righteye Flounders are north temperate fishes, found in the Arctic, Pacific, and Atlantic Oceans, with some entering brackish water. They are known as dabs, flounders, plaices, turbots, and, incorrectly, soles, some of these names being used for the same species. There are 5 subfamilies, 21 genera, and about 60 species, including 31 species in Canada, of which 9 are in the Arctic. There have been various recent taxonomic changes in which certain genera (Limanda, Liopsetta, Pseudopleuronectes) have been lumped under Pleuronectes. These changes have not been accepted by all workers. Of the nine species known from the Canadian Arctic, Pleuronectes glacialis is most characteristic of Arctic and sub-Arctic waters. Reinhardtius hippoglosoides has the most extensive range of the Arctic flatfishes, is concentrated in Davis Strait, Ungava Bay, and Hudson Strait, but is also found off Banks Island in the west and as far north as northern Ellesmere Island. Flatfishes (Pleuronectiformes) are a unique order of fishes in which the larva metamorphoses as it develops so that one eye migrates across the head due to differential development of cranial bones. As a juvenile the fish settles on the bottom and then, as an adult, lies on either its right or its left side. Species that have their right side uppermost are called dextral (right-eyed); those that have their left side uppermost are sinistral (left-eyed). The lower, or blind side, is usually non-pigmented. The Righteye Flounders are medium to large (maximum total lengths ranging from 30 cm to 270 cm), mainly piscivorous species found in temperate waters of the northern hemisphere. Almost all the species are benthic. They have eyes on the right or eyed-side, but some species may have left-eyed individuals in a population, and some populations may be all left-eyed fish. The teeth are strongest on the blind side since these fish mostly feed on the bottom. The pectoral fin is largest on the eyed or upper side where it can move more freely. The fins lack spines, and the dorsal fin origin is above or anterior to the eye. The dorsal and anal fins run the length of the body. The pelvic fins are symmetrically placed on each side of the body. The scales are often ctenoid on the eyed



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side and cycloid on the blind side but may be ctenoid or cycloid on both sides. The egg yolks usually lack an oil globule. Flatfishes are a successful group in part due to their utilization of extensive coastal and continental shelf habitat. Species in Canadian Arctic waters can be classified as shallow, coastal, or deep species. All these species are exclusively benthic, with the exception of the Greenland Halibut and occasionally the Atlantic Halibut, which are known to feed in schools in mid-water. The smaller species eat invertebrates and clam siphons; the larger species feed on fishes. Typically they conceal themselves with a layer of sand or mud that they disturb by flapping their fins. Some may migrate long distances. These fishes take in water through the mouth, passing it over both gills for oxygen extraction, and ejecting it only on the upper or eyed side. A special chamber connects the two sets of gills, allowing water that would normally exit on the blind side to be transferred to the eyed side. Flatfishes have the ability to change colour quickly and match their surroundings for concealment from predators and prey. Individuals placed on a chess board make a valiant attempt to match the black and white squares. This family contains many economically important species. Seven of the twelve most commonly encountered fish species in surveys of the Bering and Chukchi Seas are flatfishes.

sources: Norman (1934); Sakamoto (1984); Cooper & Chapleau (1998); Evseenko (2004).

Glyptocephalus cynoglossus (Linnaeus, 1758)

Witch Flounder, plie grise

common names: A local name is Skærising (Danish/Greenlandic). Other common names are Craig Fluke, Gray Sole, Pale Flounder, Pole Flounder, White Sole, Witch, and flet. taxonomy: The genus comes from the Greek glyptos (sculptured, carved) and kephale (head), referring to the mucus pits on the underside. The species name comes from the Greek kynos (dog) and glossa (tongue), in reference to the elliptical body shape; in Latin cynoglossus refers to a sole. Glyptocephalus acadianus Gill, 1873, described from Eastport, Maine, is a synonym. description: This species is characterized by up to 16 mucous

pits (blister-like cavities or craters) on the blind side of the head and by a distally black, eyed-side pectoral fin in adults. The body is very thin and narrowly elliptical; posterior to the head, the dorsal and anal margins of the body form almost straight lines to the caudal peduncle. The caudal fin is rounded. Mucous pits are also found on the eyed side but are much less distinctive than those

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Glyptocephalus cynoglossus

on the blind side. Both the head and the mouth are very small, the head making up a fifth of the total body length. The dorsal profile is noticeably convex. The lateral line is straight with a slight upward curve over the pectoral fin in some fish. The pectoral fin is short. The dorsal fin-rays number 95–120, the anal fin-rays 82–102, the pectoral fin-rays 9–13, and the vertebrae 36. The anal spine projects in front of the anal fin, although sometimes it is very small and buried in the skin. The cycloid scales number 110–140 along the flank above the lateral line. The gill rakers number 6–9 on the lower arch. The colour of the eyed side is reddish brown, brownish, or greyish brown. The body and the fins have numerous minute black dots on the eyed side; the black dots on the blind side body and fins are larger and more randomly spaced. Indistinct blotches are evenly spaced on the margins of the dorsal and anal fins on the eyed side. Unlike most flatfishes, the blind side is only slightly lighter than the eyed side. The eyed-side pectoral fin has dusky to black membranes with a thin, light, distal margin. The species reaches a maximum of 78.1 cm in length and 5.0 kg in weight, but specimens over 60.0 cm and 2.5 kg are now uncommon.

habitat: The species is distributed in the deeper, cold waters

of the North Atlantic. It is found mainly in deep water and offshore at 18–1,569 m, mainly at 46–274 m, and from −1°C to 13°C on mud, mud-sand, and clay, often in channels and deep holes. It has

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been caught in Davis Strait at 685–939 m and at 847–1,295 m and 3.5°C–3.7°C. Deep-water populations have been found at 273–1,371 m in southern New England slope waters and are somewhat distinct from populations in the usual depth range; they differ in growth and maturation rates. The larvae are pelagic (0–36 m), juveniles are found at 180–288 m, and adults in the summer are found at 36–270 m and in the winter at 90–432 m. Adults are found in deep-water concentrations in the winter. They are uncommon in waters warmer than 10°C.

biology: The small body and mouth size of this species means that

dietary items are restricted to polychaetes, crustaceans, snails, molluscs, brittle stars, and small fishes. A wide variety of polychaetes (23 families) form 59.0% of the diet by weight, bivalves 20.4%, and crustaceans 13.9% in a deep shelf trough of the North Sea. On Georges Bank, Witch Flounder fed intensively on polychaetes, which made up 75%–91% of the total diet; crustaceans, by contrast, comprise 2%–11%. It is eaten by Thorny and Smooth Skates, Spiny Dogfish, White Hake, Greenland Halibut, and Goosefish, as well as by Harp Seals. This species is slow growing and long lived to over 36 years. In the Gulf of St Lawrence the mean length at age 12 is 40 cm (males) and 41 cm (females). These slow growth rates make the species vulnerable to over-exploitation. Juveniles occupy different depths at

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different seasons from adults, due to differential prey availability. Females are longer lived than males, grow larger, and exhibit faster growth. Maturity is generally attained at 25–30 cm in length and 4.2–6.0 years of age for males, and at 40–50 cm and 8.4–10.2 years for females. In winter months Witch Flounder populations are found in dense pre-spawning concentrations in deep, relatively warmer water. Spawning occurs from spring to summer, depending on the area. In the eastern Newfoundland and Labrador region intensive spawning occurred during March–June. In the more westerly areas spawning occurred intensively during January–February. Spawning occurs in deep waters, sometimes in channels. Females have a high fecundity, producing up to 3 million floating eggs of ca. 1.45 mm in diameter. The pelagic stage of up to one year is the longest among pleuronectids, and the size at metamorphosis is correspondingly large, over 50 mm. This species is sedentary, with no significant migration reported. It is parasitized by 3 protozoans, 13 trematodes, 5 nematodes, 2 acanthocephalans, and a copepod and by the sealworm (Phocanema decipiens). Exposure to polycyclic aromatic hydrocarbons (PAHs) or polychlorinated biphenyls (PCBs) in Witch Flounder and other flatfish species has led to a higher prevalence of lesions, a significantly lower condition factor and lymphocyte levels, and elevated hepatic and splenic hemosiderosis (iron metabolism disorder), compared to reference sites.

importance: The largest catches in Canadian waters are taken at

2°C–6°C and 185–366 m. Reported catches of Witch Flounder in the northwest Atlantic by multiple countries peaked at 38,000 t in the 1980s and steadily decreased to 3,000 t in 2009. As a slow-growing, long-lived species, the Witch Flounder is vulnerable to exploitation. This species was economically important in the late 1950s and early 1960s as a by-product of Haddock fisheries on the Grand Banks. Later it became an important by-product of the Atlantic Cod, American Plaice, and redfish fisheries. Landings now remain at about 2,000 t per year. The highest catches are now on the Scotian Shelf and in the Gulf of St Lawrence. Decades of fishing pressure have reduced the average size and age of Witch Flounder. The flesh is white and ranks highly among flatfishes as a food fish. This flounder is well regarded by anglers. Witch Flounder abundance remains low, under moratoria or regulatory limitations. In the Gulf of Maine these catches are minor. Landings in the Gulf of St Lawrence and western Newfoundland area in 2011 were 442 t. That fishery is now a directed seine fishery.

distribution: It is found in Davis Strait. Glyptocephalus cynoglossus is an accidental Atlantic stray to the Arctic. There has been no evidence of spawning in Arctic waters. It is also found in southwest and southeast Greenland and in both the eastern and the western Atlantic. The range extends south to Georgia in the west, and to Iceland, the Faroe Islands, and northwestern Europe to the Barents Sea. In eastern Canada the distribution encompasses the Gulf of Maine, the Scotian Shelf, the Gulf of St Lawrence, and the Grand Banks, with fewer occurrences north to Labrador.



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Distribution of Glyptocephalus cynoglossus

sources: Mansfield (1967); Powles & Kennedy (1967); Evseenko

& Nevinsky (1975); Bowering (1990); Burnett, Ross, & Clark (1992); Wrigley (1998b); Bowman, Stillwell, Michaels, & Grosslein (2000); Skjæraasen & Bergstad (2000); Rabe & Brown (2001); Bergstad, Wik, & Hildre (2003); Khan (2003); Robson, O’Dywer, & King (2003); Wigley & Burnett (2003); Jørgensen et al. (2005); FAO (2011); Department of Fisheries and Oceans (Swain) (2012).

Hippoglossoides platessoides (Fabricius, 1780)

American Plaice, plie canadienne

common names: Local names are Okôtak and Oquutaq (Green-

landic). Other common names are American Dab, Blackback, Canadian Plaice, Dab, Flounder, Long Rough Dab, Rough Dab, Sand Dab, Sole, and faux flétan.

taxonomy: The genus comes from the  Greek ippos (horse) and glossa (tongue) and the Latin -oides (like, resembling). The species name comes from the Latin platessa (plaice) and the Latin suffix -oides (like, resembling). In the western Atlantic this plaice is regarded as a subspecies, Hippoglossoides platessoides platessoides, and in the eastern Atlantic as Hippoglossoides platessoides limandoides (Bloch, 1787).

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Hippoglossoides platessoides

description: This species is characterized by a large mouth,

extending to or past the mid-eye on the eyed side; the teeth are pointed and about equal in extent in both jaw halves; the lateral line on the eyed side is straight and is gently bent above the pectoral fin; the scales are ctenoid on the eyed side; the caudal fin is not emarginate (the middle rays being longest); the dorsal fin-rays number 76–101, usually more than 80; the anal fin-rays number 60–79; and the branchiostegal rays number 8. The American Plaice has a projecting lower jaw, often with a knob on the chin. The head is moderate in size, a third or a quarter of body length. The eyes are large, the upper eye being larger than the snout. The interorbital space is narrow with a raised, scaled ridge. There is a strong pre-anal spine present. There are 9–12 gill rakers on the lower branch. Cycloid scales are present on the blind side, with ctenoid scales on the rear of the body and fin bases. The lateral line has two accessory branches continuing to the upper eye. The curves of the posterior dorsal and anal fins are convex. The pectoral fin is short, less than half head length, and is longer and more rounded on the eyed side, with 1–2 more rays. The caudal peduncle is as long as it is deep. The caudal fin is broad and distinctly rounded. The pectoral rays number 9–12, the lateral-line scales 85–97, and the vertebrae 42–48. The eyed side is russet or reddish brown to grey brown with occasional darker spots of varying sizes, and the blind side is white

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or bluish white. The fins are paler than the body. The margins of the dorsal and anal fins are white. The young have 3–5 dark spots on the dorsal and anal margins of the body. As with all pleuronectid flatfishes, some specimens are partly dark on the blind side. The species reaches 82.6 cm in length and 6.4 kg in weight, but averages 38–40 cm and 0.9–1.4 kg.

habitat: American Plaice are continental-shelf, deep-water flatfish

found in a depth range of 18–3,000 m, on sand or muddy bottoms, usually at temperatures from −1.5°C to above 5°C and as high as 13°C. They are found mostly at depths of 50–250 m but may enter brackish areas of rivers. Arctic depth ranges include 140–375 m in Hudson Strait, and 241–3,000 m in Davis Strait and southern Baffin Bay at −0.1°C to 5.4°C. In the Gulf of St Lawrence, American Plaice densities peaked at 70–90 m, being closely linked to temperature. On the Grand Banks of Newfoundland the species was most abundant at 91–140 m (from −0.5°C to 1.0°C) and at 183–229 m (from 1°C to 2°C). Research surveys of the Grand Banks recorded American Plaice at 100–200 m (from −1.1°C to −0.5°C) during the summer and autumn. Their preferred habitat is a fine gritty mix of sand and mud that is characteristic of the Atlantic continental shelf. Despite that preference, they have been known to feed heavily on urchins on bedrock bottoms.

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Hippoglossoides platessoides

Females are consistently found in warmer water than are males in September. Both sexes move away from cold inshore waters to deeper water in winter to avoid subzero temperatures. Newly settled juveniles and young fish can be found in estuarine environments where coastal bays are near offshore banks in Canadian waters, although this habitat use has not been documented in New England. Although they are bottom dwellers, American Plaice have been captured in the water column, moving off the bottom at night. These movements relate to the diurnal movements of Sand Lance, a prey species that remains buried in sand during the day and rises into the water column in the evening hours to feed on plankton.

biology: The broad distribution of the American Plaice results

from its great adaptive abilities, particularly its feeding habits. This species is an opportunistic predator with the ability to change its niche depending on prey availability and interspecific competition. Variation in adult feeding is due to seasonal availability and distribution of prey items. Young American Plaice feed on polychaetes, crustaceans, and cumaceans, making the change to echinoderms, molluscs, and fishes as they become larger. Adults have been known to feed heavily on the green sea urchin  (Strongylocentrotus droebachiensis) in the southwestern Gulf of St Lawrence and eastern Newfoundland. In the winter, American Plaice in the Gulf of St



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Lawrence do not feed at all or somewhat minimally. On the southern portion of the Grand Banks, Sand Lance is the predominant fish prey species consumed in summer months; brittle stars are consumed in the winter. On Sable Island Bank a variety of crustaceans, including cumaceans, hyperiid, and gammarid amphipods, mysids, and pagurids make up the bulk of the diet. In the Barents and Norwegian Seas there is a large range of fish species eaten, including Clupea harengus, Mallotus villosus, Gadus morhua, Boreogadus saida, Melanogrammus aeglefinus, stichaeids, Ammodytes, Sebastes, Triglops, and young American Plaice. American Plaice distribution is very similar to that of the northern Atlantic Cod stock, and the two species compete with each other, markedly so as juveniles; larger cod prey on American Plaice. Predators of American Plaice include large piscivorous fishes, such as Goosefish, Atlantic Cod, White Hake, Spiny Dogfish, Sea Raven, Thorny Skate, Acadian Redfish, and Halibut (in order of stomach-content occurrences). They are intensively eaten by large cod in the Magdalen Shallows. They are also eaten by Greenland Sharks at higher latitudes. Seal predators include Gray and Harp Seals. Its lifespan is up to 30 years. Sexual maturity is attained at age 6 (25 cm) for males and age 10 (41 cm) for females in the Magdalen Islands area. Half of the females from the Flemish Cap reach maturity at 7.8 years, while for males the range is 5.3–7.5 years. Growth is slower in

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colder, northern waters. Spawning individuals move inshore from deeper water to spawn. Spawning follows a north–south gradient, occurring from as early as March and ending in June in the northwest Atlantic in water temperatures of −1.3°C to 3.5°C, and ending in July and August in populations off Labrador and west Greenland. Individuals generally return to the same areas to spawn. The pelagic eggs lack an oil globule but are buoyant, having a characteristic large perivitelline space. Eggs number up to 2.2 million and are 2.8 mm in diameter. They hatch from 11 to 14 days at 3.9°C. Larvae hatch at 2.4 mm and transform at between 6.2 and 7.5 mm. Larval settling occurs at 30–40 mm. Numerous species of parasites have been documented in American Plaice, including 2 species of protozoans, 2 copepods, 7 nematodes, 3 acanthocephalans, and 2 cestodes.

importance: American Plaice is the most abundant flatfish in the

northwest Atlantic and is one of four major species of flatfishes in Canadian Atlantic fisheries. Plaice are very important economically in Canada and comprise about half of all flatfish landed. They are sold as fresh or frozen fillets with white, tasty flesh. They are harvested by offshore otter trawls, seine nets, and long lines. Until 1948 the landings were minor, but they reached 27,215 t in 1960 and 60,521 t in 1985. Landings from 1999 to 2003 were 4,384 t, 3,924 t, 3,834 t, 3,120 t, and 3,004 t, respectively. There has been an observed decline in this stock. Recruitment, abundance, and total and spawning biomass remain low. Declining trends in American Plaice abundance resulted in the Committee on the Status of Endangered Wildlife in Canada assessing the Maritime and Newfoundland and Labrador populations as “Threatened,” and the Arctic population as “Data Deficient.” By-catch off Labrador from the Greenland Halibut fishery has increased since 1999. The average size and age at maturity declined through the mid-1990s. Stock-rebuilding prospects remain extremely poor. Attempts to reduce American Plaice by-catch from the long-line cod fishery have thus far been unsuccessful. There is no documentation of the Plaice being caught by anglers as its range is generally offshore and at great depth.

distribution: It has been reported from Baffin Bay, Davis Strait, Frobisher Bay, eastern Hudson Strait, Ungava Bay, and eastern Hudson Bay and is found on all coasts of Greenland. The American Plaice has an Arctic boreal distribution and is found on both sides of the Atlantic. The European subspecies (H. p. limandoides) is encountered off Iceland, around the Faroe Islands, around the British Isles, in the North Sea, and to the western part of the Baltic Sea. The North American subspecies (H. p. platessoides) is found from Greenland and Baffin Bay south to Delaware.

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Distribution of Hippoglossoides platessoides

sources: Government of Newfoundland and Labrador (n.d.); Pitt

(1963, 1966, 1967, 1989); Powles (1965); Beamish (1966); Fahay (1983); Margolis & Kabata (1984–96); Macdonald & Green (1986); Benoit & Bowen (1990); Hudon (1990a); Keats (1991); Khan, Lee, & Whitty (1991); Zamarro (1992); Køie (1993); Ponton et al. (1993); Lawson, Stenson, & McKinnon (1994); Lile, Halvorsen, & Hemmingsen (1994); Martell & McClelland (1994); Walsh (1994); Berestovskiy (1995); Swain (1997); Swain & Morin (1997); D.L. Johnson, Berrien, Morse, & Vitaliano (1999); Kulka & Tillman (2000); Rountree (2001); González, Román, & Paz (2003); D.B. Stewart & Lockhart (2004, 2005); Jørgensen et al. (2005); Osborne, Treble, Dueck, & Cosens (2005); González, Paz, Román, & Hermida (2006); Rodger (2006); Dwyer, Morgan, Parsons, Brodie, & Healey (2007); Busby et al. (2008); Morin, LeBlanc, Chouinard, & Swain (2008); COSEWIC (2009); Department of Fisheries and Oceans (Morgan) (2011).

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common names: Local names are Nadalna, Natanak, Natarnak, and Natarnaq (Inuktitut).

Hippoglossoides robustus Gill and Townsend, 1897

taxonomy: The species name is the Latin robustus (robust). Synonymy of this species with H. elassodon Jordan and Gilbert, 1880 (Flathead Sole, plie à tête plate), by some authors is based on the difficulty in distinguishing the two species in the field and on molecular evidence. However, genetic and anatomical evidence warrant its place as a distinct species.

Bering Flounder, plie de Béring

description: This species is characterized by a large mouth,

extending to or past the mid-eye on the eyed side; the teeth are pointed, about equal in extent in both jaw halves; the lateral line on the eyed side is straight or gently bent above the pectoral fin; the scales are ctenoid on the eyed side; the caudal fin is not emarginate (the middle rays being the longest); the dorsal fin-rays number 66–80, usually less than 80; the anal fin-rays number 51–64; and the branchiostegal rays number 7. The upper eye is lateral, not on the head margin, and is larger than the fixed eye. The interorbital space lacks a crest but has 2–4 scale rows. The body is markedly ovate, and the dorsal profile is concave above the eye. The body is thick. The scales on the head are separate and rarely touching. The nasal tubes are very short. The scales are ctenoid or cycloid on the blind side. The anal spine is pronounced. The pectoral fin-rays number 8–12, and the pelvic fin-rays 6. The gill rakers on the lower arch are 9–13, with the total count being usually Hippoglossoides robustus

Hippoglossoides robustus



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less than 17. There are 85–95 lateral-line scales. The vertebrae number 44–47. In colour the eyed side is a light, brownish yellow, reddish brown, or greyish brown, with reddish- to greyish-brown speckles and large dark spots along the dorsal and anal fins and on the mid-body. The dorsal and anal fins are not blotched. The caudal fin membranes are clear but are sometimes smudged. The blind side is white with a silvery sheen mainly below the lateral line. The species attains 30.0 cm in length, to 52.0 cm in Alaskan waters.

habitat: The Bering Flounder is found in deep, cold water from 18

m to 530 m and on soft bottoms, usually shallower than 150 m. Bering Flounder are found in the Bering Sea, but in very low numbers and densities.

biology: Fish of 6–13 years of age and 14.4–25.5 cm length are

mature. Spawning occurs in summer in shallow water. Arctic spawners may have low success as their eggs can be carried into unsuitably cold areas by currents. Growth is slow in such waters, with Bering Flounders taking 11 years to reach 23 cm. The Bering Flounder was the most abundant pleuronectid by numbers in 1991, and the second most abundant in 1990, in experimental trawls south of St Lawrence Island. In some years it is one of the three or four most abundant fish species in the Chukchi Sea.

Distribution of Hippoglossoides robustus

importance: Hippoglossoides species (both Bering Flounder

sources: Fadeev (1986); Allen & Smith (1988); Spencer, Walters,

and Flathead Sole) are managed as a unit stock in the Bering Sea and Aleutian Islands and were formerly a constituent of the “other flatfish” catch component. This species is too small and rare to be economically important in Canadian waters.

distribution: The two Canadian Arctic records in the Bathurst Inlet and Melville Sound area (CMNFI 1966-0076 at 67°51'48" N, 107°50' W; and CMNFI 1976-0119 at 68°20' N, 107°41' W) may be strays; more sampling in the Beaufort Sea should show if this is part of a continuous population. Thirty-two specimens were trawled over a wide area of the Alaskan Beaufort Sea. This is an Arctic and boreal Pacific species found from the northern Sea of Japan off Hokkaido, the Sea of Okhotsk, the East Siberian Sea to the Chukchi Sea, north off Akutan Island in the Alaska Peninsula, and the western Bering Sea.

& Wilderbuer (2004); Mecklenburg et al. (2007); Kartavtsev et al. (2008); Cui et al. (2009); Rand & Logerwell (2011).

Hippoglossus hippoglossus (Linnaeus, 1758)

Atlantic Halibut, flétan atlantique

common names: A local name is Nataarnaq (Greenlandic).

Other common names are Common Halibut and Giant Halibut. Names for colour variants are Cherry Bellies and Grays or Greys for individuals having the blind side mottled with red or grey, respectively.

taxonomy: The genus and species names come from the Greek ippos (horse) and glossa (tongue), in reference to the halibut shape. At one time this species was thought to be in both the North Atlantic and the Pacific oceans, but the Atlantic Halibut is now recognized as distinct from the Pacific Halibut (H. stenolepis Schmidt, 1904). This divergence occurred in the Pliocene, 1.7–4.5 million years ago. Synonyms are Hippoglossus vulgaris Fleming, 1828, described from rivers and estuaries of England; Hippoglossus gigas Swainson, 1839 (no locality given); and Hippoglossus americanus Gill, 1864, described from the eastern coast of North America.

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Hippoglossus hippoglossus

description: This species is characterized by a large mouth

extending to or past the mid-eye on the eyed side; the teeth are pointed, being about equal in extent in both jaw halves; the lateral line on the eyed side is strongly arched above the pectoral fin; the scales are cycloid on the eyed side; and the caudal fin is obviously emarginate. The Atlantic Halibut is the largest Atlantic flatfish. The body is thick but narrow. The head is large with a very large and nearly symmetrical mouth. The eyes are noticeably separate, with a broad flat area between the two. The upper eye can project slightly above the head margin. The dorsal fin originates above the upper eye, elongating to its greatest length halfway along the body, then tapering to the caudal peduncle. The anal fin originates at the pectoral fin, reaching its maximum length midway or more posteriorly along the body. The dorsal fin-rays number 92–110, the anal fin-rays 69–85, and the vertebrae 50–51. There is a strong anal spine, sometimes hidden by skin in older fish. The lateral-line scales number about 160. The body is slimy with mucus. The eyed side is greenish to dark brown or black, and the blind side lacks pigmentation. There is usually some blotchy mottling on the eyed side. In New England waters they are described as being chocolate to olive or slaty brown, whereas in European waters they are uniformly dark brown or black. Young Atlantic Halibut are marbled or have spots with paler marks. The blind side of both adults and young is pearly white but may have grey mottles (hence “greys”) or red mottles (“cherries”). As with other flatfishes, partial or full ambi-colorate specimens are sometimes found. The species attains 250.0 cm in length and 330 kg in weight.



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habitat: Atlantic Halibut is a continental slope species found

at 37–2,000 m, usually at 160–230 m when adult but in shallower water when immature. In northern Davis Strait and southern Baffin Bay, Atlantic Halibut are found at 490–813 m at temperatures of 3.7°C–4.5°C, and down to 1,062 m in southern Davis Strait. They prefer temperatures above 2.5°C, entering shallower water in summer, where they are captured in directed fisheries, and retreating to deeper water in winter. The species is found on sand, gravel, and clay bottoms, not on mud or rock bottoms. Adults seldom enter water less than 60 m deep. It is benthic but sometimes is found in the water column, where it ascends for the purpose of feeding. This species prefers high salinities and is not generally found in estuaries. Its movements are still not well known. There is summer–winter movement in Davis Strait in which adults are likely seeking the fish prey that is more numerous on the shoals than off the continental shelves. There is an extensive movement of Atlantic Halibut throughout the Canadian portion of the North Atlantic, with younger fish moving more than adults. There is no distinct migration, but individual fish occasionally make long journeys. One fish was seen to have moved 2,575 km from Anticosti Island to Iceland over a seven-year period; others travelled 241 km and 805 km, but these were exceptional.

biology: The Atlantic Halibut is not a distinctly Arctic species as

it prefers cold boreal waters. It is one of four high-latitude pleuronectids characterized by deep-water habitat, having large eggs, a long development time, low mortality, and late reproduction. The Atlantic Halibut has been described as a voracious predator, generally on benthic or demersal fishes.

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Adult halibut over 80 cm in length feed actively, ascending into the water column to prey on a wide variety of fishes. Large Atlantic Halibut feed almost exclusively on other fishes, including Skates, Atlantic Herring, Capelin, Atlantic Cod, Grenadiers, Silver Hake, Sculpins, American Sand Lance, Wolffishes, Atlantic Mackerel, redfish, Pollock, Haddock, American Plaice, and Lumpfish. Halibut have been observed striking cod with their tail to stun them. As well, adults eat crabs, shrimps, lobsters, clams, mussels, and even seabirds. Smaller Atlantic Halibut feed on annelid worms and crustaceans such as hermit crabs, other crabs, and mysids almost exclusively until reaching about 30 cm, then gradually adding fish to their diet as they grow. In the northwest Atlantic there are three main groups in the diet: bony fishes (66% by weight), squid (18%), and crustaceans (15%). Predators of Atlantic Halibut include seals, Spiny Dogfish, Goosefish, and notably Greenland Sharks, which consume them regularly. Growth rates vary among populations within Canada and between stocks in the western and eastern Atlantic Ocean. Females exhibit faster growth than do males (also reaching larger maximum sizes), and the species grows faster than other Atlantic flatfishes. The maximum recorded age is 50 years. The average age of maturity is about 10 years; females reach maturity at 10–14 years, whereas males do so at 8–10 years. Size at maturity declined from 84 cm for males and 98 cm for females in 1959–69 to 66 cm and 70 cm in 1970–9 as a result of heavy fishing. Atlantic Halibut are annual, group-synchronous spawners. They home to the same spawning site in successive years. Spawning occurs during late winter and early spring, with peak spawning having been reported during November to December in Canadian waters. Spawning in the western Atlantic is believed to occur on the slopes of the continental shelf and on offshore banks, at depths of at least 180 m, over a rough or rocky bottom. In European waters, Atlantic Halibut spawn in December to April at 300–100 m at temperatures of 5°C–7°C on a soft clay or mud bottom. Off west Greenland, spawning occurs in late spring. Larger females over 90 kg produce up to 7 million eggs. With an egg diameter of up to 4.0 mm, the Atlantic Halibut has among the largest of planktonic fish eggs. The eggs are neutrally buoyant and sink slowly as they develop, floating at 54 m or as deep as 800 m. They hatch in 13–20 days with larval lengths of 6–7 mm. Larvae rise to the surface and drift into shallow inshore areas, in Atlantic Canada for example around the Sable Island Gully, Browns Bank, and the southern extent of the Gulf of St Lawrence, which may serve as nurseries. Parasites include 3 species of monogeneans, 10 digeneans, 1 isopod, 4 copepods, and 1 amphipod, as well as 1 external flagellate in Norway.

peaked at 6,000 t in 1950. Catches of Atlantic Halibut are now regulated by TAC (total allowable catch), which includes directed and by-catch. The TAC stood at 1,700 t for the Scotian Shelf and the Grand Banks in 2009. Annual by-catch from the Greenland Halibut fishery in Davis Strait is about 400 kg. The fishery is important in Canada, and halibut fetches a high price. Larger halibut are caught on long lines, and smaller ones by otter trawl. In 1994 a minimum-size limit of 81 cm was imposed for Atlantic Halibut in Canadian waters. It has been caught by anglers, and fights well, but is uncommon inshore. Atlantic Halibut are being grown in aquaculture facilities in Norway on a limited scale. In Canada experimental aquaculture is being done but is far from being economically viable.

distribution: The species is found in Baffin Bay, in Davis Strait, on both sides of the North Atlantic Ocean, and in portions of the Arctic Ocean. H. hippoglossus is an accidental Atlantic stray to the Arctic. In Europe it is found from the Bay of Biscay to the Barents Sea. Westward it is found in Iceland, in northwest, southwest, and southeast Greenland, at Uummannaq, and as far north as Upernavik (latitude 72.8° N) in western Greenland. Atlantic Halibut in the Arctic are much more numerous in Greenland compared to Canadian waters as the warm West Greenland Current extends its range northwards. On the Canadian side of Davis Strait the cold Labrador Current is a deterrent to this boreal species. Its range extends south through Atlantic Canada and the Gulf of St Lawrence to Virginia.

importance: In Canada, Atlantic Halibut commands the high-

est price of all the flatfishes. They are taken in bottom long lines, and most of these fish are sold in the United States. There is currently no directed fishery in the United States, and all landings occur as by-catch in groundfish fisheries. Atlantic Halibut were first commercially fished in the late-nineteenth century, and in the early part of the twentieth century 1,000–3,000 t were caught annually. From 1945 onwards, landings

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Distribution of Hippoglossus hippoglossus

sources: Fleming (1828); Swainson (1839); McCracken (1958);

W.S. Grant, Teel, Kobayashi, & Schmitt (1984); Fevolden & Haug (1988); Godø & Haug (1988); Stobo, Neilson, & Simpson (1988);

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J.M. Miller, Burke, & Fitzhugh (1991); Neilson, Kearney, Perley, & Sampson (1993); Trumble, Neilsen, Bowering, & McCaughran (1994); Cargnelli, Griesbach, & Morse (1999); Bowman et al. (2000); Jørgensen et al. (2005); Isaksen, Karlsbakk, & Nylund (2007); Department of Fisheries and Oceans (Armsworthy) (2009); Trzcinski et al. (2011); Department of Fisheries and Oceans (2013j).

Limanda proboscidea Gilbert, 1896

Longhead Dab, limande carline

head margin above the eye is strongly concave; and the lateral line is steeply arched over the pectoral fin; it is also distinguished by finray counts. This species has a long snout, and the jaws converge almost to a point, hence the common name of Longhead Dab. The head and snout become relatively longer and deeper in larger individuals. The body is deep and thick. The lower eye is noticeably in advance of the upper. The nostrils are widely separated. A rough crest is present behind the eyes, and, on the blind side, there is a corresponding rough area. The scales are moderately ctenoid or cycloid on the eyed side, and cycloid on the blind side. The dorsal and anal fins reach their greatest length posterior to the midline of the body, forming a diamond-shaped outline. The dorsal fin originates over the migrated eye. The anal spine is prominent, although at times it is small and buried in the skin. The caudal peduncle is short and widens posteriorly. The caudal fin is fan shaped, considerably widening posteriorly. The dorsal fin-rays number 61–77, the anal fin-rays 45–58, the pectoral fin-rays 9–13, and the pelvic finrays 6. The lateral-line scales number 73–95. The upper-arch gill rakers number 4–5, and the lower-arch gill rakers 8–11. Total vertebrae number 36–40. The coloration of the eyed side is a pale to olive grey brown with indistinct whitish speckles. The blind side is lemon yellow. Dark spots are scattered over the body. In some individuals the dark spots coalesce to form blotches on the dark side. There are fewer, lighter spots on the fins. The tips of some of the median rays are bright yellow. The species reaches 41.0 cm in length, more commonly 17.0 cm.

habitat: This species is found on soft bottoms at 10–125 m, very

rarely deeper, and mostly shallower than 50 m, in well-mixed areas with strong influences from coastal areas, river flows, and currents. It is an inner-middle-shelf or sublittoral species.

biology: Relatively little is known of the biology of the Long-

Limanda proboscidea

common names: Local names are Nadalna and Natanak

(Inuktitut).

taxonomy: Limanda is an old name for the European Dab, Limanda limanda, which in turn is derived from the Latin limus (mud). The species name comes from the Greek proboskis (snout) and the Latin idea (form), in reference to the long snout or proboscis. This species has also been placed in the genera Myzopsetta and Pleuronectes. description: This species is distinguished by a small mouth,

extending almost to below the anterior part of the eye on the eyed side; the teeth have a greater extent on the blind side of the jaws; the



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head Dab. Its diet consists of amphipods, benthic copepods, other benthic crustaceans, and bivalves. It is eaten by Pacific Halibut, Pacific Cod, Skates, Bearded and Ringed Seals, Stellar Sea Lions, and Beluga. This species has been found in low densities in areas where it has been surveyed. Estimated abundance in the Chukchi Sea is five individuals per square kilometer. Only three individuals were captured during annual summer sampling from 2004 to 2009 at Cooper Island in the Alaskan waters of the western Beaufort Sea. Longhead Dab made up a very small proportion of total flatfish biomass in the eastern Bering Sea. In the eastern Bering Sea, Longhead Dab decreased in abundance from 1979 to 2002 as average sea temperatures increased.

importance: It is not economically important and is mostly

caught as by-catch. In the eastern Pacific it is part of an “other flatfish species” complex (along with Flathead, Rock, and Yellowfin Soles and Alaska Plaice) that is managed as a fisheries unit.

distribution: The species has been found in the western Arc-

tic where there are only three records: Parry Bay near Bathurst Inlet at 68º20' N, 107º41' W (CMNFI 1976-0119); the south coast of Banks Island (NRM 45561); and off the Wollaston Peninsula of Victoria

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Limanda proboscidea

Island (NRM 45562). It is an accidental Pacific stray to the Arctic. It has also been reported from the Alaskan Beaufort Sea based on a single museum record – off Pingok Island at 70º34' N, 149º35' W (CMNFI 1982-1026) – and rare survey records. This species is likely at the eastern extent of its range in Parry Bay, Nunavut. It is also found in the Arctic and North Pacific boreal region, from the Beaufort and Chukchi Seas, the Sea of Okhotsk, and south to Unimak Island in the Bering Sea.

sources: Brodeur & Livingston (1988); Barber, Smith, Vallarino,

& Meyer (1997); Lang, Livingston, & Dodd (2005); Hoff (2006); S.W. Johnson, Thedinga, Neff, & Hoffman (2010); Quakenbush & Bryan (2010); Quakenbush, Citta, & Crawford (2011a, 2011b); Sinclair, Johnson, Zeppelin, & Gelatt (2013).

Distribution of Limanda proboscidea

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Platichthys stellatus (Pallas, 1788)

Starry Flounder, flet étoilé

Platichthys stellatus

common names: Local names are Nataarnaq and Natangnak (Inuvialuktun) and Nataaznak (Inuktitut). Other common names are Diamond Flounder, Emerywheel, Flounder, Great Flounder, Grindstone, Leatherjacket, Rough Jacket, and Swamp Flounder.

taxonomy: The genus comes from the Greek platys (flat) and

ichthys (fish), and the species name is the Latin stellatus (starry). Catalog of Fishes gives the date of authorship as 1787, but the American Fisheries Society list gives 1788, corrected from an earlier edition that had 1787. This species has been placed in the genera Platessa and Pleuronectes. Platichthys rugosus Girard, 1854, described from San Francisco, California, is a synonym. In North American waters the Starry Flounder hybridizes with the English Sole (Parophrys vetulus Girard, 1854). This right-eyed hybrid is known as the Hybrid Sole or Forkline Sole and was given the name Parophrys ischyrus Jordan and Gilbert, 1880. In the northwest Pacific, P. stellatus hybridizes with Kareius bicoloratus (Basilewsky, 1855). Two subspecies of the Starry Flounder, P. stellatus stellatus and P. stellatus rugosus Girard, 1854, have been distinguished by degree of dextrality and numbers of pyloric caeca. The former is the sinistral, Eurasian form, and the latter the North American form.

description: This species is distinguished by its large, rough,

star-shaped tubercles and distinctive dark bands separated by 4–7 white to orange bands on the dorsal and anal fins. The body is deep, and the caudal peduncle is narrow. The caudal fin is slightly rounded. The snout is pointed with a small terminal, oblique, asymmetrical mouth in which the majority of the teeth are on the blind side. The maxilla reaches the anterior portion of the eye, which is noticeably anterior to the migrated eye. The stellate tubercles are scattered over the whole eyed side and in bands along the eyed and blind side dorsal and anal fin bases, with cycloid scales scattered



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between tubercles mostly on the posterior part of the eyed and blind sides. The lateral line, which is slightly curved over the pectoral fin, has 58–83 scales. The dorsal fin-rays number 51–68; the anal fin-rays 35–51, preceded by a strong spine that can sometimes be buried in the skin; the pectoral fin-rays 9–12; the pelvic fin-rays 6; and the vertebrae 34–38. The pectoral fin is bluntly pointed. In the southeast extent of the species’ distribution there are decreased numbers of dorsal and anal fin-rays. There is little or no sexual dimorphism. Unlike all other Arctic flatfishes, Starry Flounders are found in both dextral and sinistral forms. In Japan almost all Starry Flounders are sinistral. In the western Pacific off Kamchatka to Japan and the Sea of Okhotsk, 100% are sinistral. From California to the eastern Gulf of Alaska, 49%–66% are sinistral; and in the western Gulf of Alaska and the eastern Bering Sea, 68% are sinistral. In the eastern Pacific waters of North America, the proportion of sinistral individuals increases from south to north. In the Canadian Arctic, 99% of individuals (80 out of 81) are sinistral. The Starry Flounder is dark brown to almost black with indistinct blotches on the eyed side, and white on the blind side, which is white to cream and sometimes blotched. Like almost all flatfishes, Starry Flounders are pigmented on the eyed side, but individuals that are ambi-colorate (pigmented equally on both sides) or partially pigmented underneath have been found in Arctic Canada. The species reaches 91.0 cm in length and 9.1 kg in weight.

habitat: This is a sublittoral, eurythermal, and euryhaline species,

found on soft bottoms from the intertidal area to as deep as 375 m, but usually shallower than 100 m. It is commonly found in shallow brackish waters such as estuaries and is often found upriver to the limit of tidal movement and as far as 120 km from the mouth, as well as in marshes and coastal lakes. It has a high tolerance for low salinities. Adults are found in a wide variety of habitats including cobble and boulder bottoms with algal and sea-grass thickets; sand bottoms; and sand, boulder, and pebble beds, although the preferred habitat is sand beds. The Starry Flounder is the most abundant flatfish in nearshore areas from California to the Beaufort Sea. Along the Tuktoyaktuk Peninsula this is the second most abundant flatfish and the third most abundant species of marine fish.

biology: The Starry Flounder has a very broad range of dietary

items. Its food includes crustaceans, such as amphipods, mysids, and crabs, as well as clams, snails, polychaete worms, nemerteans, chironomids, priapulids, brittle stars, and small fishes. Larvae (5–12 mm) feed on copepods and nauplii, cladocerans, and barnacle larvae. In the Eskimo Lakes, Liverpool Bay area, the adult diet consists of 49% amphipods, 22% gastropods, and 4% tunicates. Isopods and mysids are the dominant food items in Tuktoyaktuk Harbour; other food items include amphipods, plant remains, polychaetes, fish remains, and nematodes. Juvenile Starry Flounders in Washington consume polychaete worms, mysids, amphipods, and clams, while adults feed mainly on nemertean and polychaete worms, small clams and crabs, and brittle stars. Feeding behaviour differs between dextral and sinistral forms, with dextral fish striking dorsally and sinistral fish striking ventrally.

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Platichthys stellatus, left-eyed

Adults do not undergo extensive migrations but make seasonal inshore or offshore movements. In the summer they are inshore and can be found in shallow water and in estuaries. In the winter they are in deeper (to 300 m) water in offshore locations and deep bays. Movement also occurs into shallower water at night and with tides to feed. Females grow faster, become slightly larger, and live longer than males. The maximum age is 42 years; this was a female from the Beaufort Sea. Sexual maturity is reached in California by males at age 2 and 30 cm and by females at age 3 and 35 cm. Feeding stops during spawning, which occurs in February to April in British Columbia and from June to July in Arctic waters at 1.5°C–6.8°C. Spawning Starry Flounders use shallow water near river mouths and sloughs in Russian waters. They spawn in June to mid-July in bays and inlets off the Tuktoyaktuk Peninsula. Subsequently there is an offshore migration in September. The eggs are pale orange with fine markings, pelagic, and ca. 1.0 mm in diameter, numbering up to 11 million. Larvae are 1.9–2.1 mm at hatching. Metamorphosis occurs at 10.5 mm when the young become asymmetrical. Young Starry Flounder first settle out in shallow water on silty and silty-sandy grounds, most commonly in estuaries, but also in the intertidal zone and in fresh water.

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No studies on the parasites of Starry Flounder have been done in the Canadian Arctic. In Alaska three species of copepods have been recorded. In Monterey Bay, California, an isopod, a copepod, a cestode, and an acanthocephalan were found, and in western Kamchatka more than 20 species of helminthes were found in juvenile Starry Flounders.

importance: The Starry Flounder is a minor (often incidental)

component of commercial fisheries and supports a small sport fishery. It comprises substantial by-catch in western Kamchatka and southeastern Sakhalin. Test fishing has been carried out near Kugluktuk, Coronation Gulf. The commercial landings are low, with 227 t captured annually in British Columbia waters. The species has been described as an excellent food fish and is a popular sport fish for anglers from wharves and along coastal areas.

distribution: It is found in Melville Sound, Bathurst Inlet, Coronation Gulf, Amundsen Gulf, and the Beaufort Sea. It is one of the most wide-spread northern flatfishes, found from the Yellow Sea, the Sea of Japan, the Sea of Okhotsk, the Bering Sea, the Pacific coast of Japan, off the Pacific coast of North America including British Columbia to southern California, and around Alaska.

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Platichthys stellatus, right-eyed

sources: Gudger (1941); Orcutt (1950); Walters (1953a); Andriashev (1954); Hunter (1975); Bond (1982); Lacho (1991); Voronina (1999); Poly & Mah (2001); Brubacher Development Strategies Inc. (2004b); Kolpakov (2005); Bergstrom & Palmer (2007); Moles (2007); Page et al. (2013); Sokolov (2010).

Distribution of Platichthys stellatus



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Pleuronectes glacialis Pallas, 1776

Arctic Flounder, plie arctique

48–64, the anal fin-rays 33–46, and the pectoral fin-rays 8–12. The gill rakers number 9–15 on the lower arch. The lateral-line scales number 73–100, and the vertebrae 37–41. In males the lateral line is complete, but in females over 20.0 cm the posterior portion becomes an open groove. A hermaphroditic individual was found at Beaufort Lagoon, Alaska, in 1984. The eyed side is dark brown, blackish, or olive, tinged with green in some specimens and very dark, verging on black, in others. Occasionally there are scattered dark spots forming dark blotches or wide bands. The blind side ranges from off white to lime green. The fins are pale brown usually with dark spots. Most are smaller than 25.0 cm in length, but exceptionally reaching 44.0 cm, with 35.0 cm being a more normal large size.

habitat: It is a highly characteristic shallow-water and cold-water

Pleuronectes glacialis

common names: Local names are Nataagnaq, Nataarnaq, Nataaznak, Natarinaq, and Puyyaqiaq (Inuktitut).

taxonomy: The genus comes from the Greek pleura (side) and nektes (swimmer). The species name is the Latin glacialis (icy), referring to the habitat of this species. Pleuronectes franklinii Günther, 1862, described from Arctic seas of America, and Platessa dwinensis Lilljeborg, 1851, described from the mouth of the River Dvina (Dwina), Arkhangelski Oblast, Russia, are synonyms. The species was formerly in the genus Liopsetta. P. glacialis and P. putnami are very similar morphologically. Early authors used morphological characters such as pectoral-fin length to distinguish Arctic and Smooth Flounders. These keys did not discriminate the two species on a total of 100 Canadian Museum of Nature specimens. Preliminary results using mitochondrial DNA have proved promising (see also the “Keys” section). description: This species is distinguished by the small mouth,

extending almost to below the anterior part of the eye on the eyed side; the teeth are incisor-like, and their extent is greater on the blind side of the jaws; the head is not strongly concave above the eye; the lateral line is almost straight over the pectoral fin; the dorsal finrays number 64 or less; and the anal fin-rays number 46 or less. It is also distinguished by distribution. The head is short, being about a quarter of the body length. The snout is short. The lower eye is somewhat in advance of the upper eye. There is a prominent postocular ridge followed posteriorly by two protuberances. The lateral line has a slight curve above the pectoral fin and a short supraorbital branch. The caudal fin is large and slightly rounded. The scales are ctenoid on both sides in males, with a patch of cycloid scales on the blind-side abdominal area. Females have cycloid scales on both sides, with ctenoid scales along the base of the dorsal and anal fins. An anal spine is present but not prominent, being small and hidden in the skin. The dorsal fin-rays number

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species of flounder found in brackish nearshore areas of bays and estuaries to about 19 m; rarely, if ever, is it found in deeper water, perhaps to 91 m. It is typically found in shallow coastal waters during summer when it often enters low-salinity habitats including the lower, fresh parts of rivers. It avoids high-salinity areas. It overwinters in offshore marine areas, moves into nearshore habits after break-up, and moves to deeper water in the summer. These movements are most obvious near river mouths and may relate to feeding. The typical habitat for this species is shallow, mud-bottomed coastal waters and river mouths. It is known to overwinter in lakes and channels of the outer Mackenzie Delta. It is wide spread and abundant in coastal waters of the Beaufort Sea and is one of the major fish species found in marginal waters along the Arctic National Wildlife Refuge. This species and the Fourhorn Sculpin are the most important demersal species in the Beaufort Sea. During the spring and summer in the Barents Sea, Arctic Flounder are found at temperatures of 1°C–12°C. Juveniles are found up to 10 m deep, and adults to 25 m. The fish are not very mobile, not making lengthy migrations, and not going more than 40 km from shore. Lagoons, bays, and estuaries of the Beaufort Sea are highly suitable as habitat for this species. Wind-driven upwelling and the capture of plankton in the eastern Canadian Beaufort shelf is much higher than in the western portion, resulting in highly productive benthic habitats for species such as the Arctic Flounder. Due to the affinity of this species for estuarine areas, its distribution is distinctly patchy. In Mason Bay, Mackenzie Delta, Arctic Flounder are one of the most abundant species in the outer lagoons. In Phillips Bay, Beaufort Sea, Arctic Flounder are the most numerous marine species in trapnet sampling. This species is third in overall abundance after Arctic Cisco and Saffron Cod in Liverpool Bay, making up 24% of the total catch, although the catch per unit effort (CPUE) was generally low, with high yields occasionally obtained. Arctic Flounder becomes less common further to the east, in Coronation Gulf, where 15 sets of a bottom gill-net yielded 236 Platichthys stellatus and 17 P. glacialis.

biology: The diet includes bivalve molluscs, crustaceans such

as isopods and amphipods, tunicates, and polychaetes, and it is reported to bury itself in mud to ambush small fishes. In the Beaufort Sea approximately half the diet is composed of polychaetes, and most of the other half is tunicates, pelecypods, isopods, and amphipods

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Pleuronectes glacialis

in that order. Juveniles (46–58 mm) eat chironomids, mysids, copepods, and oligochaetes. In Phillips Bay, Beaufort Sea, the isopod Mesidotea entomon is eaten in great numbers. Polychaetes are the dominant food item in Tuktoyaktuk Harbour, with pelecypods second, oligochaetes third, and then isopods and amphipods. Other food items included nematodes, plant remains, and anthozoans. It is eaten by Bearded Seals. The maximum lifespan generally is about 26 years. Females grow faster than males. Sexual maturity is reached by about the fourth or fifth year, and as late as eight years for females in the Beaufort Sea; the maximum age was 12 years at 29.0 cm in a small sample. The maximum age in the Barents Sea was 22 years, but 82% of the sample was 4–10 years; males reached sexual maturity at 3–4 years, and females at 4–8 years. They range from 10.4 cm to 28.2 cm in Oruktalik Lagoon of the Arctic National Wildlife Refuge, Alaska. In the outer Mackenzie Delta the majority are 20–30 cm. At older ages, females are numerically dominant and are 1.5 times the weight of males of the same age. The strength of year classes was positively correlated with air temperatures, as water temperatures in the shallow feeding grounds of juveniles respond more to air than offshore water temperatures. Spawning occurs under ice from January to March, sometimes as late as June at depths of 5–10 m where there are strong tidal currents



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and temperatures below −1°C. Females produce up to 230,000 eggs of 1.6 mm diameter. The Arctic Flounder is restricted to warm low-salinity areas in the larval and young stages.

importance: It is of limited economic importance, being edible

but with little flesh. In Alaska it was used in coastal subsistence fishing. If exploited, it is used for dog food and was, at least historically, a regular catch in the Inuit fishery in Kugluktuk (Coppermine), Coronation Gulf.

distribution: It is found in Queen Maud Gulf, Bathurst Inlet,

Coronation Gulf, Amundsen Gulf, and the Beaufort Sea. The Arctic Flounder is roughly circumpolar in its distribution, extending from the White Sea, east to and including the Bering Sea and the Sea of Okhotsk, the Aleutian Islands, and the Alaskan portion of the Beaufort Sea.

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Distribution of Pleuronectes glacialis

sources: Ellis (1962); Percy (1975); Morrow (1980b); Hunter

(1981); Bond (1982); Craig (1984); Martell, Dickinson, & Casselman (1984); Goldberg, Yasutake, & West (1986); Bond & Erickson (1989, 1993); Crawford (1989); Kobelev (1989); Atkinson & Percy (1991); Lacho (1991); Wiswar, West, & Winkleman (1995); Voronina & Evseenko (2001); Hiscock & Grant (2004); Quakenbush et al. (2011b); Conlan et al. (2013).

Pleuronectes putnami (Gill, 1864)

Smooth Flounder, plie lisse

common names: Other common names include Blackback,

Christmas Flounder, Eelback, Foolfish, Plaice, Smooth Plaice, and Smoothback.

taxonomy: The species is named after Frederic Ward Putnam (1839–1915), curator of ichthyology, Boston Society of Natural History, and assistant in ichthyology, Museum of Comparative Zoology of Harvard University under Louis Agassiz. It has also been placed in the genera Liopsetta and Euchalarodus, the latter its synonym. description: This species is characterized by a small mouth,

extending almost to below the anterior part of the eye on the eyed side; the teeth are not pointed, and their extent is greater on the

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blind side of the jaws; the head between the eyes is not scaled, and the head is not strongly concave above the eye; the lateral line is almost straight over the pectoral fin; the dorsal fin-rays number 59 or less; the anal fin-rays number 41 or less; and it is found in Frobisher Bay. The mouth is small, with the eyed-side jaws having one set of teeth, the blind-side two sets. The head is moderate, and the snout is short. The interorbital ridge has a prominent broad, tuberculate ridge that is without scales and is smooth. The fixed eye is only slightly in advance of the migrated eye. The eyes are close to the mouth. The anterior nostril tube is prominent and long. The lateral line is nearly straight with a very slight curve over the pectoral fin. A blind-side lateral line is present, and there are accessory branches present over the eyes. The dorsal fin-rays number 48–59, the anal fin-rays 35–41, the pectoral finrays 10–11 on the eyed side; and the vertebrae 34–40. The dorsal and anal fins are highest posterior to the midpoint of the body, and the body is ovate and relatively thick. This species is sexually dimorphic. Males have much longer (about four-fifths as long as the head) and more pointed pectoral fins than do females. The scales are ctenoid on both sides in males, with a patch of cycloid scales on the blind-side abdominal area. Females have cycloid scales on both sides, with ctenoid scales along the base of the dorsal and anal fins. The eyed side can range from uniformly grey brown to blackish or an indistinct marbled green grey, at times with dusky, indistinct, brown blotches in arcs along the body. The blind side is yellowish beige. The dorsal and anal fins have prominent rectangular dark spots, which are also present on the caudal fin. Females reach a maximum of 33.0 cm in length, and males reach 27.0 cm, but average fish do not exceed 20.0 cm.

habitat: The Smooth Flounder is found in estuaries (often year-

round in the Gulf of St Lawrence and in New England) and shallow, sheltered bays on mud and silt bottoms down to a maximum of 27 m, but more usually 10 m, where temperatures and salinities are variable. They can tolerate sea water at its freezing point and highs to 32°C. It is an unusual pleuronectid for the western Atlantic as it is the smallest species and restricted to inshore habitats. Its movement is very limited. The Miramichi estuary population of Smooth Flounder does not migrate outside of that area.

biology: The diet of the Smooth Flounder is wide ranging and

includes small prey such as marine worms, molluscs, amphipods, and shrimps. Small bivalve molluscs are the main prey in the Miramichi estuary, New Brunswick, while sand shrimp and crabs make up the remainder of the diet. Polychaetes and copepods are the primary food items in the St Lawrence estuary. Juveniles in the Bay of Fundy eat primarily harpacticoid copepods. The Smooth Flounder does not feed during the winter. In Nova Scotia, Blue Heron eat Smooth Flounders that are smaller than 10 cm almost exclusively. Maturity is at age 1+ years. Spawning occurs from December to January in Canadian waters and as late as March in more southern localities such as Maine. In the Miramichi estuary, spawning occurs when temperatures are near or below 0°C. Fecundity for females in New Hampshire is 4,600–52,000 eggs. They are 1.1–1.4 mm in diameter and do not float, lacking oil globules. In New Hampshire

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Pleuronectes putnami

26 species of protozoan and metazoan parasites were recorded from this species.

importance: It is not economically important, because of its small size, but it is excellent to eat. At one time the Smooth Flounder was used as feed on fox farms in Atlantic Canada.

distribution: It has been found from a single record (CMNFI 1977-1501H) at 62°59'30" N, 65°55' W, in Peter Force Sound in Frobisher Bay on southern Baffin Island. The previously known northern limit of P. putnami was mid-Labrador. These two specimens may well represent strays, or their apparent rarity is an artifact of collecting methods. Department of Fisheries and Oceans surveys in Baffin Bay, Davis Strait, or Hudson Strait do not usually start until 100 m or deeper (Margaret Treble, DFO, pers. comm.). The Smooth Flounder is found in the western North Atlantic from southern New England (Connecticut), throughout the Maritimes, and north to central Labrador. This is a restricted range compared to that of other pleuronectids. There are no verifiable records from Ungava Bay, despite repeated citations to that effect (Backus, 1957; Laroche, 1981; Collette & Klein-MacPhee, 2002). Distribution of Pleuronectes putnami

sources: Margolis & Arthur (1979); Burn (1980); Quinney &

Smith (1980); Imrie & Daborn (1981); Laroche (1981); Dutil & Fortin (1983); Armstrong & Starr (1994); Hanson & Courtenay (1997).



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Reinhardtius hippoglossoides (Walbaum, 1792)

Greenland Halibut, flétan du Groenland

common names: Local names include Nat-ah-nuh and Natarnak

(Inuktitut); and Kaleralik, Netarnârak, and Qaleralik (Greenlandic). Other common names are Greenland Turbot, Newfoundland Turbot, Turbot, Bastard Halibut, Black Halibut, Blue Halibut, Lesser Halibut, Little Halibut, Mock Halibut, and flétan noir. Halibut exported from Canada to U.S. markets are labelled “Greenland Turbot” due to a 1968 protest by Pacific Halibut processors. Greenland Halibut is the name generally accepted by the Northwest Atlantic Fisheries Organization, the American Fisheries Society, and Fisheries and Oceans Canada.

taxonomy: The genus is named after the zoologist Johannes Christofer Hagemann Reinhardt (1776–1845), first professor of zoology at the University of Copenhagen, who described new species from Greenland. The species name comes from the Greek ippos (horse) and glossa (tongue) and the Latin -oides (like, resembling). Hippoglossus groenlandicus Günther, 1862, and Pleuronectes pinguis Fabricius, 1824, both described from Greenland, are synonyms. It was placed in the genus Platysomatichthys Bleeker, 1862, in older literature. Atlantic and Pacific populations may constitute distinct subspecies. Reinhardtius matsuurae Jordan and Snyder, 1901, described from Japan, was found not to differ from R. hippoglossoides on the basis of morphometrics and meristics, but it is still considered a valid subspecies in the Pacific Ocean by some Russian workers. The subspecific status of Atlantic and Pacific populations is supported by a study of protein loci that suggested genetic divergence at that level. description: This species is characterized by a large mouth,

extending to or past the mid-eye on the eyed side; the teeth are pointed, about equal in extent in both jaw halves; the uppermost eye lies in a notch on the upper head margin; and the blind side of the body is dark in adults. The body is elongate and laterally compressed, with well-developed musculature on the blind side, reflecting the active movement and feeding habits of this species. The eyes are more separated than those of any other pleuronectid species. The preopercle has a right angle at its posterior margin and 4 or 5 pores near the edge. The dorsal fin origin is posterior to the upper eye. The caudal fin is concave or slightly forked to truncate. The dorsal fin-rays number 83–109, the anal fin-rays 62–84 with no pre-anal spine present, the pectoral finrays 11–15 on the eyed side, and the pelvic fin-rays 5–7. The gill rakers number 10–16 on the lower arch and are massive and short. The lateral-line scales number about 109–119, the lateral line being straight and slightly decurved above the pectoral fin. The scales are cycloid on both sides. The vertebrae number 59–64. The eyed side ranges from rust-coloured or purplish brown to dark or greyish brown to black

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with a green sheen. The blind side is initially creamy white but then becomes grey to dark grey or blue greenish, or even quite dark in adults. In the southern extent of the Atlantic population the eyed side is yellowish or greyish brown. As with other flatfishes, it is occasionally found reversed (left-eyed). The species attains 1.3 m in total length and 44.5 kg in weight.

habitat: This is a deep-water species found off soft bottoms at

depths of 14–2,000 m, usually at 50–650 m, with the larger specimens at the greater depths, at temperatures from −0.5°C to 7.0°C. Greenland Halibut are often numerous in areas of −1°C to 3°C water. Temperatures below −1°C can be lethal. They may be found in shallow waters in the north but are in deeper waters in the south. In its northern home range this species, especially as breeding adults, is found in deep water, beyond the continental shelf. Greenland Halibut in the northern Atlantic develop at uniform, near-bottom temperatures on the continental slope at depths of 600–1,000 m. They are recorded at 146–1,487 m and −0.08°C to 5.4°C in Davis Strait and southern Baffin Bay for 65 collections; and to 435 m in the Canadian Beaufort Sea, at 1,000–2,000 m in Mittimatalik (Pond Inlet), to 1,067 m in Cumberland Sound on Baffin Island, at 290– 1,483.5 m and from −0.1°C to 5.4°C in southern Baffin Bay and Davis Strait generally for up to 1,099 fish per tow, and at 0–530 m in Hudson Strait. This fish makes extensive vertical movements and spends less time on the bottom than do other flatfishes. Greenland Halibut tagged in White Bay, Newfoundland, have been recaptured off Baffin Island. Greenland Halibut are most abundant in areas associated with the shrimp Pandalus borealis, and with Sebastes mentella and Glyptocephalus cynoglossus at depths of 150–450 m, with temperatures between 4.5°C and 6°C in the Gulf of St Lawrence. In Davis Strait, species associations were with Boreogadus saida, Arctogadus glacialis, and Amblyraja hyperborea, Liparidae, and Lycodes spp. Greenland Halibut migrate extensively between feeding grounds and spawning areas. Spawning fish migrate northward to Davis Strait and remain in deep water. There is no evidence of migration between west Greenland fiords and Davis Strait, confirming the isolation of those populations. Greenland Halibut in northwestern Greenland and Cumberland Sound remain resident in those areas. North of the Baffin-Greenland Rise the recruitment may come from Baffin Bay.

biology: As the Greenland Halibut is a deep-water species with

a specific temperature preference and a main large spawning area, there is no marked variation in the Atlantic populations. Halibut in Davis Strait, northern Labrador, and eastern Newfoundland could be distinguished from Gulf of St Lawrence and Fortune Bay populations in southwest Newfoundland based on blood parasites. Multivariate meristic analysis has shown that Greenland Halibut form two distinct populations: Gulf of St Lawrence (which receives migrants from the Labrador area through the Strait of Belle Isle between Québec and the island of Newfoundland) and Davis Strait. Halibut from west Labrador are probably spawned in Denmark Strait. Another multivariate analysis has distinguished eight stocks

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Reinhardtius hippoglossoides

in the northwest Atlantic when it had been demonstrated that there was one. This was due to north-south clinal variation in morphometric characters. A study based on meristic and allozyme variation suggested a single interbreeding stock in the northwest Atlantic with partial isolation of populations in west Greenland fiords. All North Atlantic Greenland Halibut were shown to be genetically homogenous based on mitochondrial DNA. However, analysis of nuclear DNA showed two groups that did not correspond to geographic populations but were both represented in the Gulf of St Lawrence, as well as within Davis Strait, suggesting an east-west differentiation of Greenland Halibut in that area. Greenland Halibut deviate from other pleuronectids in several ways: almost identical coloration of both body sides; positioning of the eyes resulting in bilateral vision; an elongate shape; and an unusual habit of often feeding in the water column. The partially migrated eye on the midpoint of the head provides a larger field of vision. This and the dark coloration on the blind side with notable musculature of the blind side indicate a secondary adaption to pelagic feeding. They are vigorous swimmers, exhibiting a bathypelagic behaviour. This behaviour has also been directly observed and inferred by the evidence of catches of Greenland Halibut in Atlantic Salmon nets. Greenland Halibut make regular excursions for several hundred meters upward through the water column. Smaller Greenland Halibut can be found further off the bottom, and pelagic activity is greatest during early autumn. Greenland Halibut have been observed in Greenland waters of 300 m depth to rise to 200 m, to be caught in handline fisheries. Halibut caught in this fishery were full of Capelin, indicating pelagic feeding.



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This species is a voracious and pelagic-to-bathypelagic feeder. Species eaten include Atlantic and Polar Cod, Capelin, Roundnose Grenadier, Witch Flounder, barracudinas, redfishes, American Sand Lance, and their own young. A variety of crustaceans, squids, and bottom invertebrates are also eaten. Hyperiidae, Boreogadus saida, and the shrimp Pandalus borealis are important within the diets of fish smaller than 60 cm. Cephalopods and Liparidae are commonly eaten by a wide range in size of Greenland Halibut. An abrupt change in diet at about 64 cm to 69 cm is related to changes in both feeding habit and geographic distribution. Sebastes mentella and their own species are important dietary components of Greenland Halibut that are larger than 75 cm. Stable isotope analyses show that, in Cumberland Sound, Greenland Halibut feed heavily on Capelin. Off southern Labrador and northeastern Newfoundland, Greenland Halibut under 20 cm eat small crustaceans and squid; fish from 20 to 69 cm eat mainly Capelin; and halibut larger than 70 cm feed on demersal fishes, predominantly redfishes, macrourids, and other Greenland Halibut. Food habits and intensity of feeding are related to several factors: ontogenetic changes, location, depth, and season. Depth and temperature are of less importance than Greenland Halibut size in prey determination. In Davis Strait the dominant prey species of large Greenland Halibut are beaked redfishes and Roundnose Grenadier, whereas in Labrador myctophids, Roundnose Grenadier, and Capelin make up the principal diet; in eastern Labrador it was Capelin, Sand Lances, and Pandalus borealis. A dominant prey item for small Greenland Halibut in the southern Labrador-Newfoundland area is hyperiid amphipods. Mid-sized fish consume Pandalus and cephalopods

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(mainly Gonatus). Larger Greenland Halibut eat larger fishes, including redfishes, gadiforms, and other Greenland Halibut. Diet choice can be flexible: when redfish disappeared from this area after 1992, Greenland Halibut switched to gadiforms. There is strong diel behaviour associated with feeding. Predators include Greenland Shark (considered the most significant predator), Atlantic Cod, Atlantic Salmon, Beluga, and Hooded and Harp Seals because the young are abundant in the shallow benthos. Greenland Halibut were the second most numerous food item in Atlantic Cod stomachs in 1983. Narwhals are known to feed intensively on Greenland Halibut, and in the Pond Inlet area Greenland Halibut can constitute on average 37%, but as much as 45%, of the total weight of stomach items. One study suggested that a total of 790 t of Greenland Halibut are eaten daily by Narwhals in two overwintering areas in Baffin Bay and northern Davis Strait. Seventy-six parasite species are reported from Greenland Halibut, with two (a myxosporidean and a copepod) being unique to that species. Growth rate varies with the area. Greenland Halibut in the Arctic environment are slower growing and longer lived than previously thought, necessitating some caution in conservation and management. They can live to 36 years. On the continental shelf, immature Greenland Halibut of both sexes exhibit the same growth rate. On the continental slope, however, there is a significant difference in males and females, with females exhibiting higher growth rates, living longer, and achieving a larger ultimate size than males. The age at 50% maturity is 9.5–15.0 years for females and 8.2–11.6 years for males in the Canadian northwest Atlantic Ocean. All fish over 90 cm are female. The major spawning location in the western Atlantic is in the deep slope area of Davis Strait along the boundary between Canada and Greenland. Spawning occurs at depths of 600–1,000 m in the relatively warm Atlantic Layer (ca. 4°C and 34.5‰ salinity) south of the Baffin-Greenland Rise that separates Davis Strait from Baffin Bay. Spawning takes place south of 64° N at depths greater than 1,200 m at temperatures of 3.0°C–3.5°C from mid-December to mid-April. Peak spawning occurs in Davis Strait from February to March, although the season can be extended beyond that. Spent females are found in June in Davis Strait deep water. Individuals in Davis Strait can be found in a mature pre-spawning condition year-round as temperatures there are particularly conducive to spawning. Not all adults spawn every year. Each female produces up to 300,000 eggs. The eggs reach 4.5 mm (range 3.7–4.7 mm) in diameter and are clear with a small perivitelline space. The development time for the fertilized eggs is about 50 days. The larvae can drift for up to four months before settling. The eggs and larvae are dispersed by currents both to the west coast of Greenland by the West Greenland Current and to the eastern Canadian coast by the Canadian Polar Current. Young Greenland Halibut in west Greenland migrate both to Davis Strait and to the deeper parts of the fiords. On reaching maturity, Greenland Halibut in the fiords migrate to the main spawning area in Davis Strait. The larval period is very long (with larval sizes to 54–57 mm). The larvae are pelagic until 80 mm. At about 70 mm they begin settling on the bottom, and metamorphosis

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begins. Larvae at 60–70 mm commence metamorphosis at depths of 200–300 m on clay-silt substrates; one- to two-year-old fish preferred clay sediment. The Baffin Bank may be a nursery area. Nursery areas at depths of 300–450 m around Disko Bay, Greenland, with soft productive bottoms are favoured. The slopes of the banks west of Disko Bay and the northern and western slopes of Store Hellefiske Bank are a nursery area for Greenland Halibut. Young Greenland Halibut are found in the area between 64°00' N and 64°45' N in high densities during the summer and autumn. This area is known for high primary and secondary production.

importance: Greenland Halibut is one of the top predators in

the northwest Atlantic and the most abundant top predator since the collapse of the northern Atlantic Cod. Greenland Halibut are caught in gill-nets, trawls, and long lines and sold as fresh-frozen fillets. They are also smoked and salted for local use. The three main fisheries are Baffin Island–West Greenland, Labrador–East Newfoundland, and Gulf of St Lawrence. Greenland Halibut were exported from Newfoundland beginning in 1857, reaching 590 t, and remaining at 250–900 t until 1964. Limitations to commercial harvest in the Arctic are the lack of suitable equipment and infrastructure, as well as transportation issues. Distribution of Greenland Halibut in deeper waters means that specialized gear is necessary to catch them. Landings from 1999 to 2003 were 16,234 t, 21,706 t, 22,083 t, 16,454 t, and 19,617 t. They constitute a by-catch in the shrimp fishery in Davis Strait. In 1965 there was an experimental halibut fishery in Killiniq, which was not continued due to the logistical difficulties of vessels’ offshore capabilities. Off Baffin Island and parts of western Greenland, the catches peaked at 18,000 t in 1992 but were stable around 10,000 t during 1993–2000. Catches increased to 24,155 t due to increased effort in NAFO division areas 0A and 1A in 2007. Catches stayed stable during 2003–2005. The Cumberland Sound Greenland Halibut fishery began in 1986, has been fished in the winter months using long lines under the ice, and is accessed by snowmobile. The total allowable catch (TAC) peaked at 430 t in 1992, declined to below 100 t in the late 1990s, and peaked again at 245 t in 2003; it was set at 500 t in 2005. From 2005 to 2008, catches were very low with harvests of 9 t, 70 t, 3 t, and 32 t, respectively. Years of low catches are related to poor ice conditions that led to low fishing effort. Long lines are preferred in this fishery as gill-nets tend to catch primarily large mature females. There is also the issue of by-catch of Greenland Shark and entanglement of marine mammals such as Narwhals, Bowhead Whales, and Belugas. Lost gillnets can turn into ghost fishing nets, catching fishes and other species for years.

distribution: This species is found in Nares Strait, Baffin Bay, Davis Strait, Cumberland Sound, Frobisher Bay, Hudson Strait, Ungava Bay, northern Hudson Bay, Fury and Hecla Strait, Lancaster and Jones Sounds, off northwest Ellesmere Island at 83° N, at western Banks Island, and in the Alaskan Beaufort Sea (CMNFI 1974-0282A). It is found on all coasts of Greenland. One of 11 specimens from the Beaufort Sea was a ripe female, suggesting that spawning is taking place. This is quite possible as spawning fish have

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been found in areas distant from main spawning areas. Deep-water zones of 4,000–5,000 m exist off the Beaufort shelf, providing habitat for adults and a potential area for spawning. This species is amphiboreal, found mainly in the North Atlantic and North Pacific Oceans, with an increasing number of recent records in the Arctic Ocean off Alaska and Canada. It is found in the Pacific Ocean from northern Japan to the Chukchi Sea and south to British Columbia, with stragglers as far south as Baja California. In the northeastern Atlantic it is found from northern Norway to southwest of Ireland, westward through Iceland, and around Greenland. In the northwest Atlantic it is found from Arctic Canada southward throughout maritime Canada including the Gulf of St Lawrence as far east as the Saguenay Fjord, and south to the Gulf of Maine. It is most abundant from the eastern Grand Banks and Flemish Cap and northwards. Sparse southern populations are made up of non-breeding expatriates from the far north.

sources: Jensen (1909, 1935); Hubbs & Wilimovsky (1964); Templeman (1965, 1973); Drainville (1970); Groot (1970); Smidt (1971); Chumakov (1975); Fairbairn (1981); Tremblay & Ayelsen (1981); Finley & Gibb (1982b); Khan, Dawe, Bowering, & Misra (1982); Misra & Bowering (1984); Bowering & Parsons (1986); Chumakov & Podrazhanskaya (1986); Gillis & Allard (1986, 1987a); Alton, Bakkala, Walters, & Munro (1988); Bowering (1988); Riget & Boje (1988); Crawford (1989, 1992); Hudon (1990); Bowering & Lilly (1992); Riget, Boje, & Simonsen (1992); Junquera & Zamarro (1994); Bowering & Brodie (1995); Chiperzak, Saurette, & Raddi (1995); Rodríguez-Marín, Punzón, & Paz (1995); Mathias & Keast (1996); Jørgensen (1997a, 1997b, 2005a, 2007, 2010); Orr & Bowering (1997); Vis, Carr, Bowering, & Davidson (1997); Stene, Gundersen, Albert, Nedreaas, & Solemdal (1998); Kulka (1999); Rideout, Maddock, & Burton (1999); Boje (2002); Boje & Simonsen (2004); Laidre, Heide-Jørgensen, Jørgensen, & Treble (2004); Osborne et al. (2005); Simonsen & Gundersen (2005); González et al. (2006); Gregg, Anderl, & Kimura (2006); Cooper, Maslenikov, & Gunderson (2007); Dyck, Warkentin, & Treble (2007); Department of Fisheries and Oceans (2008, 2013q); Department of Fisheries and Oceans (Treble) (2008a); Healey & Mahé (2008); Pomilla, Treble, Postma, Lindsay, & Reist (2008); Treble, Campana, Wastle, Jones, & Boje (2008); Vollen & Albert (2008); Dennard et al. (2009); Treble & Stewart (2009); Dwyer, Buren, & Koen-Alonso (2010); Gundersen et al. (2010).

Distribution of Reinhardtius hippoglossoides



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GLOSSARY Brian W. Coad

This glossary provides descriptions of technical terms and some commoner words that have a special meaning in ichthyology, along with terms related to the Arctic environment. Structures used in fish identification are illustrated in the “Keys” section. The website www.briancoad.com has an extensive dictionary of ichthyology. 1+. Age, meaning fish between one and two years old (and 2+, 3+, etc.). ‰. Parts per thousand (ppt), used for measuring salinity, for example; see also psu.

A

abdominal. Pertaining to the abdomen. The pelvic fins have an abdominal fin position when they are found on the abdomen and are remote from the pectoral fins. abdominal cavity. The part of the body containing the viscera or guts, liver, ovaries, testes, kidneys, etc. absolute fecundity. The total number of eggs in a female. abyssal (adjective of abyss). Water below 4,000 meters or 2,000 fathoms (3,660 m), down to 6,000 m, where light does not penetrate. It is occasionally used for depths below 2,000 m. It is a constant environment with temperatures that are usually 0°C–2°C or uniform. abyssobenthic. The depth zone of the ocean floor between 4,000 m and 6,000 m, or from about 3,700 m downward, or below the 4°C isotherm. abyssopelagic. Living pelagically in deep waters between 4,000 m and 7,000 m, deeper than the bathypelagic layer. AC. A series of ventro-lateral photophores extending between a vertical at the anal fin origin and the end on the caudal peduncle. The AC row may begin posterior to the anal fin origin if it is offset from other ventro-lateral photophores. accessory lateral line. A branch of the lateral line in certain flatfishes, originating at the head and extending along the upper flank near the base of the dorsal fin. adfluvial. Living in lakes and migrating into streams to spawn; juveniles feed in streams but as subadults migrate to lakes for feeding. adhesive disc. A sucker, found on the belly of Cyclopteridae (Lumpfishes) and Liparidae (Snailfishes) and modified from the pelvic fins. adipose eyelid. Transparent membranes over the anterior and posterior parts of the eye, e.g., in the Clupeidae (Herrings). adipose fin. A small, tab-like fin on the back behind the dorsal fin. It is fleshy and lacks rays and spines. See also ventral adipose fin. aestivate. The ability to remain dormant in the dry season.



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age group 1+. Age, meaning fish between one and two years old (and 0+, 2+, 3+, etc). alar spines, alar thorns. Large spines near the tips of the upper surface of the pectoral fins of male Rajidae (Skates). alevin. A young fish with a yolk sac; larva of species in which post-larval stages are not recognized, that is, in which the yolk-bearing larva transforms directly into the juvenile, e.g. in Salmonidae (Trouts and Salmons); the stage from hatching to the end of dependence on the yolk sac as the primary source of nutrition. allopatric. Referring to populations or taxa whose ranges do not overlap; geographically separated. allozyme. A different structural form of the same enzyme (coded by a different allele at the same locus). ambi-coloration. Pigmentation on both the eyed and the blind sides of flatfishes that normally have only the upper (or eyed side) pigmented. ammocoete. The larval stage of Petromyzontidae (Lampreys), characterized by the absence of teeth, disc, and eyes and by the presence of an oral hood. amphiboreal. Pertaining to an interrupted northern circumpolar distri­bution. amphidromy (adjective, amphidromous). Fishes that regularly migrate between the sea and fresh water (or vice versa) at some definite stage in their life cycle but not for the purpose of reproduction. amphitropical. Pertaining to a distribution of temperate species interrupted by the tropics. ampullae of Lorenzini. Jelly-filled canals on the snout and head of sharks and rays that are involved in electroreception. anadromous (noun, anadromy). Said of fishes that spawn in fresh water but spend part of their life in the sea, e.g., Salmonidae (Trouts and Salmons). The opposite is catadromous. anal fin. The fin behind the vent and anus on the lower, rear part of the body. Some fishes may have two anal fins. anal papilla. A fleshy protuberance before the anal fin through which the end of the intestine passes. It is well developed in some members of the Cottidae (Sculpins). It is also called genital papilla or urogenital papilla. annulus (plural, annuli). A ring or rings on a fish scale or in a bony or cartilaginous structure corresponding to a year of growth. In a scale it usually consists of closely arranged ridges (circuli). An accessory annulus is a ring caused by retarded or temporarily terminated growth that does not represent an annual cycle.

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anterial. One of the teeth on the anterior field of the oral disc of Petromyzontidae (Lampreys). See also endolateral; exolateral; infra-oral; supraoral. anterior. In front of; towards the front (opposite of posterior). anthropogenic peturbation. Changes in the environment caused by human actions. anti-freeze. Chemical components that lower the freezing point of fish body fluids below that of sea water (−1.9°C), enabling survival in cold waters. antitropical. Found in cooler waters north and south of the tropics. anus. The posterior opening of the digestive system through which faeces are voided. AO. The anal photophores, a series of light organs above the anal fin base and along the lower caudal peduncle. arch. See gill arch. archibenthic. The waters on the slope beyond the outer edge of the continental shelf at depths between 200–400 m and 1,000–1,100 m or below the 4°C isotherm. archipelago. A group of islands or an expanse of water with scattered islands. Arctic. A polar region at the northernmost part of the Earth. The Arctic consists of the Arctic Ocean and parts of Canada, United States (Alaska), Russia, Finland, Sweden, Norway, Iceland, and Greenland. Arctic Deep-Water Layer. Water below the Atlantic Layer from 800 m depth to the bottom. Arctic Surface Layer. The uppermost layer of water extending to about 200 m depth. Atlantic Water Layer. Waters of 200-800 m in depth entering the Arctic Ocean via the Barents Sea and the Fram Strait. attached. Not free, e.g., said of a structure that is connected to another, such as dorsal and anal fins being attached to the caudal fin. auditory capsule. Cartilaginous skeleton about the inner ear; also called otic capsule. author. The person to whom a published work or zoological name is attributed. axil. The angle between the upper side of the pectoral or pelvic fins and the body. axillary scale or process. A scale found at the base or axil of the pectoral or pelvic fins. It may serve to streamline the fin.

B

backwater. A still-water section of a stream or river beside the main flow but separated by a ridge of land (or an arm of the sea similarly separated from the open ocean), or habitat at the margin of a riffle or a run. It is sometimes used for water that has backed up compared to its normal flow or for an area off the main part of a lake; often separated from the source during dry seasons. band. A vertical block of pigment (a stripe is horizontal). bar. Elongated pigment distributed vertically (compare stripe). barb. Another term for spinules, e.g., in the Cottidae (Sculpins). barbel. A thin, fleshy appendage near the mouth used for touch or taste senses. bar-coding. See DNA bar-coding. basibranchial. Median bones at the base of the gill arches. batch spawner. A fish that sheds eggs more than once in a spawning season rather than within a short period (a fractional spawner). See also serial spawner. bathyal. The area of the continental slope, a benthic habitat from 200 m to 4,000 m deep. bathybenthic. Pertaining to life on the deep sea floor. bathydemersal. Living and feeding on the bottom below 200 m. bathylittoral. That part of the marine sublittoral zone that is devoid of algae.

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bathypelagic. Referring to mid-waters between 1,000 m and 4,000 m, below the mesopelagic zone. beach seine. A net used to encircle fish in shallow water; usually operated by two people wading out from shore; the net has lead weights to keep the bottom on the sea floor and floats to keep the top of the net at or near the surface; there may be a bag extending back from the centre of the net’s length to increase capture efficiency. The seine may be set from a boat but hauled in from the land. Beaufort Gyre. A wind-driven, clockwise-rotating ocean current in the Beaufort Sea over the Canada Basin, composed of both surface water and ice. benthic. Bottom dwelling. benthivore. A feeder on benthos. benthopelagic. Pertaining to fishes that swim just above the sea bed. benthos. Organisms that live on the bottom of a water body, in it, or near it (and also the bottom of a body of water including the sediment); compare epibenthic. Beringia. An area comprising the land around the Bering Strait. Historically, the ice-free Bering land bridge connected Asia with North America during the Pleistocene. bicuspid. With two points or cusps, as in teeth. bifid. Divided in two. bifurcated. Divided in two, forked. bilaterally symmetrical. With most organs identical on each side of the midline; left and right halves almost identical. bioluminescence. The production of light by living organisms using a chemical reaction. biomass. The weight of living material in a given area or sample or for a given species. biota. All living organisms of a region. birth river, birth stream. See natal river, natal stream. blind side. The side of flatfishes without eyes, lying on the sea bed. Opposite of eyed side. block. A frozen fish fillet in a rectangular shape. bone. A hard supporting tissue made up of cells, fibres, and calcium and phosphate salts; compare cartilage. boreal. Referring to the north temperate region between the true Arctic and tropical regions. bottom trawl. A bag-shaped net that is dragged along the sea bed. BR. Photophores along the lower jaw or on the branchiostegal rays. brackish. Water with a salinity intermediate between fresh and salt water, usually 0.5‰–30.0‰. branchial. Relating to the gills. branchiostegal membrane. The membrane below the operculum that helps to enclose the gill chamber. This membrane is separate when the two sides are not joined to each other or to the isthmus beneath the head. It is united and free from the isthmus when both sides are joined with each other and have a narrow or wide margin posteriorly that is unattached to the isthmus. It is joined when both sides fuse to the isthmus without a free posterior margin. branchiostegal ray. The curved, strut-like skeletal support for the branchiostegal membrane. breast. The area anterior to the abdomen on the ventral surface of the fish. broadcast spawning. The release of eggs and sperm into the water for external fertilization without parental care. buccal gland. The gland in Petromyzontiformes (Lampreys) that secretes a saliva-like fluid having anti-coagulant, haemolytic, and cytolytic properties; the secretion is called lamphredin. bulb. The rounded swelling that forms the main body of the esca or bait at the end of the illicium or fishing rod in anglerfishes.

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by-catch. Species taken incidentally in a fishery. Usually the by-catch is less valuable and is often discarded even though it may contain young of commercial species.

C

caeca. Pyloric caeca are finger-like blind sacs attached to the junction of the stomach and the intestine that serve to aid in digestion. caisson. An airtight chamber used in underwater work, open at the bottom and containing air under sufficient pressure to exclude the water. cancellous. Spongy, porous, or reticulate, usually said of bone. canine teeth. Large, pointed teeth, usually few in number. cannibalism. Eating members of one’s own species. carrion. Animals used by fish as food when the animals are dead and often partially decomposed. cartilage. A flexible supporting tissue made up of cells and fibres but low in calcium and phosphate salts. Also known as gristle; compare bone. caruncle. A fleshy superficial outgrowth or knob. catadromous. Said of fish that spawn in the sea but spend most of their life in fresh water, e.g., possibly Pungitius pungitius (Ninespine Stickleback). catch per unit effort (CPUE). A term for the catch in numbers or weight taken for a given amount of fishing effort over time using specific gear, expressed as a ratio. It is often considered an index of fish biomass or abundance; a decline in CPUE usually indicates a decline in the stock. It may be used as a measure of economic efficiency of fishing. caudal fin. The tail fin. See also diphycercal; heterocercal; homocercal. caudal luminous gland. See infracaudal luminous gland; supracaudal luminous gland. caudal peduncle. The part of the body between the end of the anal fin and the base of the tail fin; the tail stem. centrum (plural, centra). The central body of each vertebra. character displacement. Forced evolution of dissimilar characters in related species where their ranges overlap. Species differ more where they occur together than when their distribution does not overlap. Usually this is detected as morphological features related to resource exploitation. characters. Structures and other features like colour used to distinguish and describe species, families, orders, etc. cheek. The area between the eye and the preoperculum. chest. The anterior part of the ventral surface of a fish just behind the head. chin. The anterior part of the underside of the head, just behind the mouth. circulus (plural, circuli). The concentric ring on the scales, laid down in growth. circumboreal. Around the northern hemisphere in the higher latitudes. circumorbital. Around the eye, often referring to a series of bones that partly encircle the eye in fishes. cirrus (plural, cirri). Fringe-like, fleshy appendages. clasper. A finger-like extension of the pelvic fins of male Rhinochimaeridae and Chimaeridae (Longnose and Shortnose Chimaeras), sharks, and rays, used to deliver sperm to the female, but not for holding as the name suggests. Also called tenaculum and myxopterygia. See also head clasper. class. The group above order and below phylum in classification. clavicular spine. A spine in the shoulder region. cleithral head spine. See head spines. cleithral symphysis. The junction of the ventral and anterior ends of the cleithra. cleithrum (plural, cleithra). The main, dermal, L-shaped bone of the pectoral girdle. cloaca. The vestibule into which empty the urogenital and digestive systems and which opens to the exterior, usually in front of the anal fin. cod trap. A pound net designed to capture cod. It consists of a net floor and walls in a box-like shape with a small opening on one wall called the door. Leader nets running from the shore or a shoal direct the fish into the net.



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coelom. Abdominal cavity or body cavity containing the guts, gonads, kidneys, etc. commercial fishery. All fishing for purposes of sale or barter. It is strictly regulated by species, by gear, by season, and by quotas. common name. The vernacular name of a species, varying from place to place, by language, and over time. Scientific names, in contrast, are in Latin or Latinized Greek worldwide and are subject to rules of usage that cannot apply to common names. Some common names of rare or deepsea species are artificial “book names” because these species are never seen by the general public. They are coined simply to provide a consistent format in books where common names are used or to provide a means of communication with people unfamiliar or uncomfortable with Latin names. Official common names are an attempt to standardize usage, and some countries have recommended lists. complex. A convenient term used to recognize a group of closely related species that have not yet been adequately distinguished by scientists. compressed. Flattened from side to side. The opposite is depressed. congeneric. Belonging to the same genus. contact organ. The dermal bony outgrowth or spicule projecting from a finray or scale margin and surrounded by the epidermis through which bony outgrowths may protrude. It is present in those parts of the body and fins of the male that come in direct contact with the female during the spawning act. It may be tactile in function. continental shelf. The area of gently sloping sea bottom from the shore out to a depth of about 200 m. It may be only a few kilometers offshore where the sea floor descends rapidly to great depths, or it may be extensive and form an accessible habitat for many commercial fishes. continental slope. The steep slope connecting the ocean basins with the shallow continental shelf waters. Also called the continental rise. continuous. Without a break or interruption, as in a continuous lateral line extending completely from head to tail along the flank. Coriolis effect or force. Motion relative to a non-inertial, uniformly rotating frame of reference such as the Earth. coronal head spine. See head spines. counter-current heat exchanger. The mechanism of blood vessels in which heat is exchanged between those vessels going to the skin and gills and those vessels going to the deep body tissues. countershading. The typical pigmentation pattern of fishes, dark above and light below. The effect is to obscure the image of the fish to predators by blending with the dark sea floor when viewed from above, with the light sky when viewed from below, and with the general diffused pattern of light when viewed from the side. CPUE. Catch per unit effort. cranium. The skull. crenate. Scalloped. crenulate. Minutely scalloped. crest. A ridge, usually on the middle of the top of the head, but it can be on a lower surface. cryophilic. Association with temporary or permanent ice. ctenoid scale. A scale with small spines or ctenii on the posterior part, giving the fish a rough feeling; it is found mainly in spiny-rayed fishes. cusp. Point, as on a tooth or a spine. cusplet. A small or secondary cusp; also a denticle. cycloid scale. A smooth-edged scale without spines, found mainly in softrayed fishes.

D

Danish seine. A seine or cone-shaped otter trawl that is hauled from an anchor buoy over an area of about two square kilometers to a stationary vessel, the very long towing ropes disturbing clouds of mud that help herd the fish into the net. deciduous. Easily detached.

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deglaciation. The “retreat” of ice by melting, uncovering land from beneath an ice sheet. demersal. Living on or near the bottom. dental plate. A flattened tooth element. dentary (plural, dentaries). The anterior, deep bone in the lower jaw, often bearing teeth and usually the only bone in the mandible with teeth. denticle. A small scale in the skin; also called placoid scale or dermal denticle. A term sometimes used for a cusplet. denticular teeth. Specialized teeth that may lie outside the mouth on the snout and lower jaw in male anglerfishes, and which are used to attach to the female. depauperate. Impoverished; said of ichthyofaunas or areas with little diversity in numbers or species. depressed. Flattened from top to bottom, e.g., Rajidae (Skates). The opposite is compressed. dermal. Relating to the skin. dermal plate. A bony plate in the skin of the flank, e.g., in Gasterosteidae (Sticklebacks). dextral. Used to denote flatfishes with eyes on the right side; compare sinistral. diagnosable. Having character(s) that enable the fish to be separated from other populations. diel. Daily, a 24-hour period. dimorphic. Having two forms, e.g., two sexes. diphycercal. A tail fin that is both internally and externally symmetrical. directed fishery. A commercial effort aimed at catching a certain species or group of species. It may also apply to a sport fishery. disc. The circular body of Rajidae (Skates). The area surrounding the mouth of Petromyzontidae (Lampreys); see oral disc. The ventral disc of Cyclopteridae (Lumpfishes) and Liparidae (Snailfishes); see adhesive disc. discard. The part of a fish catch that is thrown overboard but which may be of important ecological or commercial value. It is also the act of throwing fish overboard. The discard typically consists of “non-target” species, damaged specimens, or undersized specimens. The fish may be alive or dead, whole or in parts. discharge. Flow of water in a river or drainage basin, measured in cubic feet per second (cfs) or cubic meters per second, passing a certain point. disjunct. Distinctly separate; said of ranges that are discontinuous so that discrete, but potentially interbreeding, populations cannot interbreed. distal. Away from the centre of the body or point of attachment. diurnal. Pertaining to a day; daily. Dn. A photophore above and in front of the eye and above the olfactory capsule. DNA. Deoxyribonucleic acid, the molecule in fish and other organisms that encodes instructions for development and function, used to distinguish and relate species and populations. DNA bar-coding. The use of a short genetic marker in an organism’s DNA to identify it to species. The mitochondrial CO1 gene is used in fish and other animals. domestic fishery. According to the Northwest Territories Fishery Regulations, domestic fishing means “fishing for personal use but not for sale or barter” (see http://lawslois.justice.gc.ca/eng/regulations/C.R.C.,_c._847/ page‐1.html). Furthermore, clause 23 states: “A domestic fishing licence may be issued to a person who (a) has resided in the Northwest Territories for at least two years; (b) needs fish for food for himself or his family; or (c) needs fish for food for his dogs.” Beneficiaries may fish without a licence and may sell, trade, or barter with “(a) other beneficiaries, fish or any portions thereof taken for subsistence usage within the Inuvialuit Settlement Region; and (b) any other person, the non‐edible byproducts of fish taken from the Inuvialuit Settlement Region for personal use.” dorsal. Of or pertaining to the back.

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dorsal fins. The fins on the midline of the back. There may be up to three of these. dorsum. The back or upper surface. dory. A boat with a narrow and flat bottom, flaring sides, and a high bow. drumming muscle. One of the muscles attached to the gas bladder that makes the contained gas vibrate and emit a drumming sound. duct. Any tube-like structure. dwarf. Atypically small.

E

ear stone. See otolith. ecophenotype. A phenotype showing adaptations associated with the habitat or environment that are not genetic. ecoregion. An ecological region, relatively large areas of land or water containing characteristic, geographically distinct assemblages of natural communities and species. ecosystem. The complex of living organisms and environmental conditions that function as a unit. ecotype. A population adapted to a restricted habitat as a result of natural selection within a local environment. ectoparasite. An external parasite, which on fishes is often a louse or leech. It may also be a parasite found in the gill cavity. eddy. A circular movement of water in which currents flow counter to each other or past obstructions. EEZ. Exclusive economic zone; waters out to 200 nautical miles (370.4 km) from shore where international waters begin. It is reserved for the country’s exploitation and management of resources. egg capsule. See egg case. egg case. The keratinous egg shell of sharks and rays. electrophoresis. The movement and separation of chemicals in a fluid medium under electrical stimulation. It is used to determine the chemical content of fishes and other organisms and thereby to distinguish and relate them. elver. Young, transparent, but round eels. emarginate. Having a slightly concave edge; often used to describe a shallowly forked tail fin. embayment. An indentation of the shoreline forming a bay. It may also refer to lagoons, fiords, etc. embedded. Enclosed by skin, lacking free edges, e.g., scales of Zoarcidae (Eelpouts). emergence. Departure of fry from the gravel into the water column. endobenthic. Infaunal, living in the sea bed. endolateral. One of the large teeth immediately on each side of the mouth opening in Petromyzontidae (Lampreys). See also anterial; exolateral; infra-oral; supraoral. engybenthic. Living or feeding near the sea bed. epibenthic. Living associated with the sea bed but just above it, from low tide to a depth of 200 m; compare benthic. epicontinental. Found in or on a continent or continental shelf. epipelagic. Living in surface waters, down to about 200 m. esca. The structure on the end of the illicium in the anglerfishes. It serves as a lure and may be worm-like, fish-like, or luminous. ethanol. C2H5OH; used as a 70%–80% solution in water for the permanent preservation and storage of fish specimens in museum collections. euryhaline. Organisms capable of withstanding a wide range of salinity and also water with a salinity of 30.1‰–40.0‰ derived from ocean salts. eurythermal. Organisms capable of withstanding a wide range of temperature. exolateral. One of the small teeth between the endolaterals (near the mouth opening) and the marginals (which line the margin of the disc) in Petromyzontidae (Lampreys). See also anterial; endolateral; infra-oral; supraoral.

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expatriate. A fish that has been removed from its usual habitat, e.g., by a current, to an environment where it cannot reproduce, such as cold Arctic waters. extralimital. Beyond the limits (of a regional study or an identification key). eyed side. The side in flatfishes bearing both eyes, the uppermost side when the fish is resting on the bottom. It is also called upper surface, but not dorsal surface as it is a flank. The opposite of blind side.

F

factory ship. A large stern trawler equipped with a plant for gutting, filleting, freezing, and storing fish, for processing fish oil and fish-meal, and sometimes for canning. It catches its own fish and takes fish from other ships to process. falcate. Sickle-shaped. family. A group of closely related genera. fang. A long, sharp tooth. farm. In respect of fishes, an aquaculture facility. fathom. 1.829 meters. fatty eyelid. Adipose eyelid. fecundity. A measure of the production of young, usually in numbers and/or weight of eggs in fishes. fetch. An area where ocean waves are being generated by the wind. filament. A thread-like projection. filamentous. Having a thread-like projection. filter feeder. A fish that obtains small particles of food by filtering them out of the water, usually with numerous, elongate, and fine gill rakers. fin fold. The median body-wall folds in members of the Myxinidae (Hagfishes); also the supposed origin of paired fins in other fishes. fingerling. An immature fish, less than one year old. fin-rays. Cartilaginous or bony rod-like supports of fins. fins. Flap-like external organs used in steering, balance, propulsion, and behaviour. They are composed of fatty tissue (hence adipose) or of rays separated by membranes. The rays may be stiffened into spines or flexible and branched (known as soft rays). Median fins include the dorsal, adipose, caudal, and anal fins, and paired fins (one on each side) are the pectoral and pelvic fins. See also under adipose fin; anal fin; caudal fin; dorsal fins; pectoral fins; pelvic fins; ventral adipose fin. fiord. A long, deep, narrow inlet of the sea between steep mountainous sides. fisher. A person participating in a fishery (gender neutral, in preference to the previously used term fisherman). Historically, most fishers were fishermen. fishing apparatus. A mechanism for attracting prey close to the mouth in anglerfishes, formed from dorsal fin spines modified into a fishing rod (illicium) with a lure (esca) at the tip. fish-meal. Ground-up fish used as fertilizer or as an ingredient in foods. flank. The side of the body. flaw lead. An opening between pack ice and land-fast ice. fluvial. Flowing (rivers); living in or migrating between main rivers and tributaries (fish). food fall. The descent of a large dead organism (such as a whale) to the sea floor, and the corpse itself on the sea floor. It then forms both directly and indirectly a nutrient supply for a variety of organisms and for fish. The term is also used for the much smaller marine snow or plankton rain. food web. The feeding connections in an ecological community. forage species. Fish that serve as food for other fish especially for commercial or sport fishes. forehead. The frontal curve of the head and the area of the head over or in front of the eyes.



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forked. In reference to the caudal or tail fin, indicating a central indentation to the posterior margin as opposed to a straight or rounded margin. The fork may be shallow or deep. fork length (FL). The distance from the anteriormost tip of the body to the innermost part of the fork of the tail fin, a measurement often used by fishery biologists. formalin. A 37%–40% solution of formaldehyde gas dissolved in water, making 100% formalin; it may contain some methanol. It is used as a fixative and preservative in fish collections because decay bacteria cannot live in a 5%–10% solution of it. It is usually replaced with ethanol for long-term storage. Formalin is a noxious chemical and should be handled with appropriate safety measures. fossa (plural, fossae). Groove, pit, or depressed area in a bone. fraenum. The tissue joining the lip to the tip of the jaw; it prevents the jaw from being projected outwards. free. Unconnected. freshet. A great rise or overflowing of a stream caused by heavy rains or melted snow. frontal. A superficial, paired, dermal bone on top of the skull above the eyes. fry. A young fish at the post-larval stage. It may include all the fish stages from hatching to fingerling. An advanced fry is any young fish at the stage from the start of exogenous feeding after the yolk has been absorbed, and a sac fry is the stage from hatching to yolk-sac absorption. In Salmonidae (Trouts and Salmons) it is the stage from the end of dependence on the yolk sac as the primary source of nutrition to the dispersal from the redd. Fish may end up in a fish fry (an indoor or outdoor party where fish are cooked and eaten). fulcral spine, fulcrum (plural, fulcra). Unsegmented, unbranched rays in front of the dorsal, anal, and caudal fins, found in Notacanthidae (Deepsea Spiny Eels) and Acipenseridae (Sturgeons). fusiform. Spindle-shaped; tapering at both ends in a streamlined fashion, e.g., Salmonidae (Trouts and Salmons).

G

gangion. A leader line or snood (a short line connecting a fishing line to the hook). gape. The mouth opening. gas bladder. A sac in the upper part of the body cavity of bony fishes, containing a mixture of gases and often used to adjust the vertical position of a fish in the water by varying the gas content. Also called swimbladder. gelatinous. Jelly-like, soft, not firm. generation time. The time required for a female to produce a reproductively active female. genital. Pertaining to the sex organs. genital palp, genital papilla. A small fleshy projection behind the anus through which eggs and sperm pass, e.g., in Cottidae (Sculpins). Also called anal papilla or urogenital papilla. genus (plural, genera). The category below family. Each genus may contain several species that are more closely related to one another than to species in other genera. The genus is the first word (always capitalized and italicized) of the scientific name. gill. A paired respiratory organ in fishes consisting of gill filaments on the gill arch in the posterior portion of the head and usually providing the primary exchange of gases between the blood and the surrounding water. gill arch. The skeletal support of the gill filaments and gill rakers. gill cavity. The space occupied by the gills. gill cover. The operculum, the bones, and the tissues covering the gills on the side of the head in bony fishes (absent in Hagfishes, Lampreys, sharks, and Skates; fleshy in chimaeras). gill filament. The blood-infused structures of the gills that take up oxygen from the water and give off waste carbon dioxide.

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gill membrane. The tissue forming the lower wall of the gill chamber, supported by the branchiostegal rays. Usually called branchiostegal membrane. gill-net. A net that is hung in the water like a curtain that catches fish by entangling them around the gills when the head is pushed through. gill opening. The single opening, or multiple openings, on the side of the head that allows inhaled water to exit after it has passed over the gills. gill pouch. The sac containing the gills and communicating with the mouth cavity, and with the exterior in Myxini (Hagfishes) and Petromyzontiformes (Lampreys). gill raker. One of a series of bony projections on the inner side of the gill that strain or retain food from inhaled water. Counts of gill rakers are used to identify some fishes, the count being of the upper-arm rakers, lower-arm rakers, or total rakers. gill slit. One of the series of openings on the head of sharks and rays that allow inhaled water to exit after it has passed over the gills. glacial maximum. The greatest extent of any glaciation in time or place, usually referring to the Pleistocene glaciation. glaciation. A period of time (thousands of years) during an ice age, marked by colder temperatures and glacier advances. glycoprotein. A natural protein in the blood of polar and cool-temperate fishes that prevents the formation of ice crystals, down to an exterior temperature of −6°C. GO2. The posterior gular organ on the chin in Platytroctidae (Tubeshoulders). gonad. The ovary and testis that produce the eggs and sperm. grilse. An immature and anadromous salmon or trout that still has the silvery body as in the sea but returns to fresh water to spawn for the first time. groundfish. Fish that live on or near the bottom, usually those sought commercially. Also called bottom fish. group-synchronous spawner. A species that mass releases eggs all at once in a group of fishes. gular. Pertaining to the gula, the region between the chin and the isthmus. gurry. Wastes from fish, such as skin, fins, viscera, mucus. gyre. An ocean current following a circular path within an ocean basin, clockwise in the northern hemisphere.

H

hadal zone. The part of the ocean below about 6,000 m, the deepest part of the ocean, named for its supposed proximity to hell. haemoglobin. A metalloprotein containing iron found in red blood cells, functioning to transport oxygen to the tissues from the gills in order to power bodily functions. halocline. The region of rapid change of salinity between two layers of different salinity. When capitalized, it refers to the lower 150 m of the Arctic Surface Layer, a complex zone with waters of varying densities. handline. A line with baited hooks on short side lines, usually laid on the bottom and set and hauled by hand. head canals. The extension of the lateral-line system on the head. The canals open to the surface through pores and contain neuromasts. The canals may be lost, and the neuromasts are exposed. head clasper. A spiny, knob-like structure in male Rhinochimaeridae (Longnose Chimaeras) and Chimaeridae (Shortnose Chimaeras) used to grasp females during copulation. head crest. A raised ridge on the head, e.g., in Pricklebacks. head spines. Spines on the head of Scorpaenidae (Scorpionfishes) and Cottidae (Sculpins). They are, from anterior to posterior over the top of the head on each side, the nasal, preocular, supraocular, postocular, tympanic, coronal (medial to the tympanic and postocular spines), parietal, and nuchal. Opercular spines are at the postero-dorsal corner of the operculum, preopercular spines line the posterior margin of the

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preoperculum, and the cleithral and postcleithral spines are just above the opercular spines on the side of the head. hermaphrodite (adjective, hermaphroditic). Having both ovarian and testicular tissue in one individual, i.e., both female and male reproductive organs. The eggs and sperm are not necessarily produced simultaneously. heterocercal. A type of tail fin in which the vertebral column turns upwards into the upper lobe, which is usually longer than the lower lobe. holostyly. A type of jaw suspension in which the upper jaw is fused to the skull, e.g., in the Rhinochimaeridae (Longnose Chimaeras) and Chimaeridae (Shortnose Chimaeras). holotype. The single specimen designated or indicated as “the type-specimen” by the author in the publication describing a new species. home range. The area over which a fish normally travels in its day-to-day activities. homing. The return to a place formerly occupied instead of to other equally probable places, e.g., a return to spawning grounds. homocercal. A type of tail fin in which the vertebral column turns upwards, but which is symmetrical when viewed externally. hoop net. A net mounted on hoops that help support the netting. horny capsule. An egg case as in the Rajidae (Skates). hybridization. The process of the interbreeding of two different species, resulting in individuals that are intermediate between the parent forms or that have a mix or an overlap of characters. hyoid arch. The series of bones between the jaws and the gill arches. hyperbenthic. Living above but close to the bottom. hypersaline. Salinity well in excess of that of sea water. hypolimnion (adjective, hypolimnetic). The cold lower layer of a stratified lake, under the epilimnion (warm uppermost layer) and beginning just below the thermocline. This layer is not directly affected by surface events. hypural plate. The flattened bones at the end of the vertebral column and the base of the caudal fin, supporting the caudal fin-rays.

I

IC. The entire ventral row of photophores running from the anterior end of the isthmus to the posteriormost photophore on the caudal peduncle. ICNAF. International Commission for the Northwest Atlantic Fisheries, which divides the northwest Atlantic into sub-areas and divisions for fishery management purposes. In Arctic waters the Canadian side of Davis Strait and Baffin Bay are Divisions 0B and 0A. illicium. A modified spine of the first dorsal fin in anglerfishes, used as a mobile fishing rod to lure prey near the mouth. incisor. A flat tooth with a more or less straight edge used for shearing, usually found at the front of the mouth. incomplete. Usually referring to the lateral line that does not extend all the way from the head to the tail fin base. Indigenous fishery. A fishery by Indigenous peoples for food, commercial, social, and ceremonial purposes. infauna. Organisms found in sea-bed sediment. inferior. Below or ventral, often used in referring to the mouth being positioned on the lower surface of the head. infracaudal luminous gland. A long median and ventral gland on the caudal peduncle of some Myctophidae (Lanternfishes). infra-oral. One of the teeth just below the mouth in Petromyzontidae (Lampreys). See also anterial; endolateral; exolateral; supraoral. infra-oral lamina. A plate bearing teeth just below the mouth in Petromyzontidae (Lampreys). infra-orbital. Below the eye, suborbital. infra-subspecific. A category or name of lower rank than subspecies and therefore not recognized by the International Code of Zoological Nomenclature.

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insertion. The posterior end of a fin at a point closest to the body; compare origin, the anterior end of the base of a fin. interorbital. The area between the eyes on top of the head; interorbital width is a measurement taken between the bony orbits over the top of the head. interspecific. Between two or more species; compare intraspecific. intertidal. The shore exposed between high and low tides. intestinal valve. A fold in the intestine that increases surface area for processing food without lengthening the intestine. Found in sharks, for example. intraspecific. Within a species. introgression. The spread of inherited characters between species by hybridization. intromittent. Capable of being inserted, as with a male reproductive organ. iridescent. Having colours of the rainbow that change depending on the angle of view. isinglass. The glutinous or gelatin-like fluid prepared from the collagen of the outer layer of gas bladders of Acipenseridae (Sturgeons) or other fishes. It is used in the clarification of wines and beers, in jams and jellies, in printing inks, and as adhesive cement. isocercal. See diphycercal. isopropyl alcohol. Used in some museum collections to preserve fish, usually at a concentration of 50% or 55%. It is less inflammable than ethanol (ethyl alcohol) and not subject to stringent controls, but may clear tissues and decalcify bones. isosmotic. Pertaining to solutions that exert the same osmotic pressure. isotherm. A map line connecting all points having the same temperature. isthmus. Tissue between the gill openings on the lower surface of the head. iteroparity (adjective, iteroparous). The condition of producing offspring in successive years or seasonal batches, as is the case in most fishes, compare semelparity. IV. A row of ventral photophores running from the anterior end of the isthmus to the origin of the pelvic fin. IVO. An interventral photophore between the pelvic fins in Platytroctidae (Tubeshoulders).

J

jellied. Also known as the jelly condition, in which the flesh has a percentage of water making it flaccid and unsuitable as food. jigging. Using a jig (one to several bare hooks attached to a weighted line). The hook(s) may have a lead head (lead moulded around the hook) and be dressed with, or have a skirt of, rubber, hair, silicone, or plastic. jugular. Of or pertaining to the throat region. The position of the pelvic fins when they lie in front of the pectoral fins on the throat.

K

karyotype. The number and appearance of the chromosomes in the nucleus of a cell. Also used for the complete set of chromosomes in a species. keel. A ridge, often on the sides of the caudal peduncle or the belly midline. keratin. A waterproof, structural protein forming horny parts of fishes, e.g., Lamprey teeth. keystone species. A species that is essential to the food web of an ecosystem, e.g., Boreogadus saida in the Arctic; a species whose loss from an ecosystem would cause a greater-than-average change in other species populations or ecosystem processes. kilo-Langley (kLy). A unit of energy distribution over an area; used to measure solar radiation or insolation (1 kilo-Langley = 11.622 kilowatt-hours per square meter). k-selection. A life history strategy characterized by slow development, low mortality, late maturity and reproduction, repeated spawnings but sometimes at intervals of several years, large body size, and long lifespan. It is an adaptation to a relatively constant environment.



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kype. The hooked and lengthened lower jaw in male Salmonidae (Trouts and Salmons).

L

labial. Pertaining to the lip; e.g., the labial furrow is a fold behind the mouth corner that provides slack skin for jaw protrusion in sharks. lacustrine. Pertaining to or inhabiting lakes or ponds. lagoon. A shallow pond or elongate channel separated from the open ocean by a sand bar or reef, or by a narrow outlet, with little or no freshwater input. lamella (plural, lamellae). A layer, a thin plate. lamina. See lamella. laminar. Blade-like. lamphedrin. An anti-coagulant fluid secreted by a gland in Petromyzontidae (Lampreys). lanceolate. Spear-shaped. land bridge. A connection between adjacent land masses, forming a dispersal or migration route for freshwater fishes or a barrier to marine fishes. It may form and recede over time. land-fast ice. Sea ice extending from the land out to sea or attached to shoals. land-locked. Living in waters shut off from the sea, or in waters with access to the sea, though such fishes are non-migrating populations. larva (plural, larvae). Young after hatching, usually different in appearance to the parents. lateral. Relating to the side or flanks. lateral line. A sense organ in a pored tube along the side of the body that detects movement, low frequency vibrations, and temperature changes. The lateral line extends onto the head in a series of head canals. lateral-line organs. Horseshoe-shaped structures left on the flank of Myctophidae (Lanternfishes) when the scales become detached. A lateral-line-organ count is often used in place of a scale count because scales are very deciduous in this family. lateral-line pores. Openings of the lateral-line organ along the side of the body. The lateral line may be completely pored from head to tail or interrupted. lateral-line scales. Scales along the lateral line, a count of which is often used to identify species. lateral-series scales. A count of scales along the flank when the lateral line is absent or incompletely pored. lead. A channel of water, especially one through ice. lectotype. One of several syntypes (every specimen in a type-series numbering two or more) designated, after the publication of a species-group name, as the type-specimen of the taxon bearing that name. Designated only where there was no original holotype. leister. A spear typically having three barbed prongs that is used in the subsistence fishery. Originally the shaft was made of wood, 12–16 feet long (3.7–4.9 m), and the prongs were made from musk-ox horn or caribou antler that was shaped after being immersed in hot water. length. See fork length; standard length; total length. leptocephalus (plural, leptocephali). The transparent, leaf-like, compressed larva of eels and their relatives such as Notacanthidae (Deep-sea Spiny Eels). level. The relationship of fin positions and other structures to each other by being level with, behind, or in front of the compared structure. light organ. Structures producing light by a chemical reaction or by means of light-producing bacteria, e.g., in the Myctophidae (Lanternfishes). line trawling. Long-line fishing. lingual. Pertaining to the tongue; or looking out from inside the mouth (opposite of labial).

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lingual lamina. A plate bearing teeth on the tongue-like piston of Petromyzontidae (Lampreys). These plates consist of an anterior or transverse lamina and a pair of posterior or longitudinal laminae. litter. Those organisms produced at multiple births, e.g., in sharks. littoral. Shore waters, down to 200 m in the sea. live-bearer. A fish giving birth to active, free-swimming young. long line. A fishing line with baited hooks set at intervals on branch lines; it may be 150 km long, have several thousand hooks, and be on the sea bed or above it supported by floats. It may be anchored or drift free and is marked by floats. longshore. Directed along the shore or existing on, frequenting, close to, or parallel to the shore. luciferase. See luciferin. luciferin. A substance concerned in bioluminescence, being oxidized in the presence of the enzyme luciferase to produce oxyluciferin and light. lumen. The cavity of any organ, duct, or sac. luminescent. Light-producing or light-reflecting. luminous organ. A light-producing structure or light organ. lunate. Crescent-shaped. lure. A general term for artificial devices meant to attract fish to strike, usually imitating fish or other prey or providing flash and movement that stimulate a strike.

M

macroalga (plural, macroalgae). Macrophyte. macrophyte. Large aquatic plants. main stem. The largest channel of a river system. malar spines. Spines found close to the edge of the disc in some male Rajidae (Skates). mandible (adjective, mandibular). The lower jaw. maxilla (plural, maxillae). A dermal bone forming part of the upper jaw. maxillary. See maxilla. medial. Towards the vertical plane running through the middle of the body. median. Situated on the vertical plane running through the middle of the body. median fins. The dorsal, adipose, caudal, and anal fins. Also called vertical fins. mediolateral. Middle of the side. melanophore. A black chromatophore or dermal pigment cell. Expansion and contraction of pigment within the cell affects the darkness or lightness of the body structure. mental crest. A crest on the chin; e.g., in Lycodes a cartilage ridge apparently used to plough through mud while blowing with the mouth to uncover buried prey. meristic character. A countable character that is serially repeated, e.g., finrays, vertebrae, scales. mermaid’s purse. An egg case of sharks and rays. meromictic lake. A permanently stratified lake, usually without oxygen (and fish) in its deeper portions, due to a density gradient and a lack of turnover. mesobenthic. Said of organisms living on the sea bed between 200 m and 1,000 m. mesopelagic. Mid-waters of the ocean between 200 m and 1,000 m, poorly lighted. metamorphosis. A marked change in the appearance and structure, as between a leaf-like larval eel and the elongate, rounded adult, or in a young flatfish before one eye migrates to the other side of the head. mid-water trawl. A net towed through the water column by one boat, the net being positioned above the bottom and below the surface. It may have otter doors that function to keep the net mouth open and depressor plates that function to make it fish deeper, or two vessels may pair trawl to keep

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the mouth open. The front net sections are often made with very large meshes or ropes, which herd the fish schools towards the net aft sections. migration. A journey to, and return from, a specific locality or habitat type. It usually occurs at a certain stage and regularly in the life cycle of a species. See anadromous; catadromous. milt. Male seminal fluid containing sperm. mitochondrial DNA. A form of DNA, the chemical basis of heredity, found in the mitochondria (energy-producing structures within cells) and used to identify and relate species. molariform. Large, flat, crushing or grinding teeth. morphometric character. A character based on measurement. morphotype. One of several morphologically distinguishable populations of a single taxon. Usually these are of no taxonomic significance, or such significance has not been determined. MS222. A fish anaesthetic applied by immersion in dosed water (tricaine methanesulphonate, methanesulfonate salt, or 3-aminobenzoic acid ethyl ester – C10H15NO5S). mtDNA. An abbreviation for mitochondrial DNA. mucus (adjective, mucous). A slippery and slimy fluid produced by membranes or glands, used to protect and moisten. multi-year ice. Ice that does not melt each summer in the Arctic as temperatures warm, and thus it has survived more than one melting season. It contains much less brine and more air pockets than first-year ice. myodome. A cavity in the skull under the brain in the postorbital region of the skull in which lodge the rectus muscles (anterior, posterior, superior, and inferior) of the eye and, anteriorly, the ethmoid for anchoring the oblique muscle (superior and inferior). myomere. A lateral body segment muscle.

N

NAFO. Northwest Atlantic Fisheries Organization, an intergovernmental agency mandated to provide scientific advice and management of fisheries in the northwestern Atlantic Ocean. NAFO Subarea 0 comprises Division 0A, Canadian waters north of 66°15' N in Davis Strait and Baffin Bay; and Division 0B, to the south of 0A almost parallel to Cape Chidley, Labrador. naked. Lacking scales, in fishes. nape. The area behind the back and top of the head. nasal. A paired dermal bone usually enclosing the nostrils. nasal capsule. The structures enclosing the nostrils. nasal head spine. See head spines. nasal lamellae. Flaps of tissue in the nostril. nasal sinus. A cavity leading into the pharynx, in which may be found one or more papillae, in Myxinidae (Hagfishes). nasohypophysial opening. An opening allowing water into the olfactory organ of Petromyzontidae (Lampreys) and Myxinidae (Hagfishes). It is on top of the head in Lampreys and in front of the head in Hagfishes. natal. Of or connected with birth, birthplace, e.g., stream or river of a fish. natal homing. Returning to the birthplace. natal river, natal stream. The waterbody in which a fish was born or hatched. nautical mile. 1,852 meters, 1.151 miles, or 1 minute latitude of the great circle of the Earth. nearshore. Shallow waters at a small distance from the shore. Also called inshore or onshore waters, although the former may be defined as nearer the coast. neoteny (adjective, neotenic). Achieving sexual maturity while the rest of the body is in the juvenile form (the juvenile form of others of the same group). neritic. Pertaining to the shallow waters on or above the continental shelf (as opposed to oceanic) from the low-tide level to a depth of 200 m; shelf fauna.

Glossary

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nest. A structure constructed by fish for deposition and protection of the eggs. neural spine. The dorsal spine on top of the neural arch of the vertebra, directed backwards. neurocranium. The part of the skull that surrounds the brain, including elements that surround the olfactory (smell), orbital (sight), and otic (hearing) capsules. neuromast. A sensory cell that detects motion or vibrations. The cell has a sensitive hair-like structure embedded in a gelatinous capsule. Neuromasts are enclosed in canals (e.g., lateral line), freely exposed, or sunk in a pit on the skin surface or in the inner ear. niche. The habitat and role of an organism (food habits, community relationships, etc.); the range of environmental space occupied by a species. nictitating eyelid. A membrane that covers the eye, an eyelid, found particularly in sharks. nocturnal. Pertaining to the night. nominate. Name-bearing taxon. North Atlantic Oscillation (NAO). Fluctuations in atmospheric pressure at sea level, controlling the strength and direction of winds in the North Atlantic Ocean. nostril. The external opening to the organ of smell. nuchal. Pertaining to the nape. nuchal head spine. See head spines. nuchal thorn. A strong spine on the nape region in Rajidae (Skates). nuptial tubercle. Small epidermal tubercles on the head, body, and fins used for contact during breeding, in defence of nests and territories, in stimulating females, and possibly for sex and species recognition. Also called breeding tubercles. nursery. An area favoured for birth or egg deposition where young can grow.

O

OA. The upper, ventro-lateral row of photophores running above the ventral series from just behind the operculum to above the anal fin. oblique. In relation to the mouth usually running from anterior to posterior at a sharp angle. Also used for other structures at an angle. occiput (adjective, occipital). The rear end of the top of the head. oceanic. Pertaining to the sea, especially when depths exceed 200 m, i.e., offshore. oceanodromous. Marine fishes that make migrations wholly in the sea. ocellus (plural, ocelli). An eye spot, a dark mark surrounded by a lighter halo. offal. A term used in fish processing for viscera, skin, bones, and trimmings that are discarded. offshore. Waters away from the shore but under a country’s jurisdiction. oil globule. A sphere of fat or oil in the yolk of some fish eggs. It varies in number, size, position, and colour and therefore is a useful character for identification. olfactory organ. The organ of smell. oligotrophy (adjective, oligotrophic). A lake condition with low productivity, hence the hypolimnion does not become depleted of oxygen. Usually having a deep, narrow basin and clear waters with little littoral vegetation. onshore. Shallow waters at a small distance from the shore. Also called inshore or nearshore waters. OP. Photophores on the gill cover, one near the anterior base of the preopercle, one in front of the anterior part of the subopercle, and one antero-dorsally to the operculum. opercle. The principal dermal bone of the gill cover. opercular flap. The fleshy edge of the whole gill cover. Also called ear flap. opercular head spine. See head spines. opercular opening. See gill opening. operculum. The main bone of the gill cover. Also called opercle. oral. Pertaining to the mouth, such as the oral chamber or the mouth cavity.



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oral disc. The circular mouth area of Petromyzontidae (Lampreys), bearing horny teeth. Also called suctorial disc. oral fimbria. One of a series of small tag-like appendages around the perimeter of the oral disc of Petromyzontiformes (Lampreys), presumed to help create an effective seal when the lamprey is attached to its host; it is also probably sensory. oral hood. Lip extensions in the form of a scoop leading to the mouth, in larval Petromyzontidae (Lampreys). ORB. Photophores near the eye; one antero-ventral to the eye is called the suborbital (or preorbital) and one postero-ventrally is called the postorbital. orbit. The skull cavity that contains the eye; also used for the eye itself. order. A group of closely related families. origin. The anterior end of the base of a fin; compare insertion, the posterior end of a fin at a point closest to the body. otic capsule. The auditory capsule. otolith. Ear stones, small calcium carbonate structures lying in the inner ear of fishes. Used to detect changes in motion and gravity by stimulation of sensory nerve cells as the fish moves. See also sagittal otolith. otophysic connection. A firm contact between the auditory capsules (inner ear) and the gas bladder, enabling the better detection of higher and lower frequencies of sound through resonation of the bladder, e.g., in Moridae (Codlings). otter trawl. A towed net that strains demersal fish out of the water. Rectangular otter boards of wood or steel on the tow ropes plane through the water and help keep the mouth open and give the trawl its name; floats on the head rope and weights on the ground line also assist in this. outlier male. A male Oncorhynchus gorbuscha (Pink Salmon) that is smaller than the dominant male, is coloured like a female, and maintains a position to one side of a spawning pair; in this fashion it is able to deposit sperm because the dominant male does not chase it away. OV. A lateral series of photophores running from behind the opercle on the head to above the pelvic fin insertion on the lower flank. ovary. The egg-producing organ. overfishing. A level of fishing effort or fishing mortality such that a reduction of this level would, in the medium term, lead to an increase in the total catch. For long-lived species, overfishing starts well before the stock becomes overfished. overwinter. Surviving winter conditions. oviduct. A tube used to carry eggs away from the ovary. oviparity (adjective, oviparous). Production of eggs that hatch and develop outside the body. ovovivaparous. Eggs develop and hatch in the mother but do not feed at maternal expense; young are born as miniature, free-swimming adults.

P

pack. Floating ice that has been driven together into a single mass. paired fins. The pectoral and pelvic fins. Other fins are called the vertical fins or median fins. palate. The roof of the mouth. palatine. One of a pair of bones on the roof of the mouth, lateral to the vomer, often bearing teeth. papilla (plural, papillae). A fleshy protuberance. parapophysis (plural, parapophyses). A long, transverse process arising from the abdominal vertebral centrum. Parapophyses serve to support epipleural ribs when present and, in Gadidae (Cods), the gas bladder. In Clupeiformes (Herrings and relatives) they are not fused to the vertebrae. parasitic males. Small male anglerfishes that live attached to the much larger female. Much of the body is degenerate and depends on the female blood supply for food. The gonads are well developed. parietal. A dermal paired bone covering the auditory region of the skull. parietal head spine. See head spines.

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parr. A young member of the Salmonidae (Trouts and Salmons) before it migrates to the sea, having dark, almost oval blotches along the side. parr mark. Round to oval or pear-shaped dark blotch on the side of young Salmonidae (Trouts and Salmons). pectoral fins. The paired fins on the side of the body just behind the gill opening or head. pedicel. A stalk-like supporting structure. pelagic. Occurring above the bottom of the sea. pelagic trawl. A net shaped like a bag that is dragged through open waters above the bottom. pelvic fins. The paired fins on the belly. They may be abdominal (remote from the pectoral fins on the belly), thoracic (on the chest), or jugular (on the throat). periglacial. Areas, objects, or processes near the edge of a glacier, or in or about glacial times. See also glaciation; deglaciation. peritoneal cavity. The coelom. peritoneum, peritoneal membrane. A membrane covering the coelom and the organs within. It may be black, silvery, speckled, or some other colour and is often used as a distinguishing character. pharynx. The part of the intestine between the oesophagus and the mouth. phenotype. The observable structural and functional properties of an organism. phenotypic plasticity. Variation in the phenotype. pheromone. A chemical secreted and released to produce a response by others of the same species. photocytes. Light-producing cells. photophore. A light-producing organ. There are various abbreviations for photophore positions, and the arrangement and counts of photophores are used in identification. See Gonostomatidae (Bristlemouths) and Myctophidae (Lanternfishes) in the “Family and Species Accounts” section for illustrations. See also AO; Dn; GO2; IVO; OA; PLO; PO; Pol; Prc; PVO; SAO; VO. phylogenetic species concept. A species is a group of organisms diagnosably distinct from other groups with a parental pattern of ancestry and descent (see Kottelat, 1997, for a discussion on this concept relevant to fishes). phylogeography. Relationships between gene genealogies (phylogenetics) and geography. physoclist (adjective, physoclistous). A species having the the gas bladder closed, with no connection to the gut. There is supposedly a connection in larvae to allow for first inflation. Adult fish must secrete gases against a pressure gradient using a gas gland and rete mirabile. The gas gland secretes lactic acid into the blood, causing a decrease in pH, which results in haemoglobin releasing oxygen that diffuses across the rete. An organ known as the oval body is the re-absorbent organ when the partial pressure of gases in the swim bladder is greater than that of the dissolved gases in the blood. physostome (adjective, physostomous). A species having the gas bladder connected by a tube to the gut. phytoplankton. Plant members of the plankton. pineal eye. A median, eye-like structure on top of the head in Petromyzontidae (Lampreys); it develops from the pineal organ. pineal organ. A light-sensitive area of the brain that influences behaviour and the melanophores. pinnate. Divided in a feathery manner with lateral processes. piscivore (adjective, piscivorous). A feeder on fish. piscivory. The act of feeding on fish (cannibalism when carried out by fish). pit organ. A neuromast set in a small depression in the skin, not enclosed in a lateral-line canal. placoid scale. The thorn-shaped scale of sharks and rays; also called denticle. It consists of a spine and a rhomboidal basal plate.

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plankton. Small animals and plants living above the bottom in the water column. Fish eggs and larvae are often members of the plankton. Zooplankton are animal plankton, and phytoplankton are plant plankton. plastic species. One showing variation in form, ecology, or behaviour, e.g., phenotypic plasticity. plate. Any flattened structure, usually an external armament in certain fishes. Pleistocene. A geological epoch of the Quaternary period, ca. 1.6 million to 10,000 years ago. plica (plural, plicae). Small skin folds. PLO. A photophore above the base of the pectoral fin. plume. Water, often turbid, beyond its usual confines, e.g., a river plume beyond the estuary or river channel. PO. Pectoral fin photophore in Platytroctidae (Tubeshoulders). Pol. One or more ventro-lateral photophores above the AO series of photophores, which itself lies along the base of the anal fin. Polar Mixed Layer. The upper 50 m of the Arctic Surface Layer. In winter it becomes hypersaline as ice forms and salt is extruded; in summer it becomes hyposaline due to ice melt and freshwater inflow from terrestrial areas. polar pack. See pack. polynya. Geographically fixed open water (or low-average sea-ice thickness) surrounded by ice in the Arctic, formed by wind displacement or warm upwelling. polyphyly (adjective, polyphyletic). Having more than one origin or line of descent, not closely related. Species may be grouped polyphyletically as a convenience until a monophyletic classification can be made. population. A local group of individuals that form a potentially interbreeding community. pore. A minute opening through which fluids can pass. post-. Behind; often used in measurements for identification, e.g., post-branchial (behind the gill region). postcleithrum (plural, postcleithra). A dermal bone of the pectoral girdle postero-ventral to the cleithrum, which supports it dorsally. There may be one to several postcleithra (absent in some species). posterior. Behind; opposite of anterior. posterior myodome. A cavity in the skull under the brain in the postorbital region of the skull in which lodge the rectus muscles (anterior, posterior, superior, and inferior) of the eye and, anteriorly, the ethmoid for anchoring the oblique muscle (superior and inferior). postglacial. After glaciation. post-larva (plural, post-larvae). A larval fish after the yolk sac has been absorbed, when the structure is still unlike the juvenile fish. postocular head spine. See head spines. postorbital. Behind the orbit. Also refers to a bone in the series around the eye. post-smolt. A stage in a salmonid’s development, from its departure from a river until the onset of wide annulus formation at the end of the first winter in the sea. post-temporal fossa. A cavity in the bone connecting the pectoral girdle to the skull. ppm. Parts per million. ppt. Parts per thousand (‰), used for measuring salinity, for example; see also psu. Prc. The photophores on the lower half of the caudal peduncle at the caudal fin base. pre-. Before, anterior. Often used in measurements for identification, e.g., predorsal length (the length from the dorsal fin origin to the snout tip); or in describing the locality of structures, e.g., pre-anal (in front of the anus or anal fin). pre-branchial. Before the gill area.

Glossary

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precaudal pit. The notch on the caudal peduncle just anterior to the tail fin. Also called caudal pit. precaudal vertebrae. Those vertebrae lying anterior to the caudal region, usually bearing ribs rather than haemal spines. predorsal. Before the dorsal fin. Also, bones located between the head and the dorsal fin above the vertebral column. pre-larva. An early stage in fish developement in which the yolk sac is present and fins are not fully developed. premaxilla (plural, premaxillae). A superficial bone on each side of the upper jaw anterior to the maxilla, usually having teeth. premaxillary. See premaxilla. preocular head spine. See head spines. preorbital spine. A spine on the lachrymal bone. prepelvic. Lying before the pelvic fin. prickle. A small sharp spine, sometimes modified from scales, e.g., in Liparidae (Snailfishes). primary production. The transfer of the sun’s energy to biomass via photosynthesis by plants. procurrent. Inclined forward. productivity. The rate of generation of biomass in an ecosystem. See also primary production; secondary production. pro-glacial lake. A lake of glacial origin beyond the limits of the glacier. prolarva. A larva still bearing yolk. protractile. Capable of being retracted. protrusible. Capable of being thrust out. pseudobranch. A small gill on the inside of the gill cover. Also called the hemibranch or pseudobranchium. pseudo-population. A non-maturing group of fish that may drift into Arctic waters as young or larvae. pseudospine. Unsegmented, unpaired, modified soft-rays in the first dorsal fin of some Macrouridae (Grenadiers) and Phycidae (Phycid Hakes), not true spines but spine-like. psu. Practical salinity units, a measurement of salinity from conductivity, temperature, and pressure (as a ratio it should not have units, but these are applied in some works). Salinities measured as parts per thousand (ppt) and estimated as practical salinity units are usually quite similar. See also ‰; ppt. pterygiophore. The skeletal base of the fins inside the body. pulse fishing. Harvesting a stock of fish, then moving on to other stocks or waiting until the original stock recovers. purse seine. A seine used to encircle a school of fish in open water. It can be set at speed from a large powered vessel while the other end is held by a small boat. PVO. Photophores below the pectoral fin and above the PO series photophores in Myctophidae (Lanternfishes). pyloric caeca. Finger-like blind sacs attached to the junction of the stomach and the intestine that serve to aid in digestion.

Q

quincunx. An arrangement of five objects in a square with one in the middle. quota. Amount of a catch or harvest allocated in a time period by a governmental authority. It may refer to a fishery as a whole or to the amount allocated to an individual or company of the total allowable catch.

R

radius (plural, radii). A groove radiating out from the centre of a scale. rays. Dermal, rod-like, and bilaterally paired supports of the fins. They are usually flexible, branched, and segmented (soft rays), but the term rays may also be used to include the stiff, unbranched, unsegmented spines. Some soft rays are inflexible and resemble true spines. Rudimentary rays are short, unbranched, and unsegmented rays found at the origin of fins.



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Ray counts (usually omitting rudimentary rays) are used in identification. Soft ray counts are usually given in the scientific literature in Arabic numerals, and spines in Roman numerals. recreational fishery. Fishing for personal use, entertainment, sport, and challenge; it does not include the sale of catch but does include the businesses associated with it. recruitment. The new members and/or the numbers of fishes born in a given year. recurved. Curved upwards. redd. A hollow made in, and covered by, gravel for egg deposition and protection by Salmonidae (Trouts and Salmons). refugium (plural, refugia). An isolated locality often surrounded by a different climate or habitat; often a centre for relicts; a refuge in which organisms survived during glaciation, having escaped major climatic change. relictual. Said of survivors (relicts) of a formerly wide-spread fauna existing in certain isolated areas or habitats. resident. Refers to non-migratory fish, usually salmonids. It may refer to fish that remain within a circumscribed area. residual. Members of a generally anadromous species that do not migrate but remain in fresh water and do not spawn; a juvenile that matures sexually before it smolts and goes to sea. reticulate. Having the appearance or form of a network. retrorse. Turned backwards. riffle. A shallow stream habitat with broken or choppy surface water and a moderate to fast current. Gradient is about 1%–4%. riverine. Pertaining to a river, river-inhabiting. roe. The mass of eggs in the ovary. roe fishery. A fishery directed to capturing fish for their eggs or roe content, e.g., Pacific Herring (Clupea pallasii). rosette. A pleated structure resembling a flower. rostral bar. An anterior extension of the skull in some Rajidae (Skates). rostral cartilage. An element in the olfactory region of the skeleton. rostrum. A snout-like extension of the head, e.g., in members of the Rajidae (Skates). ROV. An abbreviation for remotely operated vehicle; one that dives in deep water to film deep-sea organisms such as fishes. It is controlled from onboard ship and carries no passengers. r-selection. A life history strategy characterized by early maturity, rapid growth, large numbers of young produced at an early age, small body size, high mortality, and short lifespan. This strategy is an adaptation to an unpredictable environment such as that found in the Arctic. run. Seasonal migration undertaken by fish, usually as part of their life history.

S

saddle. Pigmentation straddling the back in the form of a saddle. sagittal otolith. The otolith in the sacculus, the largest of the otoliths in most species. salinity. Salt content of water, usually measured directly and chemically, expressed as parts per thousand (ppt or ‰); full sea water is 35‰. See also psu. salter. Sea-run Brook Trout (Salvelinus fontinalis). SAO. A ventral-lateral row of photophores above the anal fin origin in Myctophidae (Lanternfishes). sapitut. A traditional fishing site, often built of rocks across a river narrows to trap migrating fish, usually Salvelinus alpinus (Arctic Char). scale pocket. The skin fold in which a scale is situated. scales. Scales form an overlapping protective covering to the body and sometimes to the head and fins. See also ctenoid scale; cycloid scale; placoid scale.

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scales in series. The scale rows running along the body from behind the gills to the base of the caudal fin. They are counted when there is no lateral line or when the lateral-line scales are of different size and the count is of the row below or above. scapular thorns. Thorns on the shoulder region (each side of the anterior midline of the body) in Rajidae (Skates). school. A group of fishes, usually of the same species, that move in unison. scientific name. The name in Latin of an animal or a plant used by scientists. It consists of two words: (1) the genus name and (2) the species name, the species epithet, or the trivial name, e.g., Boreogadus saida, the scientific name of the Arctic Cod. Convention demands that this name be underlined, italicized, or put in bold face or in some other fashion to distinguish it from the rest of the printed text. scute. A modified scale with a sharp or blunt ridge as in Clupeidae (Herrings) and Sturgeons (Acipenseridae); as a bony flank armour as in Gasterosteidae (Sticklebacks); at the snout tip in Macrouridae (Grenadiers). sea-run. A freshwater population that enters the sea for feeding, e.g., Salvelinus fontinalis (Brook Trout) in Hudson Bay. seasonal ice. Ice that melts away and reforms annually; compare multi-year ice. secondary production. Production by animals. sedentary. Fish that do not move far. segmented rays. Fin-rays that are divided into segments along their length. semelparity (adjective, semelparous). Organisms having only one brood per lifetime, the adult dying after spawning, e.g., Pacific salmons. sensu lato. In the broad sense; e.g., using a taxon inclusively to embrace two or more taxa that other authors consider distinct. serial spawner. A fish that spawns more than once in a season. See also batch spawner. sexual dimorphism. Exhibiting differing body parts between males and females. shelf. Any anatomical structure that protrudes and has a flattened dorsal surface. The shallower waters inshore below 200 m. shelf zone. The shallower waters inshore below 200 m. shoulder. The area in Rajidae (Skates) behind the eyes on each side of the midline of the back; the area on the flank behind the head in Platytroctidae (Tubeshoulders). shoulder organ. A black-lined pit or sac under the cleithrum in Platytroctidae (Tubeshoulders) that is connected to the exterior by a tube visible above the pectoral fin. The sac produces a luminous fluid that can be discharged to confuse predators. sill. Shallow entrance to an enclosed body of deeper water or between two large bodies of deep water. simultaneous hermaphrodite. Both sexes in the same individual at the same time. sinistral. Used to denote flatfishes with eyes on the left side; compare dextral. sinus. A space in the tissues, usually a blood cavity as in Myxinidae (Hagfishes). smolt. A young salmonid with silvery flanks obscuring the parr marks, migrating to the sea. smoltify. The process of transforming from a parr to a smolt. The functioning of the gills and the kidneys must be reversed. smoltification. See smoltify. sneaker. A small non-dominant male fish that attempts to fertilize eggs by darting suddenly onto the nest site. snout. The tip of the head in front of the eyes. soft rays. See rays. sp. (plural, spp.) An abbreviation for species, used in the sense of the specific part or species epithet of the scientific name to indicate an undescribed species or unidentified species. spatulate. Spoon-shaped, with a concave depression.

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spawn. Laying and fertilizing eggs; also the eggs of fishes. spearing. Using a spear to catch fish. See also leister. species. The scientific name of an animal or plant that represents a group of populations of interbreeding or potentially interbreeding individuals that are reproductively isolated from other groups. species complex. A convenient term used to recognize a group of closely related species that have not yet been adequately distinguished by scientists. species name. The species epithet or trivial name; combined with the genus name it forms the scientific name unique to a species. sphenotic spines. A spine borne on the bone that separates the orbital and otic regions of the skull at the back of the head in Oneirodidae (Dreamers). spine. A stiff, pointed, and sharp process on the head or supporting a fin. spinulated. Having spinules. spinule. Small spine. spiracle. A small cleft between the eye and the gill slits in sharks and Skates. Skates use it to inhale water. spiral valve. A spiral or helical fold in the mid-gut of Petromyzontidae (Lampreys), sharks, chimaeras, and Acipenseridae (Sturgeons). It functions to increase the surface area for processing food without lengthening the intestine. sport fishery. The capture of fishes using rod and reel; regulated by licences for residents and non-residents. Almost entirely in the fresh waters of the Arctic. squalene. A low specific gravity hydrocarbon found in the livers of squaloid sharks and used in cosmetics, lipsticks, hair-setting preparations, moisturizers, and lubricants for fine machinery. squamation. The arrangement of scales. stable isotope analysis. Using the distribution of certain isotopes in muscle tissue, for example, to determine diet, trophic level, etc. Muscle tissue can be used because it shows the assimilated nutrients in the diet as stomachs may be empty. The three major isotopes used in aquatic ecosystem foodweb analysis are 13C, 15N, and 34S. stamukhi. Pressure ridges of sea ice, parallel to the coast in water depths of 15–50 m, forming the boundary between moving Arctic pack ice and coastal ice. standard length (SL). The distance from the anterior tip of the body to the end of the hypural bones or the vertebral column. This posterior point can be found by flexing the caudal fin laterally. Standard length is the usual scientific measurement of body length in preserved fishes because the caudal or tail fin may often have been broken off. stay. A fleshy, cartilaginous, or bony supporting strut, as in suborbital stay, found in Scorpaenidae (Scorpionfishes). stellate. Star-shaped. stenohaline. Said of organisms that are able to withstand only small changes or ranges in the salinity derived from ocean salts. Opposite of euryhaline. stenothermy (adjective, stenothermic). The ability to withstand only a narrow temperature range. stern chasers. Luminous glands on the upper and lower surfaces of the caudal peduncle in Lanternfishes. They are believed to help distract predators that strike at this attractive region and are confused when the lights are extinguished, and the lanternfish darts away. stern trawler. A trawler whose nets are retrieved from the stern with a derrick or a gantry, up a ramp, over a roller, or over the bulwark. stick. Fish marketed in the form of rectangular sticks that are cut from a block of frozen fish fillets, breaded, and fried in fat or sold frozen for cooking. stock. A distinct population or quantity of fish in a given area. stray. A fish that has wandered away from its normal habitat or distribution. See also vagrant. stripe. Pigment in elongated and horizontal arrangement (contrast bar). stylophthalmoid. The larvae of Lanternfishes characterized by eyes on stalks.

Glossary

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subgenus (plural, subgenera). A category of the genus-group subordinate to genus or an individual taxon of the category “subgenus.” subfamily. A category of the family-group subordinate to family or an individual taxon of the category “subfamily,” e.g., Salmoninae. The recommended ending is inae. submental. Under the chin. suborbital. Below the eye. suborbital stay. See stay. subsistence fishery. A fishery in which the fish are consumed directly by the families of the fishers rather than being bought by middlemen and sold at the next larger market. subsistence harvest. Subsistence fishery. subspecies (adjective, subspecific). A geographic group of local populations differing from, and isolated from, other such groups. The groups do not usually meet but can interbreed if they do, unlike most species. The subspecies name is the third Latinized word in the scientific name. The current tendency is to recognize any diagnosable population as a species, such that subspecies are not necessary. substrate. The sea bottom or bottom materials. subterminal. Before the end and ventral. Often refers to the mouth position in fishes or a tail fin notch in sharks. suctorial disc. See oral disc. sulcus. A groove or channel, found in some otoliths. superior. Above, dorsal. supplemental. Additional. supplemental pore. Additional pore found outside those of the lateral line on the flank. supracaudal luminous gland. A long median and dorsal gland on the caudal peduncle of some Myctophidae (Lanternfishes). supracleithral head spine. See head spines. supracleithrum. The paired dermal bone of the secondary pectoral girdle above the cleithrum and below the post-temporal bone. supralateral. Above the lateral region. supramaxilla. The bone on the upper, rear end of the maxilla. supraocular head spine. See head spines. supraoral. Above the mouth; also said of teeth above the oral opening in Petromyzontidae (Lampreys). See also anterial; endolateral; exolateral; infra-oral. supraoral lamina. A plate bearing teeth above the mouth in Petromyzontidae (Lampreys). supraorbital photophore. A light organ on the postero-dorsal margin of the orbit in the Myctophidae (Lanternfishes). Abbreviated as SuO. sustainable yield. The quantity of fish that can be taken from a stock (usually on an annual basis) without severely depleting or eliminating that stock. Sverdrup. A unit of flow used in oceanography for ocean currents. One Sverdrup equals one million cubic meters per second. Named after the Norwegian oceanographer and Arctic explorer H.U. Sverdrup (1888–1957). sweep net. A large net usually payed out from a boat in a circle and then pulled to shore. swimbladder. See gas bladder. sympatric. Sharing, at least in part, the same geographical range. symphysis. Cartilaginous joint where two bones are united, e.g., the two halves of the lower jaw. synonym. Each of two or more names with different spelling applied to the same taxon.

T

t. Metric ton or tonne (1,000 kilograms). Sometimes abbreviated as mt. TAC. Total allowable catch, the weight of fish of a given species or type allowed to be caught by commercial fishermen in a year according to government regulation.



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tagging. Attachment of numbered and addressed labels or tags, usually of plastic or metal, to a live fish that is then released and hopefully recaptured at a later date. Alternatively, fins may be clipped in a marking system. Used to estimate growth, stock size, mortality, and movement. tail fin. The fin at the end of the body. Also called caudal fin. tapetum lucidum. A silvery layer in the choroid of the eye (in Elasmobranchii, some marine teleosts, and many bathypelagic species), formed of guanine. It probably acts to reflect light back through the retina to increase the stimulus. taxon (plural, taxa). Any taxonomic unit such as a species, genus, or family. taxonomy. The process of distinguishing and naming taxa. TEK. Abbreviation for traditional ecological knowledge. telescopic. Resembling a telescope, capable of magnification. tenaculum. A supplemental clasper in Holocephali (Chimaeridae and Rhinochimaeridae, Shortnose and Longnose Chimaeras); one in front of the pelvic fin (prepelvic tenaculum or clasper) and one on the forehead (cephalic tenaculum or head clasper). tentacle. A short, fleshy appendage, e.g., in Scorpaenidae (Scorpionfishes). terete. Round in cross-section and tapering. terminal. At the end. Said of the mouth when it is at the tip of the body rather than ventral or subterminal. testis (plural, testes). The sperm-producing organ. tetraploid. Having a chromosome complement of 4n, twice the usual number of most organisms. Not uncommon in fishes. thermocline. The zone of rapidly changing temperature between the warm upper layer (epilimnion) and the lower cold layer (hypolimnion) of water. Characterized by a temperature change of 1C° or more per meter. THO. The thoracic photophore in Platytroctidae (Tubeshoulders). thoracic. Pertaining to the thorax, the area anterior to the abdomen. Said of the pelvic fins when they are under the pectoral fins. thorn. A sharp, often curved, prickle or spine. thornlet. A small thorn. throat. The anterior ventral surface of the fish, under the head. tide pool. An area of seashore covered and exposed by the tide daily. tongue. A fleshy structure attached to the floor of the mouth that may bear teeth. Not usually extensible in fishes. tooth plate. A flattened structure bearing teeth, or a type of tooth that is in the form of a flattened plate, e.g., in Rhinochimaeridae (Longnose Chimaeras), Chimaeridae (Shortnose Chimaeras), and Myxinidae (Hagfishes). total length (TL). The greatest length of a fish from the anterior tip of the head to the tip of the tail. traditional ecological knowledge (TEK). Traditional knowledge of Indigenous people regarding local environmental resources. It can be very useful, for example, in remote regions that have little recorded scientific data. trammel net. A net made up of a fine-meshed net between two large-meshed nets that are extended between two ropes, the fish being trapped by becoming wrapped in the fine-meshed net that forms bags around the fish. transformation. The change in structure from a larval form to an adult, in Petromyzontidae (Lampreys). transplantation. The process in which a species is intentionally or accidentally transported and released by humans into an environment inside its present range. Transpolar Drift. Movement of surface water (i.e., to 50 m depth, consisting of Pacific waters, entrained fresh water, and ice) across the Asian side of the North Pole from the area of the Bering Strait to the Fram Strait and the Barents Sea. trawl. A bag-shaped net that is towed behind a ship either along the sea floor or in mid-water, having a buoyed head rope and a weighted foot rope to

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keep the net mouth open. It may be towed by one or two ships. See also bottom trawl; otter trawl; pelagic trawl. tribe. A category of the family group that is subordinate to the subfamily; or an individual taxon of the category “tribe.” Its recommended name ending is ini. trifid. Divided into three. trifurcated. Divided into three. trilobed. Divided into three lobes. trivial name. The species name; combined with the genus name it forms the scientific name unique to a species. It is also used for the vernacular name. troller. A vessel used in trolling. trolling. Trailing a fishing line (or trolling line) behind a moving boat. trophic. Pertaining to nutrition, the gathering and eating of food. trophic level. The position that an organism occupies in a food chain (organisms eating and being eaten). Trophic level 1 is the primary producer (plants and algae), level 2 is herbivores, level 3 is predators, level 4 is carnivores eating other carnivores, and level 5 is apex predators that have no predators. Since organisms feed at more than one trophic level, their score may be fractional and not a simple whole number. trophic web. The interrelationships of food and the organisms that feed on it. truncate. Square-cut; said of the tail fin when it has a straight edge. tubercle. A small projection from a surface. See also nuptial tubercle. tusk. An enlarged tooth. tympanic head spine. See head spines. type subspecies. The original populations described as a new species; they bear the species name as the subspecies name. A subspecies distinct from the original species requires a new, subspecific name, the third Latinized word in a scientific name.

U

uncinate. Hooked at the tip. unculi. Horny projections arising from single cells. underslung jaw. Subterminal mouth or jaw. underyearling. A fish less than one year of age. unicuspid. Teeth having a single point or cusp. unsegmented ray. A soft ray, usually small, without segments and found at the beginning of a fin. upper slope. The upper part of the continental slope, a steep slope connecting the deep ocean basins with the shallow continental-shelf waters. upwelling. An upward movement of cold, nutrient-rich water from the ocean depths, often associated with great production of fish and fisheries. urea. (CH2)2CO, a waste product of metabolism excreted via the kidneys but also found in Elasmobranchii blood to maintain osmotic balance. It is degraded to ammonia by enzymes, leading to a characteristic pungent odour in spoiled Elasmobranchii. urogenital papilla. See anal papilla; genital papilla. urostyle. The small upturned posterior tip of the vertebral column, generally formed of a slender, pointed rod of cartilage; fused vertebrae and associated elements (in homocercal caudal fins of Teleostomi); or the fan-like series of bones articulating with the last true vertebra, including preural centra, ural centra, epurals, and hypurals. Counted as one vertebra in some vertebral counts, but not counted in others.

V

vagrant. A species that has strayed beyond its natural range but has not established reproducing populations. validate. To confirm the accuracy of ageing methods that use skeletal structures like otolith zones by showing that the zones do refer to age. VAV. A ventro-lateral row of photophores running from the pelvic fin insertion to the anal fin origin.

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veil. A large mucous sheet containing eggs. velar apparatus. A structure lying at the junction of the oesophagus (above) and the blind-ending branchial tube (below) in adult Petromyzontidae (Lampreys). It bears the velar tentacles that project anteriorly into the pharynx. velar tentacles. Tentacles at the junction of the pharynx and the oesophagus in Petromyzontidae (Lampreys), probably used to deflect large food particles from the branchial region. venom. The poison secreted by a gland. vent. The posterior opening of the intestine, gonads, and kidney ducts in front of the anal fin. ventral. Of or pertaining to the lower body. ventral adipose fin. A fin anterior to the anal fin on the belly. ventral photophores. A row of light organs along the abdomen on each side of the midline. ventro-lateral. On the lower side of the flank. ventrum. The underside of the body, the belly. vermiculation. Worm-track marking. vertebra. A segment of the spinal column composed of a central body (the centrum), with a neural arch and a spine above and a haemal arch and a spine below. vertical fins. The dorsal, caudal, and anal fins that stand erect from the body. vertical migration. The upward and downward movement of fish in the ocean. Movement is usually into surface waters at night to feed and into deeper waters during the day to avoid predators. villiform. In the shape of slender, finger-like processes, often forming a patch or band with the appearance of velvet. visceral cleft. The jaw, hyoid, and gill arches. Vn. A photophore in front of the eye below the olfactory capsule of Myctophidae (Lanternfishes). VO. A row of photophores on the abdomen behind the pelvic fins. vomer. A dermal bone in the middle of the roof of the mouth.

W

water column. The water mass between the surface and the bottom. weir. A structure of stone or branches used to trap migratory fishes on the spawning run. window. A clear area; e.g., in species of Macrouridae (Grenadiers) with a light organ this is an area without scales. wings. The enlarged, wing-like pectoral fins of Rajidae (Skates).

Y

year-class. All the individuals of a population of fishes born or hatched in the same year, e.g., 1995. After this brood has been recruited to the fishery, it appears year after year until all of its members die, and it is referred to as the 1995 year-class throughout its life. The biomass of a year-class initially increases as fish grow; reaches a peak at a certain age depending on the species, food availability, and mortality; and then starts to decrease as fish die off and growth slows. yolk-sac larva. An early-stage larval fish with a sac containing yolk used for nourishment. young of the year (YOY). Members of age group zero, from transformation to juvenile, until the first of January in the northern hemisphere or the first of July in the southern hemisphere.

Z

zooplankton. Animal members of the plankton.

Glossary

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BIBLIOGRAPHY Brian W. Coad and James D. Reist

An extensive bibliography on marine fishes of Arctic Canada, comprising over 5,340 references, was published separately (Coad and Reist, 2016) and is available at http://waves-vagues.dfo-mpo.gc.ca/ Library/364139.pdf. This bibliography is meant as a reference guide to literature on marine fishes in Arctic Canadian waters. It is a source for the works on which the species accounts and the general text of this book are based, and it is also one of the sources for mapping, along with museum collections and anecdotal reports. The online version is searchable. Several synoptic works have been consulted and provide data on fishes found in Arctic Canada. They include works that have information on more than one species; systematic and taxonomic works that review families or other taxa; and extralimital studies that are sources of information on poorly known species. The following bibliography contains these synoptic works to avoid constant repetition and also contains some works mentioned in the introduction. Allen, M.J., & Smith, G.B. 1988. Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific. (U.S. National Oceanic and Atmospheric Administration, National Marine Fisheries Service Technical Report No. 66). iii + 151 pp. Anderson, M.E. 1988. Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere: I. The Antarctic and Subantarctic regions. In L.S. Kornicker (Ed.), Biology of the Antarctic Seas XIX (Antarctic Research Series No. 47) (pp. 59–113). Washington, DC: American Geophysical Union. Andriashev, A.P. 1955. A contribution to the knowledge of the fishes from the Bering and Chukchi seas. (Special scientific report, U.S. Fish and Wildlife Service Fisheries No. 145). 81 pp. Andriyashev, A.P. 1954. Ryby severnykh morei SSSR [Fishes of the northern seas of the USSR]. Akademii Nauk SSSR, Opredeliteli po Faune SSSR 53, Izdatel’stvo Akademii nauk SSSR, Moscow and Leningrad. 566 pp. (Translation, 1964, by Israel Program for Scientific Translations, Jerusalem. 617 pp.). Andriyashev, A.P., & Chernova, N.V. 1995. Annotated list of fishlike vertebrates and fish of the Arctic Seas and adjacent waters. Journal of Ichthyology, 35(1), 81–123. Backus, R.H. 1957. The fishes of Labrador. Bulletin of the American Museum of Natural History, 113(4), 273–338. Berkes, F. 2012. Sacred Ecology (3rd ed.). New York: Routledge.



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Boschung, H.T., Williams, J.D., Gotshall, D.W., Caldwell, D.K., & Caldwell, M.C. 1983. The Audubon Society field guide to North American fishes, whales, and dolphins. New York: Alfred A. Knopf. 848 pp. Carpenter, K.E. (Ed.). 2002. The living marine resources of the Western Central Atlantic. (FAO Species Identification Guide for Fishery Purposes, and American Society of Ichthyologists and Herpetologists Special Publication No. 5). Rome: Food and Agriculture Organization of the United Nations. 3 volumes, xiv + 2,127 pp. Coad, B.W. 1992. Guide to the marine sport fishes of Atlantic Canada and New England. Toronto: University of Toronto Press. xiii + 307 pp. (Also available in French as Guide des poissons marins de pêche sportive de l’Atlantique canadien et de la Nouvelle-Angleterre. La Prairie, QC: Broquet. xii + 400 pp.). Coad, B.W. 1995. Expedition field techniques: Fishes (2nd ed.). London: Expedition Advisory Centre, Royal Geographical Society. v + 97 pp. Coad, B.W. 2011. Bibliography of Donald Evan McAllister. Canadian Field-Naturalist, 124(4)(2010), 336–56. Coad, B.W. 2017. Fishes of Canada: Annotated checklist. www.briancoad.com. Coad, B.W. & Reist, J.D. 2004. Annotated list of the Arctic marine fishes of Canada. (Canadian Manuscript Report of Fisheries and Aquatic Sciences No. 2674). iv + 112 pp. Coad, B.W. & Reist, J.D. 2016. Bibliography on the marine fishes of Arctic Canada. (Canadian Manuscript Report of Fisheries and Aquatic Sciences No. 3101). v + 509 pp. Retrieved from http://waves-vagues.dfo-mpo. gc.ca/Library/364139.pdf. Coad, B.W., with Waszczuk, H. & Labignan, I. 1995. Encyclopedia of Canadian fishes. Ottawa: Canadian Museum of Nature and Canadian Sportfishing Productions. viii + 928 pp. Collette, B.B., & Klein-MacPhee, G. 2002. Bigelow and Schroeder’s fishes of the Gulf of Maine (3rd ed.). Washington, DC, and London: Smithsonian Institution Press. xxxiv + 748 pp. Cook, F.R., & Coad, B.W. 2002. Donald Evan McAllister: Curatorial and research contributions at Canadian Museum of Nature. Sea Wind, 15(1/2) (2001), 46–53. Cook, F.R., Coad, B.W., & Renaud, C.B. 2002. Donald Evan McAllister, 1934– 2001. Copeia, 2002(3), 890–4. Cook, F.R., Coad, B.W., Renaud, C.B., Gruchy, C.G., & Alfonso, N.R. 2011. Donald Evan McAllister, 1934–2001: The growth of ichthyological research at the National Museum of Canada / Canadian Museum of Nature. Canadian Field-Naturalist, 124(4)(2010), 330–5. Cook, F.R., Gruchy, C.G., & Coad, B.W. 2001. Donald Evan McAllister, 1934– 2001: A tribute. Amphipacifica, 3(2), 1–2.

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Encyclopedia of Life. 2013. Encyclopedia of Life. www.eol.org. Eschmeyer, W.N. (Ed.). 1998. Catalog of fishes (Vols. 1–3). San Francisco: California Academy of Sciences. Also in CD-ROM and continuously updated online at the California Academy of Sciences website, http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. Eschmeyer, W.N., & Herald, E.S. 1983. A field guide to Pacific Coast fishes of North America from the Gulf of Alaska to Baja California. (Peterson Field Guide Series No. 28). Boston: Houghton Mifflin. xii + 336 pp. Ford, J., McDowell, G., & Pearce, T. 2015. The adaptation challenge in the Arctic. Nature Climate Change, 5, 1046–53. Froese, R., & Pauly, D. (Eds.). 2013. FishBase. www.fishbase.org. Gebruk, A.V., Krylova, E.M., Lein, A.Y., Vinogradov, G.M., Anderson, E., Pimenov, N.V., Cherkashev, G.A., & Crane, C. 2003. Methane seep community of the Håkon Mosby mud volcano (the Norwegian Sea): Composition and trophic aspects. Sarsia, 88(6), 394–403. George, C., Moulton, L., & Johnson, M. 2009. A field guide to the common fishes of the North Slope of Alaska. Barrow, AK: North Slope Borough, Department of Wildlife Management. ii + 93 pp. Grey, M. 1956. The distribution of fishes found below a depth of 2000 meters. Fieldiana, Zoology, 36(2), 73–337. Hart, J.L. 1973. Pacific fishes of Canada. (Bulletin of the Fisheries Research Board of Canada No. 180). ix + 740 pp. Hildebrandt, N., Bergmann, M., & Knust, R. 2011. Longevity and growth efficiency of two deep-dwelling Arctic zoarcids and comparison with eight other zoarcid species from different climatic regions. Polar Biology, 34(10), 1523–33. Hureau, J.C., & Monod, T. (Eds.). 1978. Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean (Vols. 1–2, with supplement). Paris: UNESCO, Paris. (First published, 1973). Integrated Taxonomic Information System. 2013. www.itis.gov/. Jackson, J.B.C., Kirby, M.X., Berger, W.H., Bjorndal, K.A., Botsford, L.W., Bourque, B.J., Bradbury, R.H., Cooke, R., Erlandson, J., Estes, J.A., Hughes, T.P., Kidwell, S., Lange, C.B., Lenihan, H.S., Pandolfi, J.M., Peterson, C.H., Steneck, R.S., Tegner, M.J., & Warner, R.R. 2001. Historical overfishing and the recent collapse of coastal ecosystems. Science, 293, 629–38. Kottelat, M. 1997. European freshwater fishes: An heuristic checklist of the freshwater fishes of Europe (exclusive of former USSR), with an introduction for non-systematists and comments on nomenclature and conservation [Supplement No. 5]. Biologia, Bratislava, 52, 1–271. Kottelat, M., & Freyhof, J. 2007. Handbook of European freshwater fishes. Cornol, Switzerland: Kottelat. Berlin: Freyhof. xiii + 646 pp. Krupnik, I., Aporta, C., Gearhard, S., Laidler, G.J. & Kielsen Holm, L. (Eds.). 2010. SIKU: Knowing our ice; Documenting Inuit sea ice knowledge and use. New York: Springer. xxxi + 501 pp. Lee, D.S., Gilbert, C.R., Hocutt, C.H., Jenkins, R.E., McAllister, D.E., & Stauffer, J.R. 1980. Atlas of North American freshwater fishes. Raleigh: North Carolina State Museum of Natural History. x + 854 pp. Legendre, V., Hunter, J.G., & McAllister, D.E. 1975. French, English, and scientific names of marine fishes of Arctic Canada / Noms français, anglais et scientifiques des poissons marins de l’Arctique canadien. Syllogeus, 7, 1–15. Majewski, A.R., Atchison, S., MacPhee, S., Eert, J., Niemi, A., Michel, C., & Reist, J.D. 2017. Marine fish community structure and habitat associations on the Canadian Beaufort shelf and slope. Deep Sea Research Part I: Oceanographic Research Papers, 121, 169–82. McAllister, D.E. 1990. A list of the fishes of Canada / Liste des poissons du Canada. Syllogeus, 64, 1–310. McAllister, D.E., Legendre, V., & Hunter, J.G. 1987. Liste des noms inuktitut (esquimaux), français, anglais et scientifiques des poissons marins du Canada Arctique / List of Inuktitut (Eskimo), French, English and scientific

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names of marine fishes of Arctic Canada. (Canadian Manuscript Report of Fisheries and Aquatic Sciences, 1932). v + 106 pp. McAllister, D.E., & Steigerwald, M.B. 1986. Bibliography of the marine fishes of Arctic Canada. (Canadian Manuscript Report of Fisheries and Aquatic Sciences, 1909). v + 108 pp. McPhail, J.D., & Lindsey, C.C. 1970. Freshwater fishes of northwestern Canada and Alaska (Bulletin of the Fisheries Research Board of Canada No. 173). 381 pp. Mecklenburg, C.W., & Anderson, M.E. 2015. Reassessment of multiple species of Gymnelus (Teleostei: Zoarcidae) in Pacific Arctic and boreal regions. Zootaxa, 3948(2), 263–88. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., & Steinke, D. 2016. Pacific Arctic marine fishes. (Conservation of Arctic Flora and Fauna Monitoring Series Report No. 23). Akureyi, Iceland. v + 377 pp., appendices. Mecklenburg, C.W., Mecklenburg, T.A., & Thorsteinson, L.K. 2002. Fishes of Alaska. Bethesda, MD: American Fisheries Society. xxxvii + 1037 pp. Mecklenburg, C.W., & Steinke, D. 2015. Ichthyofaunal baselines in the Pacific Arctic region and RUSALCA study area. Oceanography, 28(3), 158–89. Milkov, A., Vogt, P., Cherkashev, G., Ginsburg, G., Chernova, N., & Andriashev, A. 1999. Sea-floor terrains of Håkon Mosby Mud Volcano as surveyed by deep-tow video and still photography. Geo-Marine Letters, 19(1), 38–47. Mistry, J., & Berardi, A. 2016. Bridging indigenous and scientific knowledge. Science, 352, 1274–5. Møller, P.R. 2001. A new zoarcid, Lycodes mcallisteri, from eastern Arctic Canada (Teleostei: Perciformes). Ichthyological Research, 48, 111–16. Møller, P.R., Nielsen, J.G., Knudsen, S.W., Poulsen, J.Y., Sünksen, K., & Jørgensen, O.A. 2010. A checklist of the fish fauna of Greenland waters. Zootaxa, 2378, 1–84. Morrow, J.E. 1980. The freshwater fishes of Alaska. Anchorage, AK: Alaska Northwest Publishing. xv + 248 pp. Nelson, J.S. 2006. Fishes of the world (4th ed.). Hoboken, NJ: John Wiley. xix + 601 pp. Nichols, T., Berkes, F., Jolly, D., Snow, N.B, & the Community of Sachs Harbour. 2004. Climate change and sea ice: Local observations from the Canadian western Arctic. Arctic, 57, 68–79. Nielsen, J., Hedeholm, R.B., Heinemeier, J., Bushnell, P.G., Christiansen, J.S., Olsen, J., Ramsey, C.B., Brill, R.W., Simon. M., Steffensen, K.F., & Steffensen, J.F. 2016. Eye lens radiocarbon reveals centuries of longevity in the Greenland shark (Somniosus microcephalus). Science, 353(6300), 702–4. Nielsen, J.G., Bertelsen, E., & Nystrøm, B.O. 1992. Fisk i grønlandske farvande. Nuuk, Greenland: Atuakkiorfik. 65 pp. + 2 pp. Nozères, C., Archambault, D., Chouinard, P.-M., Gauthier, J., Miller, R., Parent, E., Schwab, P., Savard, L., & Dutil, J.-D. 2010. Guide d’identification des poissons marins de l’estuaire et du nord du golfe du Saint-Laurent et protocoles suivis pour leur échantillonnage lors des relevés par chalut entre 2004 et 2008 / Identification guide for marine fishes of the estuary and northern Gulf of St Lawrence and sampling protocols used during trawl surveys between 2004 and 2008. (Rapport technique canadien des sciences halieutiques et aquatiques / Canadian Technical Report of Fisheries and Aquatic Sciences No. 2866). xi + 157 pp. Okamura, O., Amaoka, K., Takeda, M., Yano, K., Okada, K. & Chikuni, S. (Eds.). 1995. Fishes collected by the R/V Shinkai Maru around Greenland. Tokyo: Japan Marine Fishery Resources Research Center. 304 pp. Orr, J.W., Kai, Y., & Nakabo, T. 2015. Snailfishes of the Careproctus rastrinus complex (Liparidae): Redescription of seven species in the North Pacific Ocean region, with the description of a new species from the Beaufort Sea. Zootaxa, 4018(3), 301–48. Page, L.M., & Burr, B.M. 1991. A field guide to freshwater fishes: North America north of Mexico. (Peterson Field Guide Series No. 42). Boston: Houghton Mifflin. xii + 432 pp.

Bibliography

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Page, L.M., Espinosa-Pérez, H., Findley, L.T., Gilbert, C.R., Lea, R.N., Mandrak, N.E., Mayden, R.L., & Nelson, J.S. 2013. Common and scientific names of fishes from the United States, Canada, and Mexico (7th ed.). (American Fisheries Society Special Publication No. 34). ix + 384 pp. Quast, J.C., & Hall, E.L. 1972. List of fishes of Alaska and adjacent waters with a guide to some of their literature. (U.S. National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Technical Report No. SSRF-658). iv + 47 pp. Rathwell, K.J., & Armitage, D. 2016. Art and artistic processes bridge knowledge systems about social-ecological change: An empirical examination with Inuit artists from Nunavut, Canada. Ecology and Society, 21(2), 21. Retrieved from http://dx.doi.org/10.5751/ES-08369-210221. Reid, W.V., Berkes, F., Wilbanks, T., & Capistrano, D. (Eds.). 2006. Bridging scales and knowledge systems: Concepts and applications in ecosystem assessment. Washington, DC: Island Press. xii + 368 pp. Robins, C.R., Bailey, R.M., Bond, C.E., Brooker, J.R., Lachner, E.A., Lea, R.N., & Scott, W.B. 1991. Common and scientific names of fishes from the United States and Canada (5th ed.). (American Fisheries Society Special Publication No. 20). 183 pp. Robins, C.R., & Ray, G.R. 1986. A field guide to the Atlantic Coast fishes of North America. (Peterson Field Guide Series No. 32). Boston: Houghton Mifflin. xi + 354 pp. Scott, W.B., & Crossman, E.J. 1973. Freshwater fishes of Canada. (Bulletin of the Fisheries Research Board of Canada No. 184). xi + 966 pp. (Reprinted in 1979 with supplements). Scott, W.B., & Scott, M.G. 1988. Atlantic fishes of Canada. (Canadian Bulletin of Fisheries and Aquatic Sciences No. 219). xxx + 731 pp.



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Sears Foundation for Marine Research (various authors). 1948–89. Fishes of the western North Atlantic. (Memoirs of the Sears Foundation for Marine Research No. 1). New Haven, CT. Soltwedel, T., Jaeckisch, N., Ritter, N., Hasemann, C., Bergmann, M., & Klages, M. 2009. Bathymetric patterns of megafaunal assemblages from the arctic deep-sea observatory Hausgarten. Deep-Sea Research, Part. I, 56(10), 1856–72. Species at Risk Public Registry. 2012. Ottawa: Government of Canada. Retrieved from www.sararegistry.gc.ca/. Steigerwald, M.B., & McAllister, D.E. 1982. List of the Canadian marine fish specimens in the National Museum of Natural Sciences, National Museums of Canada. Syllogeus, 41, 1–30. Whitehead, P.J.P., Bauchot, M.-L., Hureau, J.-C., Nielsen, J., & Tortonese, E. (Eds.). 1984–6. Fishes of the north-eastern Atlantic and the Mediterranean (Vols. 1–3). Paris: UNESCO. Wienerroither, R., Johannesen, E., Dolgov, A., Byrkjedal, I., Bjelland, O., Drevetnyak, K., Eriksen, K.B., Høines, Å., Langhelle, G., Langøy, H., Prokhorova, T., Prozorkevich, D., & Wenneck, T. 2011. Atlas of the Barents Sea Fishes. (Institute of Marine Research and Polar Research Institute of Marine Fisheries and Oceanography Joint Report Series No. 1-2011). 272 pp. Wilimovsky, N.J. 1954. List of the fishes of Alaska. Stanford Ichthyological Bulletin, 4, 279–94. Wilimovsky, N.J. 1958. Provisional keys to the fishes of Alaska. Juneau, AK: Fisheries Research Laboratory, U.S. Fish and Wildlife Service. 113 pp.

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ILLUSTRATION CREDITS Brian W. Coad

Data sources and licences follow for maps in the introduction. Illustrations are then summarized according to the three sections in which they appear: the introduction; “Keys”; and “Family and Species Accounts.” Copyright of figures, illustrations, and photographs from Fisheries and Oceans Canada (DFO) sources is ultimately retained by the department (copyright, Her Majesty the Queen in Right of Canada, as represented by the Minister of Fisheries and Oceans). The copyrighted items from the Canadian Museum of Nature are noted as “CMN” or “CMNFI” (for the fish collection).

Abbreviations BC Royal British Columbia Museum, Victoria, BC CAS California Academy of Sciences, San Francisco CMN Canadian Museum of Nature, Ottawa CMNFI Fish Collection, Canadian Museum of Nature, Ottawa DFO Fisheries and Oceans Canada IMS Institute of Marine Science, University of Alaska Fairbanks ROPOS Remotely Operated Platform for Ocean Sciences SL standard length total length TL UAM University of Alaska Museum Fairbanks Zoological Institute, Russian Academy of Sciences, ZIN St Petersburg

GIS Data Sources Bathymetry • Bathymetry for central Arctic Ocean: International Bathymetric chart of the Arctic Ocean (IBCAO); discrete depth layers derived from IBCAO by Humfrey Melling • Bathymetry for North American waters: Government of Canada, Natural Resources Canada, Canada Centre for Remote Sensing. 2006. The Atlas of Canada. Ottawa: Retrieved from http://www.arcgis.com/home/item.html?id=5d7f1b87d34a4fb29248c2d6885114a0

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Geopolitical Boundaries • “World countries,” developed by Esri. Retrieved from http:// www.arcgis.com/home/item.html?id=­3864c63872d84aec9193361 8e3815dd2 • North American geopolitical boundaries: Government of Canada. 2006. The Atlas of Canada • Canadian territorial seas and Exclusive Economic Zone: Atlas of Canada 1,000,000 National Frameworks Data: Administrative Boundaries. Retrieved from http://geogratis.gc.ca/api/en/ nrcan-rncan/ess-sst/0d2b6f01-fe48-521f-aa7c-a177613c56dd. html

Hydrology • World rivers and lakes: http://www.naturalearthdata.com/ downloads/10m-physical-vectors/10m-rivers-lake-centerlines/ • Canadian rivers and lakes: Atlas of Canada 1,000,000 National Frameworks Data: Hydrology; Drainage Network; Arctic. Retrieved from http://geogratis.gc.ca/api/en/nrcan-rncan/esssst/9e21be32-8e11-53bb-b713-20a5fa8103cd.html • Canadian drainage basins: data derived from Atlas of Canada 1,000,000 National Frameworks Data: Hydrology; Drainage Areas (WSC sub-sub drainage areas). Retrieved from http://geogratis. gc.ca/api/en/nrcan-rncan/ess-sst/30b33615-6dda-51a5-a9dd308802714a28.html

Snow and Ice • Polar pack minimum ice extent, 1969–99 median: North American Atlas: Sea Ice. Retrieved from http://open.canada.ca/data/ en/dataset/9588a081-6175-4390-bac5-49bfadd33f6d • September monthly median sea ice extent: data from National Snow and Ice Data Center • Glaciated areas: National Snow and Ice Data Center

Illustration Credits

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Licences • Atlas of Canada, North American Atlas, and Canadian Ice Service: “Open Government Licence Canada,” http://open.canada. ca/en/open-government-licence-canada • National Snow and Ice Data Center: data usage and citation policy, http://nsidc.org/about/use_copyright.html

• Esri: redistribution rights, http://www.esri.com/~/media/Files/ Pdfs/legal/pdfs/redist_rights_103.pdf?la=en • Natural Earth data: terms of use, http://www.naturalearthdata. com/about/terms-of-use/

SECTION OPENERS Illustration

Source

Title page Large male; photograph by Neil Mochnacz, DFO

Salvelinus malma

Introduction Davis Strait

CMN; photograph by Roger D. Bull, 26 July 2013

Checklists of Species Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax

Careproctus reinhardti

Keys CMNFI 1959-0380, Québec, Ungava Bay, Koksoak River at Fort Chimo; drawing by John L. Tottenham

Gasterosteus aculeatus

Family and Species Accounts Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax

Artediellus atlanticus

Introduction Illustration

Source

Environment 1. Polar projection of the Arctic Ocean

Drawing by Shannon MacPhee with assistance from H. Melling, DFO

2. Typical temperature and salinity profiles

Drawing by Michelle Kamula, DFO

3. Water masses present across the Arctic Ocean

Drawing by Michelle Kamula, DFO

4. Surface currents and ice movements of the Arctic Ocean

Drawing by Elizabeth Worden and Shannon MacPhee, DFO

5. Current flows affecting Pacific waters in the Arctic

Drawing by Elizabeth Worden and Shannon MacPhee, DFO

6. Current flows affecting the Atlantic Layer in the Arctic

Drawing by Elizabeth Worden and Shannon MacPhee, DFO

7. Canadian Arctic marine waters with contributing freshwater drainage basins

Drawing by Shannon MacPhee and Chantelle Sawatzky, DFO

8. Polynyas and shore leads of the Canadian Arctic

Drawing developed by Oceans Management, DFO, and modified by Chantelle Sawatzky (after Hannah, Dupont, & Dunphy, 2009)

9. Eastern Arctic marine food web – humans

Drawing by Chantelle Sawatzky, DFO (after Dick & Chambers, 2005)

10. Eastern Arctic marine food web – marine mammals

Drawing by Chantelle Sawatzky, DFO (after Dick & Chambers, 2005)

11. Eastern Arctic marine food web – Centroscyllium

Drawing by Chantelle Sawatzky, DFO (after Dick & Chambers, 2005)

12. Western Arctic – estuarine food web

Drawing by Chantelle Sawatzky, DFO (after Mathias, 2013)

13. Western Arctic – ice-edge food web

Drawing by Chantelle Sawatzky, DFO (after Mathias, 2013)

14. Western Arctic – marine food web

Drawing by Chantelle Sawatzky, DFO (after Mathias, 2013)



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Illustration

Source

15. Map of northern Canada

Natural Resources Canada (Open Government Licence, Canada, http://open. canada.ca/en/open-government-licence-canada (http://www.nrcan.gc.ca /earth-sciences/geography/atlas-canada/selected-thematic-maps/16886)

16. Ecoregions of Canadian marine waters

Drawing after Coad and Reist (2004)

History of Research 17. Stenodus leucichthys (as Salmo mackenzii)

After Richardson (1836a)

18. Gymnelus viridis

After Richardson (1855)

19. Myoxocephalus scorpius

CMNFI 1958-0258, from Amadjuak Bay, Baffin Island; watercolour drawn by J. Dewey Soper, August 1926

20. Boreogadus saida

CMN, from Amadjuak Bay, Baffin Island; watercolour drawn by J. Dewey Soper, July 1926

21. MV Calanus contending with ice en route to the Belcher Islands, 12 July 1959

Canadian Meteorological and Oceanographic Society photograph archives, and H.E. Grainger

22. MV Salvelinus, icebound in Starvation Cove on the south shore of Victoria Island, 23 August 1968

Canadian Meteorological and Oceanographic Society photograph archives, and J. Boulva

Habitats 23. Beaufort Sea ice, August 2014

Kelly Young, DFO

24. Banks and Victoria Islands

Wikimedia Commons, NASA, 14–16 June 2002

25. Sea ice off Baffin Island

Wikimedia Commons, NASA Earth Observatory, 30 July 2009

26. Spring in the Canadian Arctic

Wikimedia Commons, NASA Earth Observatory, 16 June 2010

27. Northeast Baffin Island

Wikimedia Commons, north of Clyde River, 7 August 1997; photograph by Ansgar Walk, under the Creative Commons licence, https://creativecommons. org/licenses/by-sa/3.0/

28. Cliffs near Arctic Bay

Wikimedia Commons, 9 July 2011; photograph by Timkal, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0/

29. Sachs Harbour (Ikahuak)

Wikimedia Commons, 6 June 2006; photograph by Timkal, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0/

30. Kugluktuk

Wikimedia Commons, 12 July 2012; photograph by Timkal, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0/

31. Davis Strait

CMN; photograph by Roger D. Bull, 20 July 2013

32. Pauline Cove, Herschel Island

Wikimedia Commons, 30 June 2005; photograph by Maedward, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0/

33. East coast of Baffin Island

CMN; photograph by Roger D. Bull, 26 July 2013

34. HMCS Labrador in ice, Baffin Bay, 1955

Canadian Meteorological and Oceanographic Society photograph archives

35. James Bay in summer and winter

Wikimedia Commons, NASA/JPL-Caltech

36. Darnley Bay, August 2015

Darcy McNicholl at DFO and University of Manitoba

37. Baffin Bay at 600 m, September 2012, Lycodonus mirabilis

Ellen Kenchington, DFO

38. Baffin Bay at 600 m, September 2012, Lyodes cf. eudipleurostictus

Ellen Kenchington, DFO

39. Baffin Bay, September 2012, bottom invertebrates and possibly a Boreogadus saida (centre)

Ellen Kenchington, DFO

40. Davis Strait, bottom habitat, October 2012

Ellen Kenchington, DFO

41. Kelp in Browns Harbour, Nunavut, 22 July 2015

Darcy McNicholl at DFO and University of Manitoba

42. Scott Inlet, 2013, epibenthic habitat

Evan Edinger and Bárbara de Moura Neves of DFO and ROPOS

43. Grounded iceberg, 15 feet depth, Resolute Bay

Kathleen Conlan, CMN

44. Habitat associations of selected Arctic marine fishes

Drawing by Sheila Atchison, DFO

592

Illustration Credits

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Illustration

Source

Arctic Fish, Northern Cultures, and Traditional Ecological Knowledge 45. Fish lure, 1921, Copper Inuit, Coronation Gulf, Nunavut, of bear tooth, sinew, and bone, 3.2 x 9.2 cm, collected by Captain Joseph F. Bernard while exploring Coronation Gulf on his schooner Teddy Bear

Photograph from the Canadian Museum of History, Gatineau, with permission

46. Sculpin, ivory, Middle Dorset Culture, Mansel Island, Nunavut

Photograph from the Canadian Museum of History, Gatineau, with permission

47. Lithograph of woman with fish, Egevadluq Ragee, 1977, 38 x 47 cm

Photograph from the Canadian Museum of History, Gatineau, reproduced with the permission of Dorset Fine Arts, Cape Dorset, Nunavut

Keys Illustrations for the “Keys” section were drawn mainly by Susan Laurie-Bourque with the exception of the illustration of teeth in the mouth by Charles H. Douglas. Images of whole fish and some body parts are mostly from the “Family and Species Accounts” section, with the following exceptions. Lethenteron camtschaticum

CMNFI, body, 229 mm TL, no locality; drawing by John L. Tottenham

Coregonus autumnalis

CMNFI 1960-0456, 321 mm SL, NWT, Peel Channel near Aklavik, Mackenzie River; drawing by John L. Tottenham

Coregonus clupeaformis

CMNFI 1960-0456, 206 mm SL, NWT, Peel Channel near Aklavik, Mackenzie River; drawing by John L. Tottenham

Coregonus laurettae

After Coad, with Waszczuk and Labignan (1995)

Coregonus nasus

CMNFI 1960-0456, 343 mm SL, NWT, Peel Channel near Aklavik, Mackenzie River; drawing by John L. Tottenham

Coregonus sardinella

CMNFI 1960-0462, 231 mm SL, NWT, Peel Channel near Aklavik; drawing by John L. Tottenham

Oncorhynchus gorbuscha

CMNFI, no locality, 533 mm SL; drawing by John L. Tottenham

Oncorhynchus keta

CMNFI, no locality, 644 mm SL; drawing by John L. Tottenham

Oncorhynchus kisutch

After Evermann and Goldsborough (1907a)

Oncorhynchus nerka

After Goode (1884)

Oncorhynchus tshawytscha

After Evermann and Goldsborough (1907a)

Salmo salar

CMNFI 1978-0113, 540 mm TL, Québec, Matamec River near Sept-Îles; drawing by Charles H. Douglas

Salvelinus alpinus

Anadromous male; composite drawing by Gerald Kuehl through a contract to DFO

Salvelinus fontinalis

CMNFI; drawing by Charles H. Douglas

Salvelinus malma

Anadromous male; composite drawing by Gerald Kuehl through a contract to DFO

Salvelinus namaycush

CMNFI 1963-0208, 360 mm SL, Nunavut, lake on Broughton Island; drawing by John L. Tottenham

Stenodus leucichthys

CMNFI 1962-0477, 433 mm SL, Yukon, Shingle Point, Mackenzie Bay; drawing by John L. Tottenham



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Family and Species Accounts The sources for the black-and-white line drawings in the “Family and Species Accounts” section are given in an abbreviated form. Lengths are given as reported or as determined from specimens where available. Further details (specimen, size, sex, locality, habitat, artist, etc.) can be found in the appropriate museum catalogue or in the literature sources cited. The principal literature sources are Günther (1887), Goode and Bean (1896), Jordan and Evermann (1896–1900), and Jensen (1904). Some of the drawings from Goode and Bean and from Jordan and Evermann were taken from

the original illustrations in the U.S. National Museum, Washington, DC, for Coad, with Waszczuk and Labignan (1995). Other drawings are composites based on several preserved fish and a variety of literature sources, and were first published in Coad, with Waszczuk and Labignan (1995). Colour drawings and photographs are noted as such with their sources. All maps in the “Family and Species Accounts” section were prepared at the Canadian Museum of Nature by Noel Alfonso and are not listed individually.

Myxine glutinosa

1. Ventral view; colour photograph from Nozères et al. (2010) 2. Body and ventral head, CMNFI 1964-0038, 537 mm TL, Newfoundland, western part of Grand Banks; drawing by Charles H. Douglas

Lethenteron camtschaticum

1. Oral disc, female, 496 mm TL, Japan, Hokkaido Island at Fukagawa; drawing by Paul I. Voevodine 2. Colour photograph, 176 mm, UAM 4622, eastern Bering Sea, 2002 3. CAS 230132, 474 mm TL, eastern Chukchi Sea, 2007; colour photograph of oral disc by Catherine W. Mecklenburg, CAS

Harriotta raleighana



After Jordan and Evermann (1896–1900)

Rhinochimaera atlantica



Composite drawing by Susan Laurie-Bourque

Hydrolagus affinis



After Goode and Bean (1896)

Apristurus profundorum



After Goode and Bean (1896)

Centroscyllium fabricii



After Goode and Bean (1896)

Centroscymnus coelolepis



After Goode and Bean (1896)

Somniosus microcephalus

1. Body and upper and lower teeth, redrawn after Bigelow and Schroeder (1948); illustration from CMNFI, but artist unknown 2. Ca. 4 m female on sea ice of Baffin Bay, 6 April 2008; photograph by Susan Dennard, with permission from Aaron Fisk, University of Windsor

Somniosus pacificus



Amblyraja hyperborea

1. Home Bay, Davis Strait (69.3631° N, 64.8467° W) at 732 m, CCGS Amundsen; colour photograph by Bárbara de Moura Neves of DFO and ROPOS 2. Dorsal view and egg case composite from Bigelow and Schroeder (1953a), Günther (1887), and Jensen (1948); drawings by John L. Tottenham 3. Ventral view from Günther (1887)

Amblyraja jenseni



Amblyraja radiata

1. Colour photograph from Nozères et al. (2010) 2. Body, ventral, and lateral views, egg case, CMNFI 1963-0133, 480 mm TL, Newfoundland, about 80 km off southeastern tip of Labrador; drawing by John L. Tottenham

Bathyraja sp.



Bathyraja spinicauda

1. Colour photograph from Nozères et al. (2010) 2. Composite drawing by Susan Laurie-Bourque

Malacoraja spinacidermis



After Coad, with Waszczuk and Labignan (1995)

Rajella bathyphila



Composite drawing by Susan Laurie-Bourque

Rajella fyllae



After Coad, with Waszczuk and Labignan (1995)

Rajella lintea



After Coad, with Waszczuk and Labignan (1995)

Acipenser fulvescens



CMNFI 1962-0474, 522 mm TL, Québec, St Lawrence River at Montréal East; drawing by John L. Tottenham

Notacanthus chemnitzii

1. Colour photograph from eastern Arctic by David Hardie, Dalhousie University, Halifax 2. After Goode and Bean (1896)

Polyacanthonotus rissoanus



After Goode and Bean (1896)

Simenchelys parasitica



Body and head after Collett (1896)

594

After Hart (1973) from Garman (1913)

After Coad, with Waszczuk and Labignan (1995)

Body and egg case, CMNFI 1977-1692, 152 mm TL, NWT, Franklin Bay, 38 km east of Cape Bathurst; drawing by Susan Laurie-Bourque

Illustration Credits

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Species

Source

Synaphobranchus kaupii

1. Colour photograph from Nozères et al. (2010) 2. Ventral view showing gill slits after Roule (1919) 3. Body after Goode and Bean (1896)

Nemichthys scolopaceus

1. Colour photograph from Nozères et al. (2010) 2. After Hart (1973), BC 62-162, 850 mm TL, California, off Point Loma; drawing by D.R. (Bon) Harriott

Serrivomer beanii

1. Colour photograph from Nozères et al. (2010) 2. After Goode and Bean (1896)

Saccopharynx ampullaceus

1. Body after Goode and Bean (1896) 2. Underside of body with filaments; open mouth, after Bertin (1934) and Harwood (1827); drawings by John L. Tottenham

Eurypharynx pelecanoides

1. After Goode and Bean (1896) 2. Colour drawing after Zugmayer (1911)

Clupea harengus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1984-0256, 297.3 mm TL, Québec, St Lawrence River, southwest of Rivière-du-Loup; drawing by Charles H. Douglas

Clupea pallasii



Argentina silus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1964-0031, 212 mm TL, Newfoundland, western part of Grand Banks; drawing by Charles H. Douglas

Bathylagus euryops



After Goode and Bean (1896)

Holtbyrnia anomala



Composite drawing by Susan Laurie-Bourque

Maulisia mauli



Composite drawing by Susan Laurie-Bourque

Maulisia microlepis



Composite drawing by Susan Laurie-Bourque

Normichthys operosus



Composite drawing by Susan Laurie-Bourque

Platytroctes apus

1. Colour drawing after Roule (1919) 2. After Günther (1887)

Alepocephalus agassizii



After Goode and Bean (1896)

Alepocephalus bairdii



After Goode and Bean (1896)

Bajacalifornia megalops



Composite drawing by Susan Laurie-Bourque

Bathytroctes sp.



Composite drawing by Susan Laurie-Bourque

Rouleina maderensis



After Markle (1978)

Xenodermichthys copei

1. Colour photograph from Nozères et al. (2010) 2. After Goode and Bean (1896)

Mallotus villosus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1960-0465, male 126 mm, female 113 mm, Yukon, Pauline Cove, Herschel Island; drawings by John L. Tottenham

Osmerus dentex

1. Adult, 110 mm FL, Phillips Bay, Yukon, August 2008; colour photograph by Colin Gallagher, DFO 2. CMNFI 1960-0484, 231 mm SL, Yukon, Thetis Bay at Simpson Point, Herschel Island; drawing by John L. Tottenham

Osmerus mordax

1. CMNFI, no locality; drawing by Charles H. Douglas 2. Nova Scotia, Lower Prospect, Halifax County; colour drawing by G.E. Johnston

Coregonus artedi



Coregonus autumnalis

1. Composite drawing by Gerald Kuehl through a contract to DFO 2. Juvenile; colour photograph from National Oceanic and Atmospheric Administration (NOAA Fisheries, Alaska Regional Office), https://alaskafisheries.noaa.gov/habitat/nearshore-fish-photos

Coregonus clupeaformis



Composite drawing by Gerald Kuehl through a contract to DFO

Coregonus nasus



Composite drawing by Gerald Kuehl through a contract to DFO

Coregonus sardinella



Composite drawing by Gerald Kuehl through a contract to DFO

CMNFI 1960-0484, Yukon, Thetis Bay at Simpson Point, Herschel Island; drawing by John L. Tottenham

CMNFI 1963-0171, 262 mm SL, Québec, stream at Povungnituk, northeast Hudson Bay; drawing by John L. Tottenham



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Species

Source

Oncorhynchus gorbuscha

1. Male silver; colour drawing by Paul Vescei through a contract to DFO 2. Male spawner; colour drawing by Paul Vescei through a contract to DFO 3. Female spawner; colour drawing by Paul Vescei through a contract to DFO

Oncorhynchus keta

1. Male silver; colour drawing by Paul Vescei through a contract to DFO 2. Male spawner; colour drawing by Paul Vescei through a contract to DFO 3. Female spawner; colour drawing by Paul Vescei through a contract to DFO

Oncorhynchus kisutch

1. Adult silver; colour drawing by Paul Vescei through a contract to DFO 2. Male spawner; colour drawing by Paul Vescei through a contract to DFO

Oncorhynchus nerka

1. Male silver; colour drawing by Paul Vescei through a contract to DFO 2. Male spawner; colour drawing by Paul Vescei through a contract to DFO

Oncorhynchus tshawytscha

1. Juvenile of 101 mm; colour photograph from National Oceanic and Atmospheric Administration (NOAA Fisheries, Alaska Regional Office), https://alaskafisheries.noaa.gov/habitat/ nearshore-fish-photos 2. Male silver; colour drawing by Paul Vescei through a contract to DFO 3. Male spawner; colour drawing by Paul Vescei through a contract to DFO

Prosopium cylindraceum



Salmo salar

1. Adult male, Gulf of St Lawrence; colour drawing by Paul Vescei through a contract to DFO 2. Male spawner; colour drawing by Paul Vescei through a contract to DFO

Salvelinus alpinus

1. Male, 670 mm and 3,300 g, from Darnley Bay near Lasard Creek, NWT, 24 July 2011; colour photograph by Colin Gallagher, DFO 2. Resident; composite drawing by Gerald Kuehl through a contract to DFO 3. Silver female; colour drawing by Paul Vescei through a contract to DFO 4. Anadromous male spawner; colour drawing by Paul Vescei through a contract to DFO 5. Salvelinus alpinus/malma, Tree River (Kogluktualuk), Nunavut (see Dolly Varden text for complexities of taxonomy and identities in this area); colour photograph by David Hardie, Dalhousie University, Halifax

Salvelinus fontinalis

1. CMNFI 1958-0248, 248 mm SL, Ontario, Algonquin Park; (inset) CMNFI 1959-0256, 88 mm TL, Québec, 8 km north of Cap-à-l’Aigle; drawings by John L. Tottenham 2. Male spawner, Rupert River watershed, Québec; colour drawing by Paul Vescei through a contract to DFO

Salvelinus malma

1. Residual male; composite drawing by Gerald Kuehl through a contract to DFO 2. Silver, 600 mm FL, Vittrekwa River, NWT (66.90° N, 135.55° W); colour drawing by Paul Vescei through a contract to DFO 3. Silver male; colour drawing by Paul Vescei through a contract to DFO 4. Silver female; colour drawing by Paul Vescei through a contract to DFO 5. Anadromous male spawner; colour drawing by Paul Vescei through a contract to DFO 6. Anadromous female spawner; colour drawing by Paul Vescei through a contract to DFO 7. Large male; photograph by Neil Mochnacz, DFO

Salvelinus namaycush

1. Oceanic; colour drawing by Paul Vescei through a contract to DFO 2. Hood River estuary; colour drawing by Paul Vescei through a contract to DFO

Stenodus leucichthys



Composite drawing by Gerald Kuehl through a contract to DFO

Gonostomatidae Family



Abbreviations for photophores; composite drawing by Susan Laurie-Bourque

Cyclothone microdon



Composite based on Clemens and Wilby (1961) and Smith (1953); drawing by John L. Tottenham

Sigmops bathyphilus

1. After Goode and Bean (1896) 2. Colour drawing after Zugmayer (1911)

Argyropelecus gigas



After Coad, with Waszczuk and Labignan (1995)

Astronesthes cf. richardsoni



After Regan and Trewavas (1929)

Borostomias antarcticus



Composite drawing by Susan Laurie-Bourque

Chauliodus sloani



After Goode and Bean (1896)

Malacosteus niger

1. After Goode and Bean (1896) 2. Colour drawing after Zugmayer (1911)

Rhadinesthes decimus

1. Composite drawing by Susan Laurie-Bourque 2. Colour drawing after Zugmayer (1911)

Stomias boa

1. After Goode and Bean (1896) 2. X-ray of CMNFI 1977-1083, Sargasso Sea, by Noel Alfonso

596

CMNFI 1958-0353, 198 mm TL adult and parr, Québec, Lake Canichico at Fort Mackenzie, Ungava; drawing by John L. Tottenham

Illustration Credits

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Species

Source

Scopelosaurus lepidus



Composite drawing by Susan Laurie-Bourque

Anotopterus pharao



After Hart (1973), BC 58-403, 318 mm TL, off British Columbia, North Pacific Weather Station (Stonetown); drawing by D.R. (Bon) Harriott

Arctozenus risso

1. CMNFI 1973-0212, 187 mm SL, Newfoundland, Gulf of St Lawrence, 97 km off coast; drawing by Charles H. Douglas 2. Colour photograph from Nozères et al. (2010)

Magnisudis atlantica



Colour drawing after Zugmayer (1911)

Paralepis coregonoides



After Goode and Bean (1896)

Myctophidae Family



Abbreviations for photophores; composite drawing by Susan Laurie-Bourque

Benthosema glaciale



CMNFI 1959-0395, 54 mm SL, Québec, Port Burwash, Ungava Bay; drawing by John L. Tottenham

Lampanyctus crocodilus



Composite drawing by Susan Laurie-Bourque

Lampanyctus intricarius



Composite drawing by Susan Laurie-Bourque

Lampanyctus macdonaldi



CMNFI 1964-0385, 97.8 mm SL, France, Bay of Biscay; drawing by Charles H. Douglas

Myctophum punctatum

1. After Coad, with Waszczuk and Labignan (1995) 2. Colour drawing after Zugmayer (1911)

Notoscopelus kroyeri

1. Colour photograph from Nozères et al. (2010) 2. After Coad, with Waszczuk and Labignan (1995)

Protomyctophum arcticum



Composite drawing by Susan Laurie-Bourque

Symbolophorus veranyi



Composite drawing by Susan Laurie-Bourque

Coryphaenoides armatus



After Hart (1973), CMNFI 1965-0327, 233 mm TL, BC, west coast of Graham Island; drawing by D.R. (Bon) Harriott

Coryphaenoides brevibarbis



Composite drawing by Susan Laurie-Bourque

Coryphaenoides carapinus



After Goode and Bean (1896)

Coryphaenoides guentheri



After Collett (1896)

Coryphaenoides rupestris



CMNFI, ca. 395 mm SL, Newfoundland, Grand Banks; drawing by John L. Tottenham

Gadomus longifilis



After Vaillant (1888)

Macrourus berglax



CMNFI 1962-0118, 386 mm SL, Newfoundland, Grand Banks; drawing by John L. Tottenham

Nezumia bairdii

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1963-0140, 354 mm SL, Newfoundland, off Spear Point, tip of Labrador; drawing by John L. Tottenham

Trachyrincus murrayi



After Günther (1887)

Antimora rostrata



After Günther (1887)

Halargyreus johnsonii



CMNFI, no locality; drawing by Charles H. Douglas

Lepidion eques



CMNFI 1971-0001, 213 mm SL, Newfoundland, east of Funk Island Bank; drawing by Charles H. Douglas

Enchelyopus cimbrius

1. Colour photograph from Nozères et al. (2010) 2. Composite drawing by Susan Laurie-Bourque

Gaidropsarus argentatus

1. After Jordan and Evermann (1896–1900) 2. Home Bay, Davis Strait (69°36'21" N, 64°84'54" W) at 711 m, CCGS Amundsen; colour photograph by Bárbara de Moura Neves of DFO and ROPOS

Gaidropsarus ensis



Phycis chesteri

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1961-0213, Québec, Gulf of St Lawrence, south of Anticosti Island; drawing by Charles H. Douglas

Redrawn after Jensen (1948)

Arctogadus glacialis



Boreogadus saida

1. CMNFI 1960-0472, 175 mm SL, Yukon, Thetis Bay, Herschel Island; drawing by John L. Tottenham 2. North of Svalbard; colour photograph by Peter Leopold

CMNFI 1958-0087, 200 mm SL, Nunavut, Port Epworth, Coronation Gulf; drawing by John L. Tottenham



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Species

Source

Brosme brosme



CMNFI 1963-0001, 535 mm TL, Nova Scotia, ca. 102 km south-southeast of Cape Sable; drawing by Charles H. Douglas

Eleginus gracilis



CMNFI 1960-0484, 174 mm TL, Yukon, Thetis Bay at Simpson Point, Herschel Island; drawing by John L. Tottenham

Gadus morhua

1. Wikimedia Commons; colour photograph by August Linnman, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0 2. CMNFI 1984-0150, 290 mm TL, Newfoundland, western arm of Red Bay; drawing by Charles H. Douglas 3. Ogac Lake Atlantic Cod; colour photograph by David Hardie, Dalhousie University, Halifax

Gadus ogac



CMNFI 1984-0214, female, 398 mm TL, Newfoundland, Red Bay harbour; drawing by Charles H. Douglas

Lota lota



CMNFI 1960-0461, 255 mm TL, Yukon, Big Joe Creek near Old Crow; drawing by John L. Tottenham

Micromesistius poutassou



Composite drawing by Susan Laurie-Bourque

Bythites fuscus



Composite drawing by Susan Laurie-Bourque

Lophius americanus



CMNFI 1962-0083, ca. 560 mm TL, Nova Scotia, west of Sable Island; drawing by Charles H. Douglas

Himantolophus groenlandicus



Composite drawing by Susan Laurie-Bourque

Chaenophryne longiceps



Composite drawing by Susan Laurie-Bourque

Oneirodes sp.



Based on Oneirodes eschrichtii; composite drawing by Susan Laurie-Bourque

Spiniphryne gladisfenae



Composite drawing by Susan Laurie-Bourque

Ceratias holboelli

1. Colour photograph from Nozères et al. (2010) 2. After Goode and Bean (1896)

Gigantactis vanhoeffeni



Scopeloberyx robustus

1. Composite drawing by Susan Laurie-Bourque 2. Colour drawing after Zugmayer (1911)

Anoplogaster cornuta

1. After Goode and Bean (1896) 2. Colour drawing after Zugmayer (1911)

Hoplostethus atlanticus

1. Colour photograph from Nozères et al. (2010) 2. Composite drawing by Susan Laurie-Bourque

Gasterosteus aculeatus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1959-0380, Québec, Ungava Bay, Koksoak River at Fort Chimo; drawing by John L. Tottenham

Pungitius pungitius

1. Wikimedia Commons; colour photograph by Piet Spaans, in public domain 2. CMNFI 1959-0471, 51 mm, Nunavut, Frobisher Bay, southern Baffin Island; drawing by John L. Tottenham

Sebastes sp.



Colour photograph from Nozères et al. (2010)

Sebastes fasciatus



Composite drawing by Susan Laurie-Bourque

Sebastes mentella



CMNFI, no locality, 314 mm; drawing by John L. Tottenham

Sebastes norvegicus



CMNFI, no locality, 318 mm; drawing by John L. Tottenham

Artediellus atlanticus

1. CMNFI 1967-0782, male, 86.3 mm SL, Québec, Saguenay Fiord, 25 km east of Chicoutimi; drawing by Charles H. Douglas 2. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax

Artediellus scaber

1. CAS 220487, 83 mm TL, western Chukchi Sea, 2004; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1960-0466, male 61.5 mm TL and female 64.0 mm TL, Yukon, south of Simpson Point, Herschel Island; drawing by John L. Tottenham

Artediellus uncinatus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1970-0290, 54.4 mm SL, Nunavut, Frobisher Bay south of Cape Sarah; drawing by Charles H. Douglas

Gymnocanthus tricuspis

1. ZIN 53511, 146 mm TL, northeastern Chukchi Sea, 2004; colour photograph by Boris A. Sheiko, Russian Academy of Sciences, and Catherine W. Mecklenburg, CAS 2. CMNFI 1970-0285, male, 129 mm SL, Nunavut, Frobisher Bay, 8 km south-southeast of Frobisher Bay; drawing by Charles H. Douglas

598

Composite drawing by Susan Laurie-Bourque

Illustration Credits

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Species

Source

Icelus bicornis



Icelus spatula

1. Colour photograph from Nozères et al. (2010) 2. CMNFI, no locality, male, 49 mm SL; drawing by John L. Tottenham

Myoxocephalus aenaeus



Myoxocephalus octodecemspinosus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1980-0224, 247.9 mm TL, New Brunswick, Miramichi inner bay; drawing by Charles H. Douglas

Myoxocephalus quadricornis

1. Head of fish on bottom, Resolute Bay; colour photograph by Kathleen Conlan, CMN 2. (Central specimen and two lower ones) CMNFI 1974-0262, female, 231 mm SL, Nunavut, King William Island, Peterson Bay, Gjoa Haven harbour; (lower left) head outline, CMNFI 1962-0102A, male, 166 mm SL, Nunavut, Peary Channel, Amund Ringnes Island; (upper left and upper right) CMNFI 1975-1935, male, 178 and 193 mm SL, Nunavut, Creswell Bay near Union River mouth; drawings by Charles H. Douglas

Myoxocephalus scorpioides



Myoxocephalus scorpius

1. CAS 230235, 237 mm TL, eastern Bering Sea, 2006; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1981-0886, body, 216.9 mm TL, Québec, St Lawrence River, Godbout quay; drawing by Charles H. Douglas 3. CMNFI 1960-0114, head views, 111 mm SL, Nunavut, Sharat River, Frobisher Bay; drawing by John L. Tottenham

Triglops murrayi

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1959-0444, female, 50 mm SL, and male, 83 mm SL, Nunavut, Ungava Bay, northeast of Inuksulik; drawings by John L. Tottenham

Triglops nybelini

1. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax 2. CMNFI 1959-0369, male, 106 mm SL, Québec, Port Burwell harbour; drawing by John L. Tottenham (reconstruction of damaged specimen with aid from Jensen, 1944).

Triglops pingelii



CMNFI 1960-0476, male, 89 mm SL, Yukon, Collinson Head, Herschel Island; drawing by John L. Tottenham

Aspidophoroides monopterygius



CMNFI 1963-0152, 155 mm TL, Newfoundland, 103 km east of northern tip; drawing by Charles H. Douglas

Aspidophoroides olrikii

1. UAM 5833, 77 mm TL, eastern Chukchi Sea, 2008; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1960-0467, 56.5 mm SL, Yukon, 11 km northeast of Collinson Head, Herschel Island; drawing by John L. Tottenham

Leptagonus decagonus

1. Colour photographs from Nozères et al. (2010) 2. CMNFI, no locality; drawing by John L. Tottenham

Cottunculus microps

1. CMNFI 1970-0306, body and anterior head view, 163 mm SL, Québec, Saguenay Fiord, Anse-auxErables; drawing by Charles H. Douglas 2. Dorsal and ventral heads after Goode and Bean (1896)

Cottunculus thomsonii

1. Composite drawing by Susan Laurie-Bourque 2. Dorsal and ventral heads after Goode and Bean (1896)

Psychrolutes phrictus

1. U.S. National Oceanic and Atmospheric Administration, via Wikimedia Commons; colour photograph modified by Chantelle Sawatzky, DFO 2. Composite drawing by Susan Laurie-Bourque

Cyclopteropsis jordani



Redrawn after Soldatov (1939) by Susan Laurie-Bourque

Cyclopteropsis mcalpini



Composite drawing by Susan Laurie-Bourque

Cyclopterus lumpus



CMNFI 1981-0885, body and disc, 221.8 mm TL, Québec, St Lawrence River south of Godbout; drawing by Charles H. Douglas

Eumicrotremus derjugini



CMNFI 1977-1456C, 65.2 mm SL, NWT, Franklin Bay, Beaufort Sea; drawing by Charles H. Douglas

Eumicrotremus spinosus



CMNFI 1966-0563, 45 mm TL, Nunavut, Evans Strait northeast of Cape Pembroke, Coats Island; drawing by John L. Tottenham

Careproctus kidoi



After Knudsen and Moller (2008), modified by Susan Laurie-Bourque

Careproctus longipinnis



Composite drawing by Susan Laurie-Bourque

CMNFI 1960-0445, male, 59 mm TL, Nunavut, Station Bay, Ellef Ringnes Island; drawing by John L. Tottenham

CMNFI 1966-0192, 192.7 mm TL, Newfoundland, Eastport Bay; drawing by Charles H. Douglas

CMNFI 1960-0483, female, 103 mm SL, Yukon, Simpson Point, Herschel Island; drawing by John L. Tottenham



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Species

Source

Careproctus reinhardti

1. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax 2. CMNFI 1962-0290, 68 mm SL, Québec, Richmond Gulf; drawing by John L. Tottenham.

Liparis atlanticus

1. CMNFI 1959-0374, body, female, 71.5 mm SL, Québec, Pitulaksitik, Ungava Bay; drawing by John L. Tottenham 2. CMNFI 1966-0181, ventral and anterior head views, Newfoundland, Squid Cove, St John Bay; drawings by John L. Tottenham

Liparis fabricii

1. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax 2. CMNFI 1960-0476, body, male, 72 mm TL; head, male, 107 mm TL, Yukon, Collinson Head, Herschel Island; drawings by John L. Tottenham

Liparis gibbus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1964-0670, 135 mm SL, Nunavut, Roes Welcome Sound off Cape Kendall; drawing by John L. Tottenham

Liparis tunicatus

1. UAM 6211, 107 mm TL, eastern Chukchi Sea, 2009; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1962-0406, male, 134 mm TL, Nunavut, Assistance Bay, Barrow Strait, Cornwallis Island; drawing by Charles H. Douglas

Paraliparis bathybius

1. After Günther (1887) 2. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax

Paraliparis copei

1. Colour photograph from Nozères et al. (2010) 2. Composite drawing by Susan Laurie-Bourque

Paraliparis garmani



Rhodichthys regina

1. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax 2. Redrawn after Goode and Bean (1896) and Jensen (1950a) by John L. Tottenham

Caristius fasciatus

1. Colour photograph from Nozères et al. (2010) 2. Composite drawing by Susan Laurie-Bourque 3. Young specimen after Borodin (1931)

Gymnelus barsukovi



Composite drawing by Susan Laurie-Bourque

Gymnelus bilabrus



Composite drawing by Susan Laurie-Bourque

Gymnelus knipowitschi



Composite drawing by Susan Laurie-Bourque

Gymnelus retrodorsalis

1. CMNFI 1975-1953, 113 mm TL, Nunavut, Strathcona Sound, Borden Peninsula; drawing by Charles H. Douglas 2. CMNFI 1975-1953, 113 mm TL, Nunavut, Strathcona Sound, Borden Peninsula; colour drawing by Aleta Karstad Schueler

Gymnelus viridis

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1960-0096, male, 148 mm TL, Nunavut, Turton Bay, Igloolik Island; CMNFI 1959-0446, male, 169 mm TL, Nunavut, Diana Bay, Hudson Strait; CMNFI 1960-0075, male, 181 mm TL, Nunavut, Hudson Strait near Cape Dorset; CMNFI 1959-0377, female, 129 mm TL, Nunavut, Fury and Hecla Strait; drawings by John L. Tottenham

Lycenchelys kolthoffi



Composite drawing by Susan Laurie-Bourque

Lycenchelys muraena



After Goode and Bean (1896)

Lycenchelys paxillus

1. Colour photograph from Nozères et al. (2010) 2. Body and ventral head after Goode and Bean (1896)

Lycenchelys sarsii



Composite drawing by Susan Laurie-Bourque

Lycodes adolfi



Composite drawing by Susan Laurie-Bourque

Lycodes esmarkii



CMNFI 1980-0416, female, 295 mm TL, Greenland, Store Hellefiskebank; drawing by Charles H. Douglas

Lycodes eudipleurostictus



Composite drawing by Susan Laurie-Bourque

Lycodes frigidus



Composite drawing by Susan Laurie-Bourque

Lycodes jugoricus



CMNFI 1960-0472, female, 161 mm TL, Yukon, Thetis Bay, Herschel Island; drawing by John L. Tottenham

Lycodes lavalaei

1. Colour photograph from Nozères et al. (2010) 2. After Goode and Bean (1896) (was L. reticulatus)

Lycodes luetkenii



600

Composite drawing by Susan Laurie-Bourque

Composite drawing by Susan Laurie-Bourque

Illustration Credits

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Species

Source

Lycodes marisalbi



CMNFI, no locality; drawing by John L. Tottenham

Lycodes mcallisteri



Holotype, female, 230 mm SL, after Møller (2001a)

Lycodes mucosus

1. Resolute Bay; colour photograph by M. Graham, CMN 2. CAS 230183, 229 mm TL, eastern Bering Sea, 2007; colour photograph by Catherine W. Mecklenburg, CAS 3. CMNFI 1962-0406, male, 150 mm TL, Nunavut, Assistance Bay, Cornwallis Island; drawing by Charles H. Douglas

Lycodes paamiuti

1. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax 2. Holotype, male, 216 mm SL, after Møller (2001b)

Lycodes pallidus



Lycodes polaris

1. ZIN 53590, 180 mm TL, western Chukchi Sea, 2004; colour photograph by Boris A. Sheiko, Russian Academy of Sciences, and Catherine W. Mecklenburg, CAS 2. CMNFI 1960-0473, female, 176 mm TL, Yukon, southeast of Herschel Island; drawing by John L. Tottenham

Lycodes reticulatus

1. Eastern Arctic; colour photograph by David Hardie, Dalhousie University, Halifax 2. CMNFI 1963-0131, ca. 195 mm TL, Québec, Saguenay Fiord at Cap à l’Ouest; drawing by John L. Tottenham

Lycodes sagittarius



CMNFI 1974-0282, holotype, male, 278 mm TL, Alaska, Beaufort Sea, about 80 km north-northeast of Brownlow Point; drawing by Charles H. Douglas

Lycodes seminudus



After Jensen (1904)

Lycodes squamiventer



After Jensen (1904)

Lycodes terraenovae



Composite drawing by Susan Laurie-Bourque

Lycodes vahlii

1. Colour photograph from Nozères et al. (2010) 2. After Jensen (1904) 3. Ventral head, after Goode and Bean (1896)

Lycodonus mirabilis



After Goode and Bean (1896)

Melanostigma atlanticum



After Goode and Bean (1896)

Zoarces americanus

1. Aquarium specimen, Wikimedia Commons; colour photograph by S.G. Johnson, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0 2. CMNFI 1979-1218, 190.6 mm TL, Newfoundland, St Margaret Bay at South Cove; drawing by Charles H. Douglas

Acantholumpenus mackayi

1. CAS 235302, 182 mm TL, eastern Bering Sea, 2011; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1965-0341C, 269 mm SL, NWT, Tuktoyaktuk, Kugmallit Bay; drawing by Charles H. Douglas

Anisarchus medius



CMNFI 1970-0279, male, 110 mm TL, Nunavut, Frobisher Bay, 8 km south-southeast of Frobisher Bay; drawing by Charles H. Douglas

Chirolophis ascanii



Composite drawing by Susan Laurie-Bourque

Eumesogrammus praecisus

1. Colour photograph from Nozères et al. (2010). 2. CMNFI 1960-0069, female, 126 mm TL, Nunavut, Cape Pembroke, Coats Island; drawing by John L. Tottenham

Leptoclinus maculatus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1959-0389, female, 130 mm SL, Québec, Ungava Bay, Port Burwell; drawing by John L. Tottenham

Lumpenus fabricii

1. IMS 23, 136 mm TL, western Chukchi Sea, 2004; colour photograph by Boris A. Sheiko, Russian Academy of Sciences, and Catherine W. Mecklenburg, CAS 2. Composite drawing by Susan Laurie-Bourque

Lumpenus lampretaeformis

1. Colour photograph from Nozères et al. (2010) 2. Composite drawing by Susan Laurie-Bourque

Stichaeus punctatus

1. Juvenile, 97 mm; colour photograph from National Oceanic and Atmospheric Administration (NOAA Fisheries, Alaska Regional Office), https://alaskafisheries.noaa.gov/habitat/ nearshore-fish-photos 2. CMNFI 1960-0186, male, 86 mm SL, Nunavut, Hudson Bay, southern Flaherty Island, Belcher Islands; drawing by John L. Tottenham

Ulvaria subbifurcata



After Collett (1880)

U.S. National Museum, Washington, DC



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I l l u st r at i on C r e di ts 601

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Species

Source

Pholis fasciata

1. CAS 220466, 159 mm TL, eastern Bering Strait, 2004; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1960-0098, male, 227 mm TL, Nunavut, Coral Harbour, southern Southampton Island; drawing by John L. Tottenham

Anarhichas denticulatus

1. CMNFI 1962-0153, 500 mm SL, Newfoundland, Grand Banks; drawing by John L. Tottenham 2. Teeth on central vomer not extending back beyond lateral palatines; drawing by Susan Laurie-Bourque

Anarhichas lupus

1. After Coad, with Waszczuk and Labignan (1995) 2. No locality; drawing by John L. Tottenham 3. Teeth on central vomer extending back beyond lateral palatines; drawing by Susan Laurie-Bourque

Anarhichas minor

1. Scott Inlet, Baffin Bay (71.5166° N, 70.2749° W) at 495 m, CCGS Amundsen; colour photographs by Bárbara de Moura Neves of DFO and ROPOS 2. CMNFI 1962-0155, 516 mm SL, Newfoundland, Grand Banks; drawing by John L. Tottenham 3. Teeth on central vomer not extending back beyond lateral palatines; drawing by Susan Laurie-Bourque

Anarhichas orientalis

1. CMNFI 1971-0108, body and frontal view of head, 454 mm TL, Nunavut, Kater Point, Bathurst Inlet; drawing by Charles H. Douglas 2. Teeth on central vomer extending back beyond lateral palatines; drawing by Susan Laurie-Bourque

Chiasmodon harteli



Body and dorsal view, after C. niger from Goode and Bean (1896)

Ammodytes dubius



CMNFI 1962-0566, 82 mm SL, Québec, Great Whale River mouth; drawing by John L. Tottenham

Ammodytes hexapterus

1. Adult; colour photograph from National Oceanic and Atmospheric Administration (NOAA Fisheries, Alaska Regional Office), https://alaskafisheries.noaa.gov/habitat/nearshore-fish-photos 2. CMNFI 1960-0470, 72 mm SL, Yukon, Thetis Bay at Simpson Point, Herschel Island; with pigment and scalation from CMNFI 1963-0227, Québec, Richmond Gulf, Clearwater River lagoon; drawings by John L. Tottenham 3. Adult in sand; colour photograph from National Oceanic and Atmospheric Administration (NOAA Fisheries, Alaska Regional Office), https://alaskafisheries.noaa.gov/habitat/nearshore-fish-photos

Aphanopus carbo



Peprilus triacanthus

1. Colour photograph from Nozères et al. (2010) 2. CMNFI 1963-0017, 235 mm TL, Nova Scotia, Georges Bank southeast of Cape Sable; drawing by Charles H. Douglas

Glyptocephalus cynoglossus



Hippoglossoides platessoides

1. CMNFI 1962-0553, 255 mm SL, Québec, Richmond Gulf, east central Hudson Bay; drawing by John L. Tottenham 2. Head; Wikimedia Commons, colour photograph by A-P. D. Picard, under the Creative Commons licence, https://creativecommons.org/licenses/by-sa/3.0

Hippoglossoides robustus

1. CAS 220440, eyed and blind sides, 185 mm TL, western Chukchi Sea, 2004; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1966-0076, Nunavut, Buchan Bay, Bathurst Inlet; drawing by John L. Tottenham

Hippoglossus hippoglossus



Limanda proboscidea

1. ZIN 54866, eyed and blind sides, 140 mm TL, eastern Chukchi Sea, 2009; colour photograph by Catherine W. Mecklenburg, CAS 2. CMNFI 1976-0119, 286 mm TL, Nunavut, northern Parry Bay off Nauyuk River; drawing by Charles H. Douglas

Platichthys stellatus

1. Ca. 250 mm, Phillips Bay, Yukon, July 2008; colour photograph by Colin Gallagher, DFO 2. Left-eyed (reversed below image) 3. CMNFI, no locality; drawing by John L. Tottenham

Pleuronectes glacialis

1. Ca. 250 mm, Phillips Bay, Yukon, July 2008; colour photograph by Colin Gallagher, DFO 2. CMNFI 1962-0353, 222 mm TL, NWT, Liverpool Bay; drawing by John L. Tottenham

Pleuronectes putnami



CMNFI 1982-0020, 245 mm TL, New Brunswick, Miramichi River northeast of Newcastle; drawing by Charles H. Douglas

Reinhardtius hippoglossoides



CMNFI 1963-0133, 430 mm TL, Newfoundland, ca. 80 km off southeastern tip of Labrador; drawing by John L. Tottenham

602

After Günther (1887)

CMNFI 1984-0267, 565 mm TL, Nova Scotia, west of Cape Breton; drawing by Charles H. Douglas

Composite drawing by Susan Laurie-Bourque

Illustration Credits

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INDEX

Page numbers in bold refer to main species account.

Scientific Names abbreviata, Chimaera, 171 abyssorum, Nematonurus, 333 acadianus, Glyptocephalus, 549 acanthias, Squalus, 81, 175 Acanthocottus labradoricus, 422, 426 Acanthocottus patris, 413 acanthognathus, Lophodolos, 82 Acantholumpenus mackayi, ix, 4, 79, 146, 515–516 Accipenser, 80, 203 Acipenser fulvescens, 3, 4, 75, 93, 203–205 Acipenser oxyrinchus, 80, 85, 203 Acipenser rupertianus, 203 aculeatus, Argyropelecus, 81 aculeatus, Clinus, 520 aculeatus, Gasterosteus, 4, 77, 101, 119, 397–399 adolfi, Lycodes, 78, 154, 484–485, 488 aegelfinus, Melanogrammus, 82 aenaeus, Myoxocephalus, 78, 114, 419–420 aeneus, Myoxocephalus, 419 aequalis, Nezumia, 81 aequoreus, Entelurus, 82 affinis, Chimaera, 171 affinis, Halargyreus, 348 affinis, Hydrolagus, 4, 75, 96, 171–173 agassizii, Alepocephalus, 76, 105, 109, 234 agilis, Gadus, 360 agnostus, Lycodes, 500 Agonus, 438 agulhensis, Lycodes, 508 alaskense, Lethenteron, 165

alba, Lycenchelys, 83 albus, Merlangus, 375 Aldrovandia phalacra, 81 Alepisaurus brevirostris, 81 Alepisaurus ferox, 81 Alepocephalus agassizii, 76, 105, 109, 234 Alepocephalus bairdii, 76, 109, 235 algibarbata, Linophryne, 82 alipes, Salmo, 283 Allocyttus verrucosus, 82 alosoides, Hiodon, 85 alpinus, Salmo, 283 alpinus, Salvelinus, 3, 4, 5, 19, 67, 76, 141, 142, 283–289, 290, 292, 297 alpinus, Salvelinus alpinus, 283 alpinus alpinus, Salvelinus, 283 alpinus erythrinus, Salvelinus, 283 alpinus oquassa, Salvelinus, 283 alpinus salvelinus, Salvelinus, 283 alpinus taranetzi, Salvelinus, 283 Amblyraja, 185, 186, 189, 197 Amblyraja hyperborea, 4, 75, 92, 137, 181, 189, 186–188, 190, 192, 201, 568 Amblyraja jenseni, 4, 75, 137, 186, 189–190, 201 Amblyraja radiata, 4, 75, 136, 181, 186, 188, 190–192 americana, Morrhua, 367 americana, Raia, 191 americanus, Ammodytes, 541 americanus, Hippoglossus, 556 americanus, Lophius, 4, 77, 94, 378–379 americanus, Lumpenus lampetraeformis, 523 americanus, Pseudopleuronectes, 83, 282 americanus, Zoarces, 79, 149, 473, 513–514 Ammocoetes aureus, 165



UTP Fishes Book 5pp04.indb 603

Ammodytes, 286, 553 Ammodytes americanus, 541 Ammodytes dubius, 79, 103, 109, 541–543 Ammodytes dubius hudsonius, 541 Ammodytes hexapterus, 5, 79, 109, 541, 542, 543–545 Ammodytes hudsonius, 541 Ammodytes marinus, 83, 541 Ammodytes sp., 291 ampullaceus, Saccopharynx, 75, 95, 216, 348 Anarhichas denticulatus, 4, 79, 110, 530–532 Anarhichas latifrons, 530 Anarhichas lupus, 4, 79, 110, 530, 532–534 Anarhichas minor, 4, 79, 110, 530, 534–537 Anarhichas orientalis, ix, 4, 79, 103, 110, 531, 537–538 andersoni, Gymnelus, 476 andriashevi, Eumicrotremus, 82 Anguilla rostrata, 81 anisacanthus, Oneirodes, 383 Anisarchus medius, 79, 147, 517–518 anomala, Holtbyrnia, 3, 4, 75, 131, 228 Anoplogaster cornuta, 77, 102, 392–394 Anotopterus arcticus, 317 Anotopterus pharao, 76, 129, 317–318 antarcticus, Borostomias, 76, 102, 148, 309 antarcticus, Somniosus, 184 Antimora, 346 Antimora microlepis, 347 Antimora rostrata, 22, 23, 77, 125, 210, 347–348 Aphanopus carbo, 79, 99, 348, 546 Aphanopus minor, 546 Aplodinotus grunniens, 85

Apristurus laurussonii, 81, 174 Apristurus manis, 174 Apristurus profundorum, 3, 4, 75, 93, 174 apus, Platytroctes, xii, 3, 76, 131, 232–233 arctica, Liparis, 464 arcticum, Protomyctophum, 76, 126, 331 arcticus, Anotopterus, 317 arcticus, Thymallus, 85 arcticus, Trachipterus, 81 Arctogadus, 358 Arctogadus borisovi, 358, 359 Arctogadus glacialis, 3, 18, 77, 118, 358–360, 568 Arctozenus risso, 76, 104, 129, 318–319 Arctozenus risso kroyeri, 318, 319 Arctozenus risso risso, 318, 319 arcturus, Salmo, 283 arenosus, Gadus, 367 argentatus, Gaidropsarus, 77, 131, 351, 353–354 Argentina silus, 75, 104, 224–225 Argyropelecus aculeatus, 81 Argyropelecus gigas, 3, 76, 101, 160, 306–307 Argyropelecus hemigymnus, 81 Argyropelecus olfersi, 81 Argyrosomus, 251, 262 armatus, Coryphaenoides, 77, 125, 333–335 armatus, Coryphaenoides armatus, 333 armatus armatus, Coryphaenoides, 333 armatus variabilis, Coryphaenoides, 333 artedi, Coregonus, 4, 5, 57, 76, 141, 251–253, 254, 256, 264 Artediellus, ix

I ndex 603

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Artediellus atlanticus, 23, 77, 113, 408–410 Artediellus gomojunovi, 82 Artediellus scaber, 77, 112, 410–411 Artediellus uncinatus, 77, 111, 113, 412–413 ascanii, Chirolophis, 79, 145, 518–519 asiatica, Lota lota, 373 asper, Macrurus, 333 aspera, Limanda, 83 Aspidophoroides borealis, 436 Aspidophoroides groenlandicus, 436 Aspidophoroides monopterygius, 78, 108, 435–436 Aspidophoroides olrikii, 5, 78, 108, 436–438 asteroides, Polyipnus, 81 Astronesthes, 308 Astronesthes cf. richardsoni, 76, 148, 308 athabascae, Coregonus, 251 Atheresthes stomias, 8 atherinoides, Notropis, 85 atikameg, Coregonus, 257 atikameg manitobensis, Coregonus, 257 atlantica, Magnisudis, 76, 130, 320 atlantica, Myxine, 163 atlantica, Rhinochimaera, 3, 4, 75, 96, 138, 169–170 atlanticum, Melanostigma, 79, 149, 473, 512 atlanticus, Artediellus, 23, 77, 113, 408–410 atlanticus, Hoplostethus, 4, 77, 100, 395–396 atlanticus, Liparis, 78, 121, 458–459 atlanticus, Lycodes, 508 atlanticus, Lycodes turneri, 500 atratulus, Rhinichthys, 85 atratus, Lycodes, 508 attrita, Rouleina, 81 aureus, Ammocoetes, 165 australis, Micromesistius, 358 autumnalis, Coregonus, 4, 21, 76, 141, 254–256 autumnalis migratorius, Coregonus, 254 Avocettina infans, 81 bairdii, Alepocephalus, 76, 109, 235 bairdii, Cottus, 85 bairdii, Gastromus, 217 bairdii, Nezumia, 77, 124, 344–345 Bajacalifornia megalops, 76, 109, 236 balushkini, Phyllorhinichthys, 82 barbatus, Mullus, 429 Barbourisia rufa, 82 barracudina, Paralepis coregonoides, 321 barsukovi, Gymnelus, 8, 78, 150, 473–474, 478

604

bartonbeani, Melanostomias, 81 bathyarcticus, Liparis, 8, 82 bathyarcticus, Liparis liparis, 462 bathybii, Paraliparis, 466 bathybius, Histiobranchus, 81 bathybius, Paraliparis, 78, 122, 466–467 Bathylaco nigricans, 81 Bathylagus euryops, 22, 75, 104, 226–227 bathyphila, Raja, 197 bathyphila, Rajella, 3, 4, 75, 138, 197–199, 201 bathyphilus, Sigmops, 76, 120, 304–305 Bathyraja, 135, 185, 193, 194 Bathyraja sp., 75, 193–194 Bathyraja spinicauda, 4, 75, 92, 135, 193, 194–195 Bathysaurus ferox, 81 Bathytroctes drakei, 236 Bathytroctes michaelsarsi, 237 Bathytroctes sp., 3, 76, 109, 237 beani, Scopelogadus, 82 beanii, Serrivomer, 75, 99, 214–215 Benthosema glaciale, 22, 23, 76, 103, 127, 322, 323–324 Benthosema glacialis, 323 berglax, Macrourus, 4, 5, 22, 77, 124, 210, 342–343 bericoides, Melanolagus, 81 bicoloratus, Kareius, 561 bicornis, Icelus, 78, 112, 415–417 bicornis, Linophryne, 82 bigelowi, Rajella, 81, 185, 197 bilabrus, Gymnelus, 8, 78, 150, 474–475, 478 blennoides, Phycis, 351 boa boa, Stomias, 313 boa ferox, Stomias, 76, 313 boa, Stomias, 76, 147, 313–315 boa, Stomias boa, 313 bolangeri, Chiasmodon, 539 borea, Raja, 186 borealis, Aspidophoroides, 436 borealis, Paralepis coregonoides, 76, 321 borealis, Petromyzon, 165 Boreogadus, 358, 375 Boreogadus pearyi, 359 Boreogadus saida, 3, 5, 20, 22, 23, 32, 48, 77, 118, 358, 360–363, 409, 541, 553, 568, 569 borisovi, Arctogadus, 358, 359 Borostomias antarcticus, 76, 102, 148, 309 Bottemannei, Gadus (Boreogadus), 359 brasiliensis, Urophycis, 351 braueri, Cyclothone, 81 brevibarbis, Coryphaenoides, 77, 125, 335–336

brevipes, Halargyreus, 348 brevipinna, Somniosus, 180 Breviraja, 197 Breviraja marklei, 199 brevirostris, Alepisaurus, 81 brosme, Brosme, 4, 77, 117, 364–365 Brosme brosme, 4, 77, 117, 364–365 brosmiana, Lota, 373 Brosmius flavescens, 364 Brosmus vulgaris, 364 brunneus, Lycodes, 508 Bythites fuscus, 3, 77, 101, 377 Caelorinchus, 333 callarias, Gadus, 367 callarias, Gadus morhua, 367 callarias marisalbi, Gadus, 371 camtschaticum, Lethenteron, 4, 75, 92, 165–167 canadensis, Sander, 85 capito, Poromitra, 82 caprodes, Percina, 85 carapinus, Coryphaenoides, 77, 106, 125, 336–337 carbo, Aphanopus, 79, 99, 348, 546 Careproctus cf. rastrinus, 82 Careproctus kidoi, xiii, 4, 78, 121, 454–455, 458 Careproctus longipinnis, 78, 120, 455–456 Careproctus reinhardti, 78, 121, 454, 455, 456–458 Caristius fasciatus, 4, 78, 99, 471–472 Caristius groenlandicus, 471 carpio, Cyprinus, 85 cataractae, Rhinichthys, 85 catervarius, Mallotus, 8 catervarius, Mallotus villosus, 240 catervarius natio schulzi, Mallotus villosus, 240 catostomus, Catostomus, 85 Catostomus catostomus, 85 Catostomus commersonii, 85 caudacuta, Motella, 351 Caulophryne jordani, 82 Centronotus gunelliformis, 528 Centrophorus coelolepis, 178 Centroscyllium fabricii, 4, 23, 75, 92, 93, 175–177 Centroscymnus, 178 Centroscymnus coelolepis, 3, 4, 75, 144, 178–180 Ceratias holboelli, 4, 77, 94, 386–387 Ceratoscopelus maderensis, 80, 81 Cetorhinus maximus, 81 Chaenophryne draco, 82 Chaenophryne longiceps, xiii, 4, 77, 95, 128, 382–383 chalcogrammus, Gadus, 82 Chalinura, 335, 336, 337 Chauliodus, 307

Chauliodus sloanei, 310 Chauliodus sloani, 76, 147, 310–311 chemnitzii, Notacanthus, 75, 100, 128, 206–207 chesteri, Phycis, 77, 106, 130, 356–357 Chiasmodon bolangeri, 539 Chiasmodon harteli, 4, 79, 96, 539–540 Chiasmodon niger, 4, 539, 540 Chimaera, 171 Chimaera abbreviata, 171 Chimaera affinis, 171 Chimaera monstrosa, 171 Chimaera plumbea, 171 Chirolophis ascanii, 79, 145, 518–519 Chirolophis galerita, 518 Chrosomus eos, 85 Chrosomus neogaeus, 85 churchillensis, Leucichthys, 251 chuss, Urophycis, 351 Ciliata septentrionalis, 81 cimbrius, Enchelyopus, 3–4, 77, 130, 351–353 clavata, Raja, 191 Clinus aculeatus, 520 Clinus unimaculatus, 519 Clupea harengus, 4, 75, 111, 188, 219–221, 223, 281, 553 Clupea pallasii, 58, 75, 105, 111, 221–223, 299 clupeaformis, Coregonus, 5, 19, 76, 140, 257–259, 260 coccineus, Lycodes, 496 coelolepis, Centrophorus, 178 coelolepis, Centroscymnus, 3, 4, 75, 144, 178–180 Coelorinchus, 333 cognatus, Cottus, 85 coheni, Liparis, 82 colliei, Hydrolagus, 172 commersonii, Catostomus, 85 compressus, Gadus, 373 confluentus, Salvelinus, 292 copei, Paraliparis, 4, 78, 123, 467–468 copei, Xenodermichthys, 76, 109, 239 coregonoides, Paralepis, 76, 130, 321 coregonoides, Paralepis coregonoides, 321 coregonoides barracudina, Paralepis, 321 coregonoides borealis, Paralepis, 76, 321 coregonoides coregonoides, Paralepis, 321 Coregonus, 67, 248, 251, 254, 277, 321 Coregonus artedi, 4, 5, 57, 76, 141, 251–253, 254, 256, 264 Coregonus athabascae, 251 Coregonus atikameg, 257 Coregonus atikameg manitobensis, 257

Index

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Coregonus autumnalis, 4, 21, 76, 141, 254–256 Coregonus autumnalis migratorius, 254 Coregonus clupeaformis, 5, 19, 76, 140, 257–259, 260 Coregonus entomophagus, 251 Coregonus huntsmani, 259 Coregonus laurettae, 4, 81, 141, 254, 256 Coregonus lavaretus, 254, 260 Coregonus macrognathus, 251 Coregonus nasus, 4, 58, 76, 140, 250, 257, 260–262 Coregonus nelsonii, 257 Coregonus pidschian, 81, 257 Coregonus pollan, 254 Coregonus preblei, 277 Coregonus pusillus, 262 Coregonus quadrilateralis, 277 Coregonus sardinella, 4, 21, 76, 140, 253, 254, 262–264 Coregonus subautumnalis, 254 Coregonus zenithicus, 251 cornuta, Anoplogaster, 77, 102, 392–394 coronata, Linophryne, 82 corporalis, Semotilus, 85 Coryphaena, 338 Coryphaenoides, 333, 335, 337, 338, 342, 344 Coryphaenoides armatus, 77, 125, 333–335 Coryphaenoides armatus armatus, 333 Coryphaenoides armatus variabilis, 333 Coryphaenoides brevibarbis, 77, 125, 335–336 Coryphaenoides carapinus, 77, 106, 125, 336–337 Coryphaenoides gigas, 333 Coryphaenoides guentheri, 77, 124, 337–338 Coryphaenoides longifilis, 81 Coryphaenoides rupestris, 4, 77, 124, 210–211, 333, 338–340, 342 Coryphaenoides variabilis, 333 corythaeola, Polymetme, 81 Cottunculus microps, 22, 23, 78, 135, 440–442 Cottunculus sadko, 440, 442 Cottunculus sp., 492 Cottunculus thomsonii, 78, 107, 135, 442–443 Cottus bairdii, 85 Cottus cognatus, 85 Cottus glacialis, 426 Cottus gobio, 440 Cottus hexacornis, 422 Cottus polaris, 422 Cottus porosus, 426

Cottus ricei, 85 Couchia edwardii, 351 couesii, Cryptopsaras, 82 Couesius plumbeus, 85 crassiceps, Poromitra, 82 Cristivomer, 297 crocodilus, Lampanyctus, 76, 127, 324–325 Cryptacanthodes giganteus, 80 Cryptacanthodes maculatus, 80 Cryptopsaras couesii, 82 Culaea inconstans, 85 cyclolepis, Nematonurus, 333 Cyclopteropsis jordani, 78, 117, 446–447 Cyclopteropsis macalpini, 447 Cyclopteropsis mcalpini, 4, 78, 117, 447–448 Cyclopterus lumpus, 78, 97, 116, 446, 449–450 Cyclopterus lumpus hudsonius, viii Cyclopterus lumpus var. hudsonius, 449 cyclostigma, Liparis, 462 Cyclothone braueri, 81 Cyclothone microdon, 76, 103, 120, 302–303 cylindraceum, Prosopium, 5, 76, 139, 277–279 Cylindraceus, Salmo, 277 cynoglossus, Glyptocephalus, 79, 132, 549–551, 568 Cyprinus carpio, 85 Dactylobatus, 197 dactylopterus, Helicolenus, 82 Danaphryne nigrifilis, 82 decagonus, Leptagonus, 78, 108, 438–439 decimus, Rhadinesthes, 76, 148, 312–313 Delolepis gigantea, 80 dentex, Osmerus, 3, 4, 76, 105, 129, 244–245, 246 dentex, Osmerus mordax, 244 denticulatus, Anarhichas, 4, 79, 110, 530–532 derjugini, Eumicrotremus, 5, 78, 117, 450–452 diceraus, Enophrys, 82 Dipturus, 201 Dipturus laevis, 189, 191 Diretmoides pauciradiatus, 82 Dissostichus eleginoides, 80, 83 Dolopichthys longicornis, 82 draco, Chaenophryne, 82 drakei, Bathytroctes, 236 dubius, Ammodytes, 79, 103, 109, 541–543 dubius hudsonius, Ammodytes, 541 dwinensis, Platessa, 564 dypterygia, Molva, 82



UTP Fishes Book 5pp04.indb 605

edentula, Einara, 81 edwardii, Couchia, 351 eggvinii, Eumicrotremus, 452 Einara edentula, 81 elassodon, Hippoglossoides, 8, 555 eleginoides, Dissostichus, 80, 83 Eleginus gracilis, 21, 77, 119, 299, 358, 365–367 Eleginus navaga, 365 Eleginus nawaga, 365 elongatus, Notoscopelus, 329 Enchelyopus, 351, 373 Enchelyopus cimbrius, 3–4, 77, 130, 351–353 Enophrys diceraus, 82 ensis, Gaidropsarus, 77, 131, 351, 355–356, 460 Entelurus aequoreus, 82 entomophagus, Coregonus, 251 Entosphenus, 165 Entosphenus tridentatus, 165 eos, Chrosomus, 85 eques, Lepidion, 4, 77, 106, 126, 349–350 erythrinus, Salvelinus alpinus, 283 eschrichtii, Oneirodes, 82, 383 esipovi, Gymnelus, 478 esmarkii, Lycodes, 78, 155, 486–487, 508 esmarkii, Trisopterus, 82 Esox lucius, 85 Etheostoma nigrum, 85 Etmopterus princeps, 81, 175 Euchalarodus, 566 eudipleurostictus, Lycodes, 78, 155, 486, 487–488, 508 eudipleurostictus, Lycodes cf., 48 Eumesogrammus praecisus, 79, 145, 519–520 Eumicrotremus andriashevi, 82 Eumicrotremus derjugini, 5, 78, 117, 450–452 Eumicrotremus eggvinii, 452 Eumicrotremus spinosus, 78, 117, 452–453 Eumicrotremus spinosus variabilis, 450, 452 euryops, Bathylagus, 22, 75, 104, 226–227 Eurypharynx pelecanoides, 75, 95, 217–218 fabricii, Centroscyllium, 4, 23, 75, 92, 93, 175–177 fabricii, Gadus, 360 fabricii, Liparis, 5, 78, 120, 121, 409, 460–462 fabricii, Lumpenus, 79, 147, 515, 522–523 fabricii, Macrourus, 342 fabricii, Spinax, 175 farrani, Nematonurus, 336

fasciata, Pholis, 79, 107, 528–529 fasciatus, Caristius, 4, 78, 99, 471–472 fasciatus, Sebastes, 4, 5, 77, 144, 402–404 ferox, Alepisaurus, 81 ferox, Bathysaurus, 81 ferox, Stomias boa, 76, 313 flagellicauda, Lycodonus, 510, 511 flavescens, Brosmius, 364 flavescens, Perca, 85 fluvialis, Petromyzon, 165 fontinalis, Salvelinus, 29, 67, 76, 142, 284, 285, 289–292 fortidens, Lycichthys, 530 franklinii, Pleuronectes, 564 frigidus, Lycodes, ix, 3, 78, 154, 485, 488–489 fulvescens, Acipenser, 3, 4, 75, 93, 203–205 fuscus, Bythites, 3, 77, 101, 377 fyllae, Rajella, 4, 75, 136, 199–201 Gadiculus, 358 Gadomus longifilis, 3, 4, 77, 123, 340–341 Gadus, ix, 286, 340, 351, 359, 360, 364, 365, 373, 375 Gadus (Boreogadus) Bottemannei, 359 Gadus agilis, 360 Gadus arenosus, 367 Gadus callarias, 367 Gadus callarias marisalbi, 371 Gadus chalcogrammus, 82 Gadus compressus, 373 Gadus fabricii, 360 Gadus heteroglossus, 367 Gadus lacustris, 373 Gadus lubb, 364 Gadus macrocephalus, 82, 371 Gadus maculosus, 373 Gadus morhua, 4, 34, 77, 106, 119, 358, 367–371, 553 Gadus morhua callarias, 367 Gadus morhua kildinensis, 367, 368 Gadus morhua macrocephalus, 367 Gadus morhua marisalbi, 367 Gadus morhua morhua, 367 Gadus morrhua, 367 Gadus nanus, 367 Gadus ogac, 35, 74, 77, 119, 358, 371–373 Gadus rupestris, 367 Gadus torsk, 364 Gadus vertagus, 367 Gadus wachna, 365 Gaidropsarus, 351 Gaidropsarus argentatus, 77, 131, 353–354, 351 Gaidropsarus ensis, 77, 131, 351, 355–356, 460

I ndex 605

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Gaidropsarus sp., 22, 23 galerita, Chirolophis, 518 garmani, Paraliparis, 78, 123, 468–469 Gasterosteus aculeatus, 4, 77, 101, 119, 397–399 Gasterosteus insculptus, 397, 398 Gastromus bairdii, 217 gemmifer, Lampanyctus, 324 gibbus, Liparis, 8, 78, 122, 462–463 Gigantactis vanhoeffeni, 4, 77, 94, 388–389 gigantea, Delolepis, 80 giganteus, Cryptacanthodes, 80 gigas, Argyropelecus, 3, 76, 101, 160, 306–307 gigas, Coryphaenoides, 333 gigas, Hippoglossus, 556 glaciale, Benthosema, 22, 23, 76, 103, 127, 322, 323–324 glacialis, Arctogadus, 3, 18, 77, 118, 358–360, 568 glacialis, Benthosema, 323 glacialis, Cottus, 426 glacialis, Pleuronectes, 8, 79, 134, 549, 564–566 glacialis, Scymnus, 180 gladisfenae, Spiniphryne, 77, 128, 385 glutinosa, Myxine, 4, 75, 92, 162–164 Glyptocephalus acadianus, 549 Glyptocephalus cynoglossus, 79, 132, 549–551, 568 gobio, Cottus, 440 gomojunovi, Artediellus, 82 Gonostoma, 304 goodii, Macrurus, 333 gorbuscha, Oncorhynchus, 3, 76, 143, 264–267 gracilis, Eleginus, 21, 77, 119, 299, 358, 365–367 gracilis, Lycodes, 509, 510 gracilis, Lycodes vahlii, 509 gracilis, Platygobio, 85 granulata, Raja, 189 groenlandica, Nansenia, 81 groenlandicus, Aspidophoroides, 436 groenlandicus, Caristius, 471 groenlandicus, Gunnellus, 528 groenlandicus, Gymnacanthus tricuspis, 413 groenlandicus, Himantolophus, 4, 77, 95, 380–381 groenlandicus, Hippoglossus, 568 groenlandicus, Myoxocephalus scorpius, 426 groenlandicus, Psednos, 82 groenlandicus, Salmo, 240 grunniens, Aplodinotus, 85 guentheri, Coryphaenoides, 77, 124, 337–338 gunelliformis, Centronotus, 528 gunnellus, Pholis, 83

606

Gunnellus groenlandicus, 528 guttatus, Lampris, 81 Gymnacanthus, 413 Gymnacanthus tricuspis groenlandicus, 413 Gymnelus, 149, 473 Gymnelus andersoni, 476 Gymnelus barsukovi, 8, 78, 150, 473–474, 478 Gymnelus bilabrus, 8, 78, 150, 474–475, 478 Gymnelus esipovi, 478 Gymnelus hemifasciatus, 476 Gymnelus knipowitschi, 8, 78, 150, 475–476, 478 Gymnelus platycephalus, 478 Gymnelus retrodorsalis, 78, 149, 476–478 Gymnelus taeniatus, 478 Gymnelus viride, 478 Gymnelus viridis, 8, 30, 78, 150, 474, 475, 478–479 Gymnelus viridis var. unimaculatus, 478 Gymnocanthus hudsonius, 413 Gymnocanthus tricuspis, 5, 78, 113, 413–415 Gymnocanthus tricuspis hudsonius, 413 Gyrinomimus myersi, 82 hacheyi, Lycodes reticulatus, 502 haeckeli, Harriotta, 81, 167 Halargyreus, 346 Halargyreus affinis, 348 Halargyreus brevipes, 348 Halargyreus johnsonii, 77, 126, 348–349 Haloporphyrus, 347, 349 Haloporphyrus viola, 347 Haplophryne mollis, 82 harengus, Clupea, 4, 75, 111, 188, 219–221, 223, 281, 553 harengus, Salmo, 251 Harriotta haeckeli, 81, 167 Harriotta raleighana, 3, 4, 75, 138, 168–169 harryi, Scopelosaurus, 316 harteli, Chiasmodon, 4, 79, 96, 539–540 harteli, Psednos, 82 hearnei, Prosopium, 277 Hearnii, Salmo, 292 Helicolenus dactylopterus, 82 hemifasciatus, Gymnelus, 476 hemigymnus, Argyropelecus, 81 herschelinus, Liparis, 464 heteroglossus, Gadus, 367 hexacornis, Cottus, 422 hexacornis, Myoxocephalus quadricornis, ix, 422 Hexagrammos stelleri, 82

hexapterus, Ammodytes, 5, 79, 109, 541, 542, 543–545 Himantolophus groenlandicus, 4, 77, 95, 380–381 Himantolophus reinhardti, 380 Hiodon alosoides, 85 Hiodon tergisus, 85 hippoglossoides, Reinhardtius, 3, 5, 12, 14, 22, 23, 79, 133, 160, 181, 182, 568–571 Hippoglossoides elassodon, 8, 555 Hippoglossoides platessoides, 79, 134, 422, 551–554 Hippoglossoides platessoides limandoides, 551, 554 Hippoglossoides platessoides platessoides, 551, 554 Hippoglossoides robustus, 8, 79, 134, 555–556 hippoglossus, Hippoglossus, 4, 5, 79, 133, 556–559 Hippoglossus americanus, 556 Hippoglossus gigas, 556 Hippoglossus groenlandicus, 568 Hippoglossus hippoglossus, 4, 5, 79, 133, 556–559 Hippoglossus stenolepis, 556 Hippoglossus vulgaris, 556 Histiobranchus bathybius, 81 holboelli, Ceratias, 4, 77, 94, 386–387 Holtbyrnia anomala, 3, 4, 75, 131, 228 Holtbyrnia macrops, 81 Hoplostethus atlanticus, 4, 77, 100, 395–396 hudsonicus, Salmo, 289 hudsonius, Ammodytes, 541 hudsonius, Ammodytes dubius, 541 hudsonius, Cyclopterus lumpus, viii hudsonius, Cyclopterus lumpus var., 449 hudsonius, Gymnocanthus, 413 hudsonius, Gymnocanthus tricuspis, 413 hudsonius, Notropis, 85 huntia, Molva, 373 huntsmani, Coregonus, 259 Hydrolagus, 171 Hydrolagus affinis, 4, 75, 96, 171–173 Hydrolagus colliei, 172 Hydrolagus pallidus, 81, 171 hyperborea, Amblyraja, 4, 75, 92, 137, 181, 186–188, 189, 190, 192, 201, 568 hyperborea, Raja, 186 Hypomesus olidus, 85 hystrix, Paraliparis, 82 Icelus, ix Icelus bicornis, 78, 112, 415–417 Icelus spatula, 78, 112, 417–418

Ichthyomyzon unicuspis, 85 inconstans, Culaea, 85 infans, Avocettina, 81 ingolfi, Macrurus, 337 ingolfiana, Raja, 201 ingolfianus, Lycenchelys, 482 inornata, Lota, 373 insculptus, Gasterosteus, 397, 398 intricarius, Lampanyctus, 76, 127, 326 ischyrus, Parophrys, 561 islandica, Normichthys operosa, 231 jaok, Myoxocephalus, 82 japonicus, Petromyzon, 165 jenseni, Amblyraja, 4, 75, 137, 185, 186, 189–190, 201 jespersenae, Myxine, 81, 164 johnsonii, Halargyreus, 77, 126, 348–349 johnsonii, Melanocetus, 82 jordani, Caulophryne, 82 jordani, Cyclopteropsis, 78, 117, 446–447 jugoricus, Lycodes, 4, 78, 152, 489–490 kamensis, Lota lota, 373 Kareius bicoloratus, 561 kaupii, Synaphobranchus, 4, 75, 98, 148, 210–211 kennerlyi, Oncorhynchus, 272 kennerlyi, Salmo, 272 keta, Oncorhynchus, 3, 76, 143, 267–270 kidoi, Careproctus, xiii, 4, 78, 121, 454–455, 458 kildinensis, Gadus morhua, 367, 368 kisutch, Oncorhynchus, 76, 143, 270–272 knipowitschi, Gymnelus, 8, 78, 150, 475–476, 478 knipowitschi, Lycodes, 496 knipowitschi panthera, Lycodes, 496 koefoedi, Liparis, 460 koefoedi, Searsia, 81 kolthoffi, Lycenchelys, 78, 151, 480 kroyeri, Arctozenus risso, 318, 319 kroyeri, Notoscopelus, 76, 126, 329–330 kroyeri, Paralepis, 318 labradorensis, Lycenchelys, 83 labradoricus, Acanthocottus, 422, 426 lacustris, Gadus, 373 lacustris, Lota lota, 373 Laemonema, 346 laevis, Dipturus, 189, 191 laevis, Raja, 191 Lamna nasus, 81

Index

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Lampadena speculigera, 81 Lampanyctus crocodilus, 76, 127, 324–325 Lampanyctus gemmifer, 324 Lampanyctus intricarius, 76, 127, 326 Lampanyctus macdonaldi, 22, 23, 76, 127, 327 Lampetra, 165 lampetraeformis americanus, Lumpenus, 523 lampetraeformis terraenovae, Lumpenus, 523 lampretaeformis, Lumpenus, 79, 147, 523–524 lampretaeformis serpentinus, Lumpenus, 523 lampretiformis, Lumpenus, 523 Lampris guttatus, 81 latifrons, Anarhichas, 530 laurentianus, Lycodes reticulatus, 502 laurettae, Coregonus, 4, 81, 141, 254, 256 laurussonii, Apristurus, 81, 174 lavalaei, Lycodes, 78, 152, 491–492, 508 lavalei, Lycodes, 481 lavaretus, Coregonus, 254, 260 Lepidion eques, 4, 77, 106, 126, 349–350 Lepidoleprus norvegicus, 338 lepidus, Scopelosaurus, 76, 104, 316 Leptagonus decagonus, 78, 108, 438–439 Leptoclinus maculatus, 79, 145, 520–521 leptura, Lota lota, 373 lerikimae, Liparis, 8 Lethenteron alaskense, 165 Lethenteron camtschaticum, 4, 75, 92, 165–167 Leucichthys, 248, 251, 254, 262 leucichthys, Salmo, 300 leucichthys, Stenodus, 4, 30, 76, 139, 248, 300–301 Leucichthys churchillensis, 251 Leucichthys nueltinensis, 251 Leucoraja, 197, 201 Limanda, 549, 559 limanda, Limanda, 559 Limanda aspera, 83 Limanda limanda, 559 Limanda proboscidea, 79, 134, 559–560 limandoides, Hippoglossoides platessoides, 551, 554 limosa, Myxine, 162, 163 Linophryne algibarbata, 82 Linophryne bicornis, 82 Linophryne coronata, 82 Linophryne lucifer, 82 lintea, Raja, 197

lintea, Rajella, 4, 75, 138, 197, 201–202 Lionurus, 336, 337, 344 Liopsetta, 8, 549, 564, 566 Liparis arctica, 464 Liparis atlanticus, 78, 121, 458–459 Liparis bathyarcticus, 8, 82 liparis bathyarcticus, Liparis, 462 Liparis coheni, 82 Liparis cyclostigma, 462 Liparis fabricii, 5, 78, 120, 121, 409, 460–462 Liparis gibbus, 8, 78, 122, 462–463 Liparis herschelinus, 464 Liparis koefoedi, 460 Liparis lerikimae, 8 Liparis liparis bathyarcticus, 462 Liparis major, 460 Liparis marmoratus, 82 Liparis rutteri, 80 Liparis sp., 428, 434 Liparis tunicatus, 5, 78, 122, 464–465 longiceps, Chaenophryne, xiii, 4, 77, 95, 128, 382–383 longicornis, Dolopichthys, 82 longifilis, Coryphaenoides, 81 longifilis, Gadomus, 3, 4, 77, 123, 340–341 longipinnis, Careproctus, 78, 120, 455–456 longirostris, Lumpenella, 83 Lophius americanus, 4, 77, 94, 378–379 Lophius piscatorius, 82, 378 Lophodolos acanthognathus, 82 lordi, Salvelinus malma, 292 loricata, Rondeletia, 82 lota, Lota, 4, 5, 19, 77, 118, 358, 373–374 Lota brosmiana, 373 Lota inornata, 373 Lota lota, 4, 5, 19, 77, 118, 358, 373–374 Lota lota asiatica, 373 Lota lota kamensis, 373 Lota lota lacustris, 373 Lota lota leptura, 373 Lota lota lota, 373 Lota lota maculosa, 373 Lota lota obensis, 373 Lota lota onegensis, 373 Lota maculosa, 373 lubb, Gadus, 364 lucifer, Linophryne, 82 lucius, Esox, 85 luetkenii, Lycodes, 78, 152, 492–493 Lumpenella longirostris, 83 Lumpenus, 515, 517, 520 Lumpenus fabricii, 79, 147, 515, 522–523 Lumpenus lampetraeformis americanus, 523



UTP Fishes Book 5pp04.indb 607

Lumpenus lampetraeformis terraenovae, 523 Lumpenus lampretaeformis, 79, 147, 523–524 Lumpenus lampretaeformis serpentinus, 523 Lumpenus lampretiformis, 523 Lumpenus lumpretaeformis, 523 Lumpenus nubilis, 522 lumpretaeformis, Lumpenus, 523 lumpus, Cyclopterus, 78, 97, 116, 446, 449–450 lumpus hudsonius, Cyclopterus, viii lumpus var. hudsonius, Cyclopterus, 449 lupus, Anarhichas, 4, 79, 110, 530, 532–534 Lycenchelys, 149, 150, 151, 480, 481, 482, 511 Lycenchelys alba, 83 Lycenchelys ingolfianus, 482 Lycenchelys kolthoffi, 78, 151, 480 Lycenchelys labradorensis, 83 Lycenchelys micropora, 8 Lycenchelys muraena, 78, 151, 481–482 Lycenchelys paxillus, 78, 151, 482–483, 484 Lycenchelys sarsii, 78, 151, 480, 482, 483–484 Lycenchelys verrillii, 80 Lycichthys fortidens, 530 Lycichthys paucidens, 530 Lycodes, 149, 150, 504, 510, 568 Lycodes adolfi, 78, 154, 484–485, 488 Lycodes agnostus, 500 Lycodes agulhensis, 508 Lycodes atlanticus, 508 Lycodes atratus, 508 Lycodes brunneus, 508 Lycodes cf. eudipleurostictus, 48 Lycodes coccineus, 496 Lycodes esmarkii, 78, 155, 486–487, 508 Lycodes eudipleurostictus, 78, 155, 486, 487–488, 508 Lycodes frigidus, ix, 3, 78, 154, 485, 488–489 Lycodes gracilis, 509, 510 Lycodes jugoricus, 4, 78, 152, 489–490 Lycodes knipowitschi, 496 Lycodes knipowitschi panthera, 496 Lycodes lavalaei, 78, 152, 491–492, 508 Lycodes lavalei, 491 Lycodes luetkenii, 78, 152, 492–493 Lycodes marisalbi, 79, 154, 493–494, 499, 504, 506 Lycodes mcallisteri, ix, 3, 79, 153, 495 Lycodes mucosus, 79, 152, 496–497

Lycodes nigricans, 505 Lycodes paamiuti, 4, 79, 155, 497–499, 506, 507, 508 Lycodes palearis, 83 Lycodes pallidus, 79, 155, 493, 497, 499–500, 506 Lycodes pallidus marisalbi, 499 Lycodes pallidus similis, 499 Lycodes pallidus var. squamiventer, 499 Lycodes panthera, 496 Lycodes perspicillum, 502 Lycodes polaris, 79, 152, 500–501 Lycodes raridens, 8, 83, 491 Lycodes reticulatus, 79, 101, 153, 181, 491, 492, 502–503 Lycodes reticulatus hacheyi, 502 Lycodes reticulatus laurentianus, 502 Lycodes rossi, 4, 502, 503 Lycodes sagittarius, ix, 8, 79, 154, 493, 504–505 Lycodes seminudus, 79, 153, 485, 505–506 Lycodes sp., 22, 23, 428, 492 Lycodes squamiventer, 4, 79, 155, 497, 499, 504, 506–507 Lycodes terraenovae, 79, 155, 486, 487, 488, 491, 508–509 Lycodes turneri, 83, 500 Lycodes turneri atlanticus, 500 Lycodes vachonii, 486 Lycodes vahlii, 79, 154, 508, 509–510 Lycodes vahlii gracilis, 509 Lycodes vahlii maculatus, 509 Lycodes vahlii vahlii, 509 Lycodes zoarchus, 509 Lycodonus, 149, 150 Lycodonus flagellicauda, 510, 511 Lycodonus mirabilis, 48, 79, 151, 510–511 macalpini, Cyclopteropsis, 447 macdonaldi, Lampanyctus, 22, 23, 76, 127, 327 Macdonaldia, 207 mackayi, Acantholumpenus, ix, 4, 79, 146, 515–516 Mackenzii, Salmo, 300 macrocephalus, Gadus, 82, 371 macrocephalus, Gadus morhua, 367 macrognathus, Coregonus, 251 macrolepidotum, Moxostoma, 85 macrops, Holtbyrnia, 81 macrosteus, Oneirodes, 82 Macrourus berglax, 4, 5, 22, 77, 124, 342–343 Macrourus fabricii, 342 Macrourus stroemii, 338 Macrozoarces, 513 Macruroplus, 337 Macrurus, 333, 335, 336, 337, 338, 342, 344

I ndex 607

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Macrurus asper, 333 Macrurus goodii, 333 Macrurus ingolfi, 337 Macrurus suborbitalis, 333 maculatus, Cryptacanthodes, 80 maculatus, Leptoclinus, 79, 145, 520–521 maculatus, Lycodes vahlii, 509 maculosa, Lota, 373 maculosa, Lota lota, 373 maculosus, Gadus, 373 maderensis, Ceratoscopelus, 80, 81 maderensis, Paracaristius, 82 maderensis, Rouleina, xii, 3, 76, 109, 238 Magnisudis atlantica, 76, 130, 320 major, Liparis, 460 Malacoraja, 185, 196, 197 Malacoraja spinacidermis, 4, 75, 136, 196–197 Malacosteus niger, 76, 147, 311–312 Mallotus catervarius, 8 Mallotus sp., 8 Mallotus villosus, 5, 76, 129, 240– 244, 281, 286, 291, 299, 553 Mallotus villosus catervarius, 240 Mallotus villosus catervarius natio schulzi, 240 Mallotus villosus villosus, 240 malma, Salvelinus, 21, 76, 142, 284–285, 287, 292–297 malma, Salvelinus malma, 4, 292 malma lordi, Salvelinus, 292 malma malma, Salvelinus, 4, 292 manis, Apristurus, 174 manitobensis, Coregonus atikameg, 257 Margariscus margarita, 85 margarita, Margariscus, 85 marinus, Ammodytes, 83, 541 marinus, Petromyzon, 81 marinus, Sebastes, 401, 404, 406, 407 marisalbi, Gadus callarias, 371 marisalbi, Gadus morhua, 367 marisalbi, Lycodes, 79, 154, 493–494, 499, 504, 506 marisalbi, Lycodes pallidus, 499 marklei, Breviraja, 199 marmoratus, Liparis, 82 matsuurae, Reinhardtius, 568 mauli, Maulisia, 3, 4, 75, 132, 229 Maulisia mauli, 3, 4, 75, 132, 229 Maulisia microlepis, 3, 75, 132, 230 Maurolicus muelleri, 81 maximus, Cetorhinus, 81 mcallisteri, Lycodes, ix, 3, 79, 153, 495 mcallisteri, Oidiphorus, ix mcalpini, Cyclopteropsis, 4, 78, 117, 447–448 medius, Anisarchus, 79, 147, 517–518 Megalocottus platycephalus, 82

608

megalops, Bajacalifornia, 76, 109, 236 megalops, Phocaegadus, 359 Melamphaes microps, 82 melanocephalus, Psednos, 82 Melanocetus johnsonii, 82 Melanocetus murrayi, 82 Melanogrammus aegelfinus, 82 Melanolagus bericoides, 81 Melanostigma, ix Melanostigma atlanticum, 79, 149, 473, 512 Melanostomias bartonbeani, 81 melas, Scymnodon, 178 mentella, Sebastes, 4, 22, 23, 77, 107, 144, 403, 404–406, 407, 568, 569 Merlangius merlangus, 82 Merlangus, 375 merlangus, Merlangius, 82 Merlangus albus, 375 Merlangus pertusus, 375 Merlangus polaris, 360 Merlangus vernalis, 375 michaelsarsi, Bathytroctes, 237 micractis, Phyllorhinichthys, 82 microcephalus, Somniosus, 4, 8, 22, 75, 93, 145, 180–183, 188, 192 microcephalus, Squalus, 180 microdon, Cyclothone, 76, 103, 120, 302–303 Microgadus, 360, 365 Microgadus tomcod, 80, 358 microlepis, Antimora, 347 microlepis, Maulisia, 3, 75, 132, 230 Micromesistius, 358 Micromesistius australis, 358 Micromesistius poutassou, 4, 77, 118, 375–376 micropora, Lycenchelys, 8 microps, Cottunculus, 22, 23, 78, 135, 440–442 microps, Melamphaes, 82 micropterus, Scymnus, 180 micruroides, Psednos, 82 migratorius, Coregonus autumnalis, 254 minor, Anarhichas, 4, 79, 110, 530, 534–537 minor, Aphanopus, 546 mirabilis, Lycodonus, 48, 79, 151, 510–511 miriceps, Trigonolampa, 81 mollis, Haplophryne, 82 mollis, Raja, 196 molva, Molva, 82 Molva dypterygia, 82 Molva huntia, 373 Molva molva, 82 monopterygius, Aspidophoroides, 78, 108, 435–436 monstrosa, Chimaera, 171 mordax, Osmerus, 3, 4, 21, 76, 129, 244, 245, 246–247

mordax dentex, Osmerus, 244 morhua, Gadus, 4, 34, 77, 106, 119, 358, 367–371, 553 morhua, Gadus morhua, 367 morhua callarias, Gadus, 367 morhua kildinensis, Gadus, 367, 368 morhua macrocephalus, Gadus, 367 morhua marisalbi, Gadus, 367 morhua morhua, Gadus, 367 Morhua vulgaris, 367 morrhua, Gadus, 367 Morrhua americana, 367 Motella, 353, 355 Motella caudacuta, 351 Motella pacifica, 351 Motella reinhardi, 353 Moxostoma macrolepidotum, 85 mucosus, Lycodes, 79, 152, 496–497 muelleri, Maurolicus, 81 Mullus barbatus, 429 muraena, Lycenchelys, 78, 151, 481–482 murrayi, Melanocetus, 82 murrayi, Trachyrincus, 77, 123, 342, 344, 345–346 murrayi, Triglops, 78, 116, 429–430 Myctophum, 316, 329, 390 Myctophum punctatum, 3, 76, 127, 328–329 myersi, Gyrinomimus, 82 Myoxocephalus, 286, 291, 408, 419, 420, 422, 423, 424, 426 Myoxocephalus aenaeus, 78, 114, 419–420 Myoxocephalus aeneus, 419 Myoxocephalus jaok, 82 Myoxocephalus octodecemspinosus, 78, 107, 114, 420–421 Myoxocephalus quadricornis, ix, 3, 4, 5, 78, 114, 421–424, 426 Myoxocephalus quadricornis hexacornis, ix, 422 Myoxocephalus quadricornis quadricornis, 422 Myoxocephalus scorpioides, 5, 78, 115, 158, 414, 424–426 Myoxocephalus scorpius, 5, 32, 78, 82, 115, 181, 414, 422, 426–429 Myoxocephalus scorpius groenlandicus, 426 Myoxocephalus sp., 434 Myoxocephalus thompsonii, ix Myoxocephalus verrucosus, 82 Myxine atlantica, 163 Myxine glutinosa, 4, 75, 92, 162–164 Myxine jespersenae, 81, 164 Myxine limosa, 162, 163 Myzopsetta, 559 namaycush, Salmo, 297 namaycush, Salvelinus, 3, 4, 5, 19, 76, 142, 283, 284, 285, 297–299

Nansenia groenlandica, 81 Nansenia oblita, 81 Nansenia sp., 81 nanus, Gadus, 367 Naresi, Salmo, 283 naresii, Salmo, 283 nasus, Coregonus, 4, 58, 76, 140, 250, 257, 260–262 nasus, Lamna, 81 nasus, Notocanthus, 206 Nautichthys pribilovius, 82 navaga, Eleginus, 365 nawaga, Eleginus, 365 nelma, Salmo, 300 nelma, Stenodus, 300 nelsonii, Coregonus, 257 Nematonurus, 333 Nematonurus abyssorum, 333 Nematonurus cyclolepis, 333 Nematonurus farrani, 336 Nemichthys scolopaceus, 75, 98, 212–213 neogaeus, Chrosomus, 85 nerka, Oncorhynchus, 4, 76, 143, 269, 272–274 nerka, Salmo, 272 nerka kennerlyi, Oncorhynchus, 272 Nezumia, 333 Nezumia aequalis, 81 Nezumia bairdii, 77, 124, 344–345 niger, Chiasmodon, 4, 539, 540 niger, Malacosteus, 76, 147, 311–312 nigricans, Bathylaco, 81 nigricans, Lycodes, 505 nigrifilis, Danaphryne, 82 nigrum, Etheostoma, 85 nitidus, Salmo, 283 Normichthys, 131 Normichthys operosus, 3, 76, 97, 131, 231 Normichthys operosa islandica, 231 norvegicus, Lepidoleprus, 338 norvegicus, Sebastes, 77, 144, 406–407 Notacanthus chemnitzii, 75, 100, 128, 206–207 Notacanthus phasgonorus, 206 Notacanthus rostratus, 207 Notocanthus nasus, 206 Notoscopelus elongatus, 329 Notoscopelus kroyeri, 76, 126, 329–330 Notropis atherinoides, 85 Notropis hudsonius, 85 nubilis, Lumpenus, 522 nueltinensis, Leucichthys, 251 nybelini, Triglops, 22, 78, 115, 431–432 obensis, Lota lota, 373 oblita, Nansenia, 81 occidentalis, Pungitius, 399 occidentalis, Pungitius pungitius, 399

Index

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octodecemspinosus, Myoxocephalus, 78, 107, 114, 420–421 ogac, Gadus, 35, 74, 77, 119, 358, 371–373 Oidiphorus, ix Oidiphorus mcallisteri, ix olfersi, Argyropelecus, 81 olidus, Hypomesus, 85 olrikii, Aspidophoroides, 5, 78, 108, 436–438 omiscomaycus, Percopsis, 85 ommatistius terraenovae, Triglops, 429 Oncorhynchus, 264 Oncorhynchus gorbuscha, 3, 76, 143, 264–267 Oncorhynchus keta, 3, 76, 143, 267–270 Oncorhynchus kisutch, 76, 143, 270–272 Oncorhynchus nerka, 4, 76, 143, 269, 272–274 Oncorhynchus nerka kennerlyi, 272 Oncorhynchus nerka nerka, 272 Oncorhynchus tschawytscha, 275 Oncorhynchus tshawytscha, 76, 143, 275–277 onegensis, Lota lota, 373 Oneirodes anisacanthus, 383 Oneirodes eschrichtii, 82, 383 Oneirodes macrosteus, 82 Oneirodes sp., 77, 128, 383–384 Onogadus, 353, 355 Onos, 351, 353, 355 Onos rufus, 355 opalescens, Platyberyx, 82 operosa islandica, Normichthys, 231 operosus, Normichthys, 3, 76, 97, 131, 231 Ophidium parrii, ix, 460 oquassa, Salvelinus alpinus, 283 orientalis, Anarhichas, ix, 4, 79, 103, 110, 531, 537–538 Osmerus dentex, 3, 4, 76, 105, 129, 244–245, 246 Osmerus mordax, 3, 4, 21, 76, 129, 244, 245, 246–247 Osmerus mordax dentex, 244 ouananiche, Salmo salar, 279 oxyrinchus, Acipenser, 80, 85, 203 paamiuti, Lycodes, 4, 79, 155, 497–499, 506, 507, 508 Pachycara, 488 pacifica, Motella, 351 pacificus, Somniosus, 3, 4, 75, 93, 145, 183–184 palearis, Lycodes, 83 pallasii, Clupea, 58, 75, 105, 111, 221–223, 299 pallidus, Hydrolagus, 81, 171

pallidus, Lycodes, 79, 155, 493, 497, 499–500, 506 pallidus marisalbi, Lycodes, 499 pallidus similis, Lycodes, 499 pallidus var. squamiventer, Lycodes, 499 panthera, Lycodes, 496 panthera, Lycodes knipowitschi, 496 parabeani, Serrivomer, 214 Paracaristius maderensis, 82 Paralepis, 320 Paralepis coregonoides, 76, 130, 321 Paralepis coregonoides barracudina, 321 Paralepis coregonoides borealis, 76, 321 Paralepis coregonoides coregonoides, 321 Paralepis kroyeri, 318 Paraliparis, 469 Paraliparis bathybii, 466 Paraliparis bathybius, 78, 122, 466–467 Paraliparis copei, 4, 78, 123, 467–468 Paraliparis garmani, 78, 123, 468–469 Paraliparis hystrix, 82 parasitica, Simenchelys, 3, 4, 75, 98, 148, 209 Parophrys ischyrus, 561 Parophrys vetulus, 561 parrii, Ophidium, ix, 460 patris, Acanthocottus, 413 paucidens, Lycichthys, 530 pauciradiatus, Diretmoides, 82 paxillus, Lycenchelys, 78, 151, 482–483, 484 pearyi, Boreogadus, 359 pelecanoides, Eurypharynx, 75, 95, 217–218 Peprilus triacanthus, 79, 98, 547–548 Perca flavescens, 85 Percina caprodes, 85 Percina shumardi, 85 Percopsis omiscomaycus, 85 perspicillum, Lycodes, 502 pertusus, Merlangus, 375 Petromyzon, 165 Petromyzon borealis, 165 Petromyzon fluvialis, 165 Petromyzon japonicus, 165 Petromyzon marinus, 81 phalacra, Aldrovandia, 81 pharao, Anotopterus, 76, 129, 317–318 phasgonorus, Notacanthus, 206 Phocaegadus megalops, 359 Pholis fasciata, 79, 107, 528–529 Pholis gunnellus, 83 Photostylus pycnopterus, 81 phrictus, Psychrolutes, 4, 78, 135, 444–445



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Phycis, 356 Phycis blennoides, 351 Phycis chesteri, 77, 106, 130, 356–357 Phyllorhinichthys balushkini, 82 Phyllorhinichthys micractis, 82 Physiculus, 346 pidschian, Coregonus, 81, 257 Pimephales promelas, 85 pingelii, Triglops, 5, 78, 116, 433–434 pinguis, Pleuronectes, 568 piscatorius, Lophius, 82, 378 Platessa, 561 platessa, Pleuronectes, 83 Platessa dwinensis, 564 platessoides, Hippoglossoides, 4, 22, 79, 134, 551–554 platessoides, Hippoglossoides platessoides, 551, 554 platessoides limandoides, Hippoglossoides, 551, 554 platessoides platessoides, Hippoglossoides, 551, 554 Platichthys rugosus, 561 Platichthys stellatus, 4, 79, 132, 561–563, 564 Platichthys stellatus rugosus, 561 Platichthys stellatus stellatus, 561 Platyberyx opalescens, 82 platycephalus, Gymnelus, 478 platycephalus, Megalocottus, 82 Platygobio gracilis, 85 Platysomatichthys, 568 Platytroctes apus, xii, 3, 76, 131, 232–233 Pleurogadus, 365 Pleuronectes, 549, 559, 561 Pleuronectes franklinii, 564 Pleuronectes glacialis, 8, 79, 134, 549, 564–566 Pleuronectes pinguis, 568 Pleuronectes platessa, 83 Pleuronectes putnami, 79, 96, 134, 564, 566–567 Pleuronectes quadrituberculatus, 83 pleurostictus, Triglops, 433 plumbea, Chimaera, 171 plumbeus, Couesius, 85 Podothecus veternus, 82 polaris, Cottus, 422 polaris, Lycodes, 79, 152, 500–501 polaris, Merlangus, 360 Pollachius virens, 80, 82 pollan, Coregonus, 254 polli, Polyipnus, 81 Polyacanthonotus rissoanus, 75, 128, 207–208 Polyipnus asteroides, 81 Polyipnus polli, 81 Polymetme corythaeola, 81 Poromitra capito, 82 Poromitra crassiceps, 82 porosus, Cottus, 426

poutassou, Micromesistius, 4, 77, 118, 375–376 praecisus, Eumesogrammus, 79, 145, 519–520 preblei, Coregonus, 277 pribilovius, Nautichthys, 82 princeps, Etmopterus, 81, 175 proboscidea, Limanda, 79, 134, 559–560 profundorum, Apristurus, 3, 4, 75, 93, 174 promelas, Pimephales, 85 Prosopium, 248, 277 Prosopium cylindraceum, 5, 76, 139, 277–279 Prosopium hearnei, 277 Protomyctophum arcticum, 76, 126, 331 Psednos groenlandicus, 82 Psednos harteli, 82 Psednos melanocephalus, 82 Psednos micruroides, 82 pseudobscura, Sternoptyx, 81 Pseudopleuronectes, 549 Pseudopleuronectes americanus, 83, 282 Psychrolutes phrictus, 4, 78, 135, 444–445 punctatum, Myctophum, 3, 76, 127, 328–329 punctatus, Stichaeus, 79, 107, 146, 524–526 pungitius, Pungitius, 4, 5, 77, 119, 399–401 Pungitius occidentalis, 399 pungitius occidentalis, Pungitius, 399 Pungitius pungitius, 4, 5, 77, 119, 399–401 Pungitius pungitius occidentalis, 399 pusillus, Coregonus, 262 putnami, Pleuronectes, 79, 96, 134, 564, 566–567 pycnopterus, Photostylus, 81 quadricornis, Myoxocephalus, ix, 3, 4, 5, 78, 114, 421–424, 426 quadricornis, Myoxocephalus quadricornis, 422 quadricornis hexacornis, Myoxocephalus, ix, 422 quadricornis quadricornis, Myoxocephalus, 422 quadrilateralis, Coregonus, 277 quadrituberculatus, Pleuronectes, 83 radiata, Amblyraja, 4, 75, 136, 181, 186, 188, 190–192 Raia, 196 Raia americana, 191 Raia scabrata, 191

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Raja, 185, 186, 191, 194, 196, 197, 199, 201 Raja (Amblyraja) sp., 193 Raja bathyphila, 197 Raja borea, 186 Raja clavata, 191 Raja granulata, 189 Raja hyperborea, 186 Raja ingolfiana, 201 Raja laevis, 191 Raja lintea, 197 Raja mollis, 196 Rajella, 185, 197, 201 Rajella bathyphila, 3, 4, 75, 138, 197–199, 201 Rajella bigelowi, 81, 185, 197 Rajella fyllae, 4, 75, 136, 199–201 Rajella lintea, 4, 75, 138, 197, 201–202 raleighana, Harriotta, 3, 4, 75, 138, 168–169 raridens, Lycodes, 8, 83, 491 rastrinus, Careproctus cf., 82 regia, Urophycis, 81 regina, Rhodichthys, 78, 122, 469–470 reinhardi, Motella, 353 reinhardti, Careproctus, 78, 121, 454, 455, 456–458 reinhardti, Himantolophus, 380 Reinhardtius hippoglossoides, 3, 5, 12, 14, 22, 23, 79, 133, 160, 181, 182, 568–571 Reinhardtius matsuurae, 568 reticulatus, Lycodes, 79, 101, 153, 181, 491, 492, 502–503 reticulatus hacheyi, Lycodes, 502 reticulatus laurentianus, Lycodes, 502 retrodorsalis, Gymnelus, 78, 149, 476–478 Rhadinesthes decimus, 76, 148, 312–313 Rhinichthys atratulus, 85 Rhinichthys cataractae, 85 Rhinochimaera atlantica, 3, 4, 75, 96, 138, 169–170 Rhinonemus, 351 Rhinoscymnus, 180 Rhodichthys regina, 78, 122, 469–470 ricei, Cottus, 85 richardsoni, Astronesthes cf., 76, 148, 308 risso, Arctozenus, 76, 104, 129, 318–319 risso, Arctozenus risso, 318, 319 risso kroyeri, Arctozenus, 318, 319 risso risso, Arctozenus, 318, 319 rissoanus, Polyacanthonotus, 75, 128, 207–208 robustus, Hippoglossoides, 8, 79, 134, 555–556 robustus, Scopeloberyx, 4, 77, 100, 390–391

610

Rondeletia loricata, 82 rossi, Lycodes, 4, 502, 503 Rossii, Salmo, 283 rostrata, Anguilla, 81 rostrata, Antimora, 22, 23, 77, 125, 210, 347–348 rostratus, Notacanthus, 207 Rouleina attrita, 81 Rouleina maderensis, xii, 3, 76, 109, 238 rufa, Barbourisia, 82 rufus, Onos, 355 rugosus, Platichthys, 561 rugosus, Platichthys stellatus, 561 rupertianus, Acipenser, 203 rupestris, Coryphaenoides, 4, 77, 124, 210–211, 333, 338–340, 342 rupestris, Gadus, 367 rutteri, Liparis, 80 Saccopharynx ampullaceus, 75, 95, 216, 348 sadko, Cottunculus, 440, 442 Sagamichthys schnakenbecki, 81 sagittarius, Lycodes, ix, 8, 79, 154, 493, 504–505 saida, Boreogadus, 3, 5, 20, 22, 23, 32, 48, 77, 118, 358, 360–363, 409, 541, 553, 568, 569 salar, Salmo, 4, 28, 76, 105, 141, 279–282 salar, Salmo salar, 279 salar ouananiche, Salmo, 279 salar salar, Salmo, 279 salar sebago, Salmo, 279 Salmo, 67, 254, 264, 279, 297 Salmo alipes, 283 Salmo alpinus, 283 Salmo arcturus, 283 Salmo Cylindraceus, 277 Salmo groenlandicus, 240 Salmo harengus, 251 Salmo Hearnii, 292 Salmo hudsonicus, 289 Salmo kennerlyi, 272 Salmo leucichthys, 300 Salmo Mackenzii, 300 Salmo namaycush, 297 Salmo naresi, 283 Salmo Naresii, 283 Salmo nelma, 300 Salmo nerka, 272 Salmo nitidus, 283 Salmo Rossii, 283 Salmo salar, 4, 28, 76, 105, 141, 279–282 Salmo salar ouananiche, 279 Salmo salar salar, 279 Salmo salar sebago, 279 Salmo trutta, 280 Salmo tullibee, 251 Salvelinus, 67, 248, 298

salvelinus, Salvelinus, 283 salvelinus, Salvelinus alpinus, 283 Salvelinus alpinus, 3, 4, 5, 19, 67, 76, 141, 142, 283–289, 290, 292, 297 Salvelinus alpinus alpinus, 283 Salvelinus alpinus erythrinus, 283 Salvelinus alpinus oquassa, 283 Salvelinus alpinus salvelinus, 283 Salvelinus alpinus taranetzi, 283 Salvelinus confluentus, 292 Salvelinus fontinalis, 29, 67, 76, 142, 284, 285, 289–292 Salvelinus malma, 21, 76, 142, 284–285, 287, 292–297 Salvelinus malma lordi, 292 Salvelinus malma malma, 4, 292 Salvelinus namaycush, 3, 4, 5, 19, 76, 142, 283, 284, 285, 297–299 Salvelinus salvelinus, 283 Salvelinus umbla, 283 Sander canadensis, 85 Sander vitreus, 85 sardinella, Coregonus, 4, 21, 76, 140, 253, 254, 262–264 sarsii, Lycenchelys, 78, 151, 480, 482, 483–484 scaber, Artediellus, 77, 112, 410–411 scabrata, Raia, 191 schnakenbecki, Sagamichthys, 81 schulzi, Mallotus villosus catervarius natio, 240 scolopaceus, Nemichthys, 75, 98, 212–213 Scopeloberyx robustus, 4, 77, 100, 390–391 Scopelogadus beani, 82 Scopelosaurus harryi, 316 Scopelosaurus lepidus, 76, 104, 316 scorpioides, Myoxocephalus, 5, 78, 115, 158, 414, 424–426 scorpius, Myoxocephalus, 5, 32, 78, 82, 115, 181, 414, 422, 426–429 scorpius groenlandicus, Myoxocephalus, 426 Scymnodon melas, 178 Scymnus glacialis, 180 Scymnus micropterus, 180 Searsia koefoedi, 81 sebago, Salmo salar, 279 Sebastes, 65, 339, 340, 344, 347, 401, 405, 533, 534, 553 Sebastes fasciatus, 4, 5, 77, 144, 402–404 Sebastes marinus, 401, 404, 406, 407 Sebastes mentella, 4, 22, 23, 77, 107, 144, 403, 404–406, 407, 568, 569 Sebastes norvegicus, 77, 144, 406–407 Sebastes sp., 224, 402 Sebastes viviparus, 82 seminudus, Lycodes, 79, 153, 485, 505–506

Semotilus corporalis, 85 septentrionalis, Ciliata, 81 serpentinus, Lumpenus lampretaeformis, 523 Serrivomer beanii, 75, 99, 214–215 Serrivomer parabeani, 214 shumardi, Percina, 85 Sigmops bathyphilus, 76, 120, 304–305 silus, Argentina, 75, 104, 224–225 Simenchelys parasitica, 3, 4, 75, 98, 148, 209 similis, Lycodes pallidus, 499 sloanei, Chauliodus, 310 sloani, Chauliodus, 76, 147, 310–311 Somniosus, 178, 180 Somniosus antarcticus, 184 Somniosus brevipinna, 180 Somniosus microcephalus, 4, 8, 22, 75, 93, 145, 180–183, 188, 192 Somniosus pacificus, 3, 4, 75, 93, 145, 183–184 sp., Ammodytes, 291 sp., Bathyraja, 75, 185, 193–194 sp., Bathytroctes, 3, 76, 109, 237 sp., Cottunculus, 492 sp., Gaidropsarus, 22, 23 sp., Liparis, 428, 434 sp., Lycodes, 22, 23, 428, 492 sp., Mallotus, 8 sp., Myoxocephalus, 434 sp., Nansenia, 81 sp., Oneirodes, 77, 128, 383–384 sp., Raja (Amblyraja), 193 sp., Sebastes, 224, 402 sp., Triglops, 427, 428 spatula, Icelus, 78, 112, 417–418 speculigera, Lampadena, 81 spinacidermis, Malacoraja, 4, 75, 136, 196–197 Spinax fabricii, 175 spinicauda, Bathyraja, 4, 75, 92, 135, 193, 194–195 Spiniphryne gladisfenae, 77, 128, 385 spinosus, Eumicrotremus, 78, 117, 452–453 spinosus variabilis, Eumicrotremus, 450, 452 Squalus acanthias, 81, 175 Squalus microcephalus, 180 squamiventer, Lycodes, 4, 79, 155, 497, 499, 504, 506–507 squamiventer, Lycodes pallidus var., 499 stellatus, Platichthys, 4, 79, 132, 561–563, 564 stellatus, Platichthys stellatus, 561 stellatus rugosus, Platichthys, 561 stellatus stellatus, Platichthys, 561 stelleri, Hexagrammos, 82 Stenodus, 248

Index

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Stenodus leucichthys, 4, 30, 76, 139, 248, 300–301 Stenodus nelma, 300 stenolepis, Hippoglossus, 556 Sternoptyx pseudobscura, 81 Stichaeus punctatus, 79, 107, 146, 524–526 Stomias, 307 stomias, Atheresthes, 8 Stomias boa, 76, 147, 313–315 Stomias boa boa, 313 Stomias boa ferox, 76, 313 stroemii, Macrourus, 338 subautumnalis, Coregonus, 254 subbifurcata, Ulvaria, 4, 79, 146, 526–527 suborbitalis, Macrurus, 333 Symbolophorus veranyi, 77, 127, 332 Synaphobranchus kaupii, 4, 75, 98, 148, 210–211 taeniatus, Gymnelus, 478 taranetzi, Salvelinus alpinus, 283 tenuis, Urophycis, 81, 351 tergisus, Hiodon, 85 terraenovae, Lumpenus lampetraeformis, 523 terraenovae, Lycodes, 79, 155, 486, 487, 488, 491, 508–509 terraenovae, Triglops ommatistius, 429 thompsonii, Myoxocephalus, ix thomsonii, Cottunculus, 78, 107, 135, 442–443 Thunnus thynnus, 83 Thymallus arcticus, 85 thynnus, Thunnus, 83 tomcod, Microgadus, 80, 358 torsk, Gadus, 364 Trachipterus arcticus, 81 Trachyrhynchus, 345 Trachyrinchus, 345 Trachyrincus, 333 Trachyrincus murrayi, 77, 123, 342, 344, 345–346 triacanthus, Peprilus, 79, 98, 547–548 tricuspis, Gymnocanthus, 5, 78, 113, 413–415 tricuspis groenlandicus, Gymnacanthus, 413 tricuspis hudsonius, Gymnocanthus, 413 tridentatus, Entosphenus, 165 Triglops, ix, 286, 408, 422, 429, 434, 553 Triglops murrayi, 78, 116, 429–430 Triglops nybelini, 22, 78, 115, 431–432 Triglops ommatistius terraenovae, 429 Triglops pingelii, 5, 78, 116, 433–434

Triglops pleurostictus, 433 Triglops sp., 427, 428 Triglopsis, 422 Trigonolampa miriceps, 81 Trisopterus, 364 Trisopterus esmarkii, 82 trutta, Salmo, 280 tschawytscha, Oncorhynchus, 275 tshawytscha, Oncorhynchus, 76, 143, 275–277 tullibee, Salmo, 251 tunicatus, Liparis, 5, 78, 122, 464–465 turneri, Lycodes, 83, 500 turneri atlanticus, Lycodes, 500

viride, Gymnelus, 478 viridis, Gymnelus, 8, 30, 78, 150, 474, 475, 478–479 viridis var. unimaculatus, Gymnelus, 478 vitreus, Sander, 85 viviparus, Sebastes, 82 viviparus, Zoarces, 522 vulgaris, Brosmus, 364 vulgaris, Hippoglossus, 556 vulgaris, Morhua, 367

Ulcina, 436 Ulvaria subbifurcata, 4, 79, 146, 526–527 umbla, Salvelinus, 283 uncinatus, Artediellus, 77, 111, 113, 412–413 unicuspis, Ichthyomyzon, 85 unimaculatus, Clinus, 519 unimaculatus, Gymnelus viridis var., 478 Urophycis, 356 Urophycis brasiliensis, 351 Urophycis chuss, 351 Urophycis regia, 81 Urophycis tenuis, 81, 351

zenithicus, Coregonus, 251 Zoarces, 513 Zoarces americanus, 79, 149, 473, 513–514 Zoarces viviparus, 522 zoarchus, Lycodes, 509

vachonii, Lycodes, 486 vahlii, Lycodes, 79, 154, 508, 509–510 vahlii, Lycodes vahlii, 509 vahlii gracilis, Lycodes, 509 vahlii maculatus, Lycodes, 509 vahlii vahlii, Lycodes, 509 vanhoeffeni, Gigantactis, 4, 77, 94, 388–389 variabilis, Coryphaenoides, 333 variabilis, Coryphaenoides armatus, 333 variabilis, Eumicrotremus spinosus, 450, 452 veranyi, Symbolophorus, 77, 127, 332 verrillii, Lycenchelys, 80 vernalis, Merlangus, 375 verrucosus, Allocyttus, 82 verrucosus, Myoxocephalus, 82 vertagus, Gadus, 367 veternus, Podothecus, 82 vetulus, Parophrys, 561 villosus, Mallotus, 5, 76, 129, 240– 244, 281, 286, 291, 299, 553 villosus, Mallotus villosus, 240 villosus catervarius, Mallotus, 240 villosus catervarius natio schulzi, Mallotus, 240 villosus villosus, Mallotus, 240 viola, Haloporphyrus, 347 virens, Pollachius, 80, 82



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wachna, Gadus, 365 Xenodermichthys copei, 76, 109, 239

English Common Names Abyssal Skate, 75, 197–199 Acadian Redfish, 77, 402–404, 553 Adolf ’s Eelpout, 78, 484–485 Alaska Plaice, 559 Alaskan Whitefish, 257 Alligatorfish, 78, 435–436 Arctic, 78, 414, 436–438 American Plaice, 79, 368, 551–554, 558 Anglemouth Spark, 76, 304–305 Veiled, 76, 302–303 Antimora, Blue, 77, 347–348 Archer Eelpout, 79, 493, 504–505 Arctic Alligatorfish, 78, 414, 436–438 Arctic Brotula, 77, 377 Arctic Char, 19, 20, 21, 24, 25, 27, 29, 32, 34, 35, 36, 39, 48, 57, 58, 61, 64, 65, 67, 76, 166, 181, 242, 248, 249, 250, 283–289, 290, 292, 292, 293, 297, 299, 361, 362, 372, 400, 407, 414, 416, 423, 424, 425, 428, 434, 437, 450, 452, 461, 465, 517, 521, 522, 524, 542 Arctic Cisco, 21, 24, 25, 43, 58, 76, 166, 244, 252, 254–256, 262, 263, 264, 295, 423, 544, 564 Arctic Cod, 20, 21, 24, 25, 27, 29, 32, 35, 36, 38, 39, 44, 48, 49, 51, 52, 57, 58, 66, 77, 241, 242, 244, 295, 358, 359, 360–363, 368, 372, 414, 416, 423, 427, 428, 434, 451, 461, 465, 501, 520, 525, 528, 541, 544 Arctic Eelpout, 79, 491, 492, 502–503

Arctic Flounder, 25, 79, 423, 564–566 Arctic Hookear Sculpin, 77, 412–413 Arctic Lamprey, xiii, 75, 165–167, 222, 245, 258, 261, 295, 300 Arctic Lumpsucker, 78, 447–448 Arctic Sculpin, 78, 158, 408, 423, 424–426, 427 Arctic Shanny, 79, 372, 524–526 Arctic Skate, 38, 51 Arctic Staghorn Sculpin, 78, 413–415, 427, 428, 437 Arctic Telescope, 76, 331 Argentine, Atlantic, 75, 224–225 Arrowtooth Flounder, 8 Atlantic Argentine, 75, 224–225 Atlantic Cod, 34, 58, 64–65, 77, 163–164, 214, 220, 224, 242, 246, 310, 315, 316, 317, 319, 324, 325, 329, 330, 334, 340, 343, 344, 358, 361, 364, 365, 367–371, 372, 375, 376, 401, 404, 409, 414, 417, 419, 421, 423, 425, 428, 429, 431, 434, 436, 451, 452, 461, 462, 478, 513, 517, 520, 521, 522, 525, 526, 531, 533, 534, 541, 542, 551, 553, 558, 570 Atlantic Footballfish, 77, 380–381 Atlantic Gymnast, 76, 239 Atlantic Hagfish, 75, 162–164 Atlantic Halibut, 79, 317, 549, 556–559 Atlantic Herring, 75, 163, 164, 166, 219–221, 281, 368, 369, 449, 558 Atlantic Hookear Sculpin, 77, 408–410 Atlantic Poacher, 78, 438–439 Atlantic Salmon, 28, 29, 38, 58, 63, 64, 65, 76, 166, 242, 246, 250, 279–282, 292, 321, 330, 361, 362, 542, 569, 570 Atlantic Seasnail, 78, 458–459 Atlantic Soft Pout, 79, 512 Atlantic Spiny Lumpsucker, 78, 427, 452–453, 531 Atlantic Warbonnet, 79, 518–519 Atlantic Wolffish, 79, 530, 532–534 Aurora Pout, 78, 476–478 Baikal Omul, 254 Banded Gunnel, 79, 528–529 Banded Manefish, 78, 471–472 Barndoor Skate, 513 Barracudina Duckbill, 76, 320 Sharpchin, 76, 317, 321 White, 76, 318–319 Barsukov’s Pout, 78, 473–474 Bering Cisco, 254, 256, 262 Bering Flounder, 79, 414, 555–556 Bering Wolffish, ix, 79, 531, 537–538 Bigelow’s Skate, 185

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Bigeye Sculpin, 78, 429, 431–432, 433 Bigeye Smoothhead, 76, 236 Bighead Searsid, 75, 228 Bigscale, Longjaw, 77, 390–391 Black Dogfish, 75, 175–177, 404 Black Scabbardfish, 79, 348, 546 Black Seasnail, 78, 466–467 Blackfin Waryfish, 76, 316 Blackline Prickleback, ix, 79, 515–516 Blacksmelt, Goitre, 75, 226–227, 343 Blacksnout Snailfish, 78, 467–468 Blob Sculpin, Giant, 78, 444–445 Blue Antimora, 77, 347–348 Blue Whiting, 77, 368, 375–376 Boa Dragonfish, 76, 313–315 Broad Whitefish, 58, 62, 65, 76, 166, 250, 257, 258, 260–262, 264, 295 Brook Trout, 29, 57, 58, 64, 67, 76, 242, 246, 250, 285, 289–292, 423, 522, 544 Brotula, Arctic, 77, 377 Bull Trout, 292 Butterfish, 79, 547–548 Canadian Eelpout, 79, 500–501 Can-Opener Smoothdream, 77, 382–383 Capelin, 24, 25, 32, 76, 192, 240–244, 246, 281, 286, 291, 299, 343, 368, 372, 423, 427, 526, 541, 558, 569 Carapine Grenadier, 77, 336–337 Cat Shark, Deepsea, 75, 174 Char, Arctic, 19, 20, 21, 24, 25, 27, 28, 29, 32, 34, 35, 36, 39, 48, 57, 58, 61, 64, 65, 67, 76, 166, 181, 242, 248, 249, 250, 283–289, 290, 292, 292, 293, 297, 299, 361, 362, 372, 400, 407, 414, 416, 423, 424, 425, 428, 434, 437, 450, 452, 461, 465, 517, 521, 522, 524, 542 Checker Eelpout, 79, 509–510 Checkered Wolf Eel, 78, 480 Chevron Scutepout, 79, 510–511 Chimaera Deepwater, 75, 171–173 Haeckel’s, 167 Knifenose, 75, 169–170 Longnose, 75, 168–169, 171 Chinook Salmon, 76, 166, 266, 275–277 Chum Salmon, 76, 166, 267–270 Cisco, 57, 76, 166, 242, 251–253, 254, 256, 264, 372, 423, 522, 544 Arctic, 21, 24, 25, 43, 58, 76, 166, 244, 252, 254–256, 262–264, 295, 423, 544, 564 Bering, 254, 256, 262 Least, 21, 24, 57, 76, 166, 252, 254, 255, 256, 262–264 Penzhina, 254 Shortjaw, 251

612

Cod Arctic, 20, 21, 24, 25, 27, 29, 32, 35, 36, 38, 39, 44, 48, 49, 51, 52, 57, 58, 66, 77, 241, 242, 244, 295, 358, 359, 360–363, 368, 372, 414, 416, 423, 427, 428, 434, 451, 461, 465, 501, 520, 525, 528, 541, 544 Atlantic, 34, 58, 64, 65, 77, 163, 164, 214, 220, 224, 242, 246, 310, 315, 316, 317, 319, 324, 325, 329, 330, 334, 340, 343, 344, 358, 361, 364, 365, 367–371, 372, 375, 376, 401, 404, 409, 414, 417, 419, 421, 423, 425, 428, 429, 431, 434, 436, 451, 452, 461, 462, 478, 513, 517, 520, 521, 522, 525, 526, 531, 533, 534, 541, 542, 551, 553, 558, 570 Greenland, 35, 38, 77, 361, 371–373 Polar, 18, 34, 77, 358–360, 569 Saffron, 21, 24, 77, 166, 242, 299, 365–367, 564 Coho Salmon, 76, 266, 270–272, 276 Common Wolf Eel, 78, 482–483 Cornerlantern, North Atlantic, 77, 332 Cusk, 77, 364–365 Cutthroat Eel, Northern, 75, 210–211, 343 Dab, Longhead, 79, 559–560 Daggertooth, 76, 317–318 Dainty Mora, 77, 348–349 Darkbelly Skate, 75, 186–188 Daubed Shanny, 79, 520–521 Deepsea Cat Shark, 75, 174 Deepwater Chimaera, 75, 171–173 Deepwater Redfish, 77, 317, 402, 403, 404–406 Diamondcheek Lanternfish, 76, 326 Doctor, Fish, 78, 427, 428, 478–479 Dogfish Black, 75, 175–177, 404 Spiny, 163, 164, 175, 211, 224, 356, 364, 369, 550, 553, 558 Dolly Varden, 21, 24, 76, 166, 283, 285, 289, 292–297, 361 Doubleline Eelpout, 78, 487–488 Dragonfish, Boa, 76, 313–315 Dreamer, Prickly, 77, 385 Duckbill Barracudina, 76, 320 Dusky Slickhead, 76, 234 Dusty Snailfish, 8 Eel Checkered Wolf, 78, 480 Common Wolf, 78, 482–483 Moray Wolf, 78, 481–482 Northern Cutthroat, 75, 210–211, 343 Slender Snipe, 75, 212–213 Snubnose, 75, 209 Snubnosed Spiny, 75, 206–207

Eelblenny Slender, 79, 372, 423, 522–523 Stout, 79, 517–518 Eelpout Adolf ’s, 78, 484–485 Archer, 79, 493, 504–505 Arctic, 79, 491, 492, 502–503 Canadian, 79, 500–501 Checker, 79, 509–510 Doubleline, 78, 487–488 Glacial, ix, 78, 488–489 Greater, 78, 486–487 Laval, 78, 491–492 Longear, 79, 505–506 Manytoothed, 8 Marbled, 8 McAllister’s, 79, 495 Newfoundland, 79, 491, 508–509 Paamiut, 79, 497–499 Pale, 79, 499–500 Pink, 78, 492–493 Saddled, 79, 423, 496–497 Scalebelly, 79, 506–507 Theologian, 78, 483–484 White Sea, 79, 493–494 English Sole, 561 Fangtooth, 77, 392–394 Fish Doctor, 78, 427, 428, 478–479 Flathead Sole, 555 Flounder Arctic, 25, 79, 423, 564–566 Arrowtooth, 8 Bering, 79, 414, 555–556 Smooth, 79, 564, 566–567 Starry, 79, 166, 242, 561–563 Witch, 79, 549–551, 569 Footballfish, Atlantic, 77, 380–381 Forkline Sole, 561 Fourbeard Rockling, 77, 351–353 Fourhorn Sculpin, 78, 244, 361, 362, 366, 372, 421–424, 427, 428, 496, 522, 544, 564 Fourline Snakeblenny, 79, 519–520, 525, 526 Gelatinous Snailfish, 78, 460–462 Ghost Shark, 171 Giant Blob Sculpin, 78, 444–445 Giant Seadevil, Northern, 77, 386–387 Glacial Eelpout, ix, 78, 488–489 Glacier Lanternfish, 76, 323–324, 331, 404 Goitre Blacksmelt, 75, 226–227, 343 Golden Redfish, 77, 403, 405, 406–407, 531 Goosefish, 77, 213, 356, 364, 378–379, 550, 553, 558 Great Lanternshark, 175 Greater Eelpout, 78, 486–487

Greater Silver Hatchetfish, 76, 306–307 Greenland Cod, 35, 38, 77, 361, 371–373 Greenland Halibut, 12, 14, 28, 34, 36, 38, 51, 52, 58, 64, 65, 79, 177, 181, 182, 188, 192, 220, 242, 317, 319, 321, 324, 339, 340, 343, 348, 355, 361, 363, 364, 365, 368, 372, 396, 401, 404, 439, 462, 525, 531, 534, 536, 549, 550, 554, 558, 568–571 Greenland Shark, 8, 34, 38, 64, 75, 178, 179, 180–183, 184, 192, 242, 450, 509, 531, 533, 534, 553, 558, 570 Grenadier Carapine, 77, 336–337 Günther’s, 77, 337–338 Rock, 77, 226, 303, 338–340, 343 Roughhead, 77, 211, 226, 242, 342–343, 458 Roughnose, 77, 345–346 Russet, 77, 333–335 Shortbeard, 77, 335–336 Threadfin, 77, 340–341 Grubby, 78, 419–420, 421, 526 Gulper Pelican, 75, 216, 217–218 Taillight, 75, 216, 217, 348 Gunnel, Banded, 79, 528–529 Günther’s Grenadier, 77, 337–338 Gymnast, Atlantic, 76, 239 Haddock, 80, 163, 164, 242, 343, 368, 436, 551, 558 Haeckel’s Chimaera, 167 Hagfish, Atlantic, 75, 162–164 Hake, Longfin, 77, 356–357 Halibut Atlantic, 79, 317, 549, 556–559 Greenland, 12, 14, 28, 34, 36, 38, 51, 52, 58, 64, 65, 79, 177, 181, 182, 188, 192, 220, 242, 317, 319, 321, 324, 339, 340, 343, 348, 355, 361, 363, 364, 365, 368, 372, 396, 401, 404, 439, 462, 525, 531, 534, 536, 549, 550, 554, 558, 568–571 Hamecon, 77, 410–411 Hartel’s Swallower, 79, 539–540 Hatchetfish, Greater Silver, 76, 306–307 Herring Atlantic, 75, 163, 164, 166, 219–221, 281, 368, 369, 449, 558 Pacific, 58, 64, 75, 166, 219, 221–223, 251, 254, 270, 275, 299, 300 Hookear Sculpin, Arctic, 77, 412–413 Humpback Whitefish, 257 Hybrid Sole, 561

Index

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Inconnu, 65, 76, 166, 222, 246, 263, 300–301, 423, 428 Irish Pollan, 254 Jewel Lanternfish, 76, 324–325 Kelp Snailfish, 78, 460, 464–465 Kido’s Snailfish, 78, 454–455 Knifenose Chimaera, 75, 169–170 Knipowitsch’s Pout, 78, 475–476 Kokanee, 272, 273, 274, 374 Lake Sturgeon, 32, 75, 203–205, 242, 544 Lake Trout, 19, 76, 166, 223, 246, 263, 278, 285, 297–299 Lake Whitefish, 19, 57, 58, 76, 166, 257–259, 260, 299, 423, 522, 544 Lamprey, Arctic, xiii, 75, 165–167, 222, 245, 258, 261, 295, 300 Lance Northern Sand, 79, 372, 541–543 Pacific Sand, 79, 542, 543–545 Lanternfish, 76, 328–329 Diamondcheek, 76, 326 Glacier, 76, 323–324, 331, 404 Jewel, 76, 324–325 Rakery, 76, 327 Spotted, 76, 328–329 Lanternshark, Great, 175 Large-Eye Snaggletooth, 76, 309 Largeye Lepidion, 77, 349–350 Laval Eelpout, 78, 491–492 Least Cisco, 21, 24, 57, 76, 166, 252, 254, 255, 256, 262–264 Leatherfin Lumpsucker, 78, 450–452 Legless Searsid, 76, 232–233 Lepidion, Largeye, 77, 349–350 Linen Skate, 75, 201–202 Longear Eelpout, 79, 505–506 Longfin Hake, 77, 356–357 Longfin Snailfish, 78, 455–456 Longhead Dab, 79, 559–560 Longhorn Sculpin, 78, 419, 420–421, 513 Longjaw Bigscale, 77, 390–391 Longnose Chimaera, 75, 168–169, 171 Longsnout Manypitshoulder, 76, 231 Loosejaw, Stoplight, 76, 311–312 Lumpfish, 78, 446, 449–450, 558 Smooth, 78, 446–447 Lumpsucker Arctic, 78, 447–448 Atlantic Spiny, 78, 427, 452–453, 531 Leatherfin, 78, 450–452 Madeiran Smoothhead, 76, 238 Manefish, Banded, 78, 471–472 Manylight Viperfish, 76, 310–311

Manypitshoulder, Longsnout, 76, 231 Manyray Smoothhead, 76, 235 Manytoothed Eelpout, 8 Marbled Eelpout, 8 Marlin-spike, 77, 344–345 Maul’s Searsid, 75, 229 McAllister’s Eelpout, 79, 495 Mora, Dainty, 77, 348–349 Moray Wolf Eel, 78, 481–482 Moustache Sculpin, 78, 429–430, 431, 433 Nebulous Snailfish, 8 Newfoundland Eelpout, 79, 491, 508–509 Ninespine Stickleback, 77, 374, 398, 399–401, 423 North Atlantic Cornerlantern, 77, 332 Northern Cutthroat Eel, 75, 210–211, 343 Northern Giant Seadevil, 77, 386–387 Northern Saillamp, 76, 329–330 Northern Sand Lance, 79, 372, 541–543 Northern Wolffish, 79, 452, 530–532 Oarjaw Wingmax, 237 Ocean Pout, 79, 473, 513–514 Omul, Baikal, 254 Orange Roughy, 77, 395–396 Paamiut Eelpout, 79, 497–499 Pacific Herring, 58, 64, 75, 166, 219, 221–223, 251, 254, 270, 275, 299, 300 Pacific Rainbow Smelt, 76, 244–245 Pacific Sand Lance, 79, 542, 543–545 Pacific Sleeper Shark, 75, 181, 183–184 Pale Eelpout, 79, 499–500 Pallid Sculpin, 78, 442–443 Patagonian Toothfish, 80 Pelican Gulper, 75, 216, 217–218 Penzhina Cisco, 254 Pink Eelpout, 78, 492–493 Pink Salmon, 76, 166, 264–267, 269, 270 Plaice, Alaska, 559 Plaice, American, 79, 368, 551–554, 558 Poacher, Atlantic, 78, 438–439 Polar Cod, 18, 34, 77, 358–360, 569 Polar Sculpin, 78, 440–442 Pollan, Irish, 254 Portuguese Shark, 75, 178–180 Pout Atlantic Soft, 79, 512 Aurora, 78, 476–478 Barsukov’s, 78, 473–474



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Knipowitsch’s, 78, 475–476 Ocean, 79, 473, 513–514 Twolip, 78, 474–475 Pouty Snailfish, 78, 468–469 Prickleback, Blackline, ix, 79, 515–516 Prickly Dreamer, 77, 385 Rabbitfish, Small-eyed, 171 Radiated Shanny, 79, 526–527 Rainbow Smelt, 21, 24, 76, 166, 246–247, 368, 374 Pacific, 76, 244–245 Rakery Lanternfish, 76, 327 Ratfish, Spotted, 172 Redfish Acadian, 77, 402–404, 553 Deepwater, 77, 317, 402, 403, 404–406 Golden, 77, 403, 405, 406–407, 531 Ribbed Sculpin, 78, 429, 431, 433–434 Richardson’s Snaggletooth, 76, 308 Rock Grenadier, 77, 226, 303, 338–340, 343 Rock Sole, 559 Rockling Fourbeard, 77, 351–353 Silver, 77, 353–354 Threebeard, 77, 355–356 Rough Sagre, 175 Roughhead Grenadier, 77, 211, 226, 242, 342–343, 458 Roughnose Grenadier, 77, 345–346 Roughy, Orange, 77, 395–396 Round Skate, 75, 199–201 Round Whitefish, 76, 277–279 Russet Grenadier, 77, 333–335 Saddled Eelpout, 79, 423, 496–497 Sadko Sculpin, 440 Saffron Cod, 21, 24, 77, 166, 242, 299, 365–367, 564 Sagre, Rough, 175 Saillamp, Northern, 76, 329–330 Salmon Atlantic, 28, 29, 38, 58, 63, 64, 65, 76, 166, 242, 246, 250, 279–282, 292, 321, 330, 361, 362, 542, 569, 570 Chinook, 76, 166, 266, 275–277 Chum, 76, 166, 267–270 Coho, 76, 266, 270–272, 276 Pink, 76, 166, 264–267, 269, 270 Sockeye, 76, 166, 266, 269, 270, 272–274 Sand Lance Northern, 79, 372, 541–543 Pacific, 79, 542, 543–545 Sawpalate, Stout, 75, 214–215 Scabbardfish, Black, 79, 348, 546 Scalebelly Eelpout, 79, 506–507

Sculpin Arctic, 78, 158, 408, 423, 424–426, 427 Arctic Hookear, 77, 412–413 Arctic Staghorn, 78, 413–415, 427, 428, 437 Atlantic Hookear, 77, 408–410 Bigeye, 78, 429, 431–432, 433 Fourhorn, 78, 244, 361, 362, 366, 372, 421–424, 427, 428, 496, 522, 544, 564 Giant Blob, 78, 444–445 Longhorn, 78, 419, 420–421, 513 Moustache, 78, 429–430, 431, 433 Pallid, 78, 442–443 Polar, 78, 440–442 Ribbed, 78, 429, 431, 433–434 Sadko, 440 Shorthorn, 78, 361, 362, 408, 413, 414, 423, 425, 426–429, 461, 478 Spatulate, 78, 415, 417–418, 428 Twohorn, 78, 415–417 Scutepout, Chevron, 79, 510–511 Sea Tadpole, 78, 343, 454, 456–458 Seadevil, Northern Giant, 77, 386–387 Searsid Bighead, 75, 228 Legless, 76, 232–233 Maul’s, 75, 229 Smallscale, 75, 230 Seasnail Atlantic, 78, 458–459 Black, 78, 466–467 Shanny Arctic, 79, 372, 524–526 Daubed, 79, 520–521 Radiated, 79, 526–527 Shark Deepsea Cat, 75, 174 Ghost, 171 Greenland, 8, 34, 38, 64, 75, 178, 179, 180–183, 184, 192, 242, 450, 509, 531, 533, 534, 553, 558, 570 Pacific Sleeper, 75, 181, 183–184 Portuguese, 75, 178–180 Sharpchin Barracudina, 76, 317, 321 Shortbeard Grenadier, 77, 335–336 Shorthorn Sculpin, 78, 361, 362, 408, 413, 414, 423, 425, 426–429, 461, 478 Shortjaw Cisco, 251 Shortspine Tapirfish, 75, 207–208 Shorttail Skate, 75, 189–190 Shulupaoluk, 78, 489–490 Silver Hatchetfish, Greater, 76, 306–307 Silver Rockling, 77, 353–354 Skate Abyssal, 75, 197–198, 199 Arctic, 38, 51 Barndoor, 513

I ndex 613

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Bigelow’s, 185 Darkbelly, 75, 186–188 Linen, 75, 201–202 Round, 75, 199–201 Shorttail, 75, 189–190 Smooth, 550 Soft, 75, 196–197 Spinytail, 75, 194–195 Thorny, 75, 190–192, 550, 553 Winter, 458 Sleeper Shark, Pacific, 75, 181, 183–184 Slender Eelblenny, 79, 372, 423, 522–523 Slender Snaggletooth, 76, 312–313 Slender Snipe Eel, 75, 212–213 Slickhead, Dusky, 76, 234 Small-eyed Rabbitfish, 171 Smallscale Searsid, 75, 230 Smelt Pacific Rainbow, 76, 244–245 Rainbow, 21, 24, 76, 166, 246–247, 368, 374 Smooth Flounder, 79, 564, 566–567 Smooth Lumpfish, 78, 446–447 Smooth Skate, 550 Smoothdream, Can-Opener, 77, 382–383 Smoothhead Bigeye, 76, 236 Madeiran, 76, 238 Manyray, 76, 235 Snaggletooth Large-Eye, 76, 309 Richardson’s, 76, 308 Slender, 76, 312–313 Snailfish Blacksnout, 78, 467–468 Dusty, 8 Gelatinous, 78, 460–462 Kelp, 78, 460, 464–465 Kido’s, 78, 454–455 Longfin, 78, 455–456 Nebulous, 8 Pouty, 78, 468–469 Threadfin, 78, 469–470 Variegated, 78, 462–463 Snakeblenny, 79, 523–524 Fourline, 79, 519–520, 525, 526 Snipe Eel, Slender, 75, 212–213 Snubnose Eel, 75, 209 Snubnosed Spiny Eel, 75, 206–207 Sockeye Salmon, 76, 166, 266, 269, 270, 272–274 Soft Pout, Atlantic, 79, 512 Soft Skate, 75, 196–197 Sole English, 561 Flathead, 555 Forkline, 561 Hybrid, 561 Rock, 559

614

Yellowfin, 559 Spark Anglemouth, 76, 304 Spatulate Sculpin, 78, 415, 417–418, 428 Spiny Dogfish, 163, 164, 175, 211, 224, 356, 364, 369, 550, 553, 558 Spiny Eel, Snubnosed, 75, 206–207 Spinytail Skate, 75, 194–195 Spotted Lanternfish, 76, 328–329 Spotted Ratfish, 172 Spotted Wolffish, 79, 530, 532, 534, 535–537 Staghorn Sculpin, Arctic, 78, 413–415, 427, 428, 437 Starry Flounder, 79, 166, 242, 561–563 Stickleback Ninespine, 77, 374, 398, 399–401, 423 Threespine, 77, 166, 372, 397–399, 400, 401, 423 Stoplight Loosejaw, 76, 311–312 Stout Eelblenny, 79, 517–518 Stout Sawpalate, 75, 214–215 Sturgeon, Lake, 32, 75, 203–205, 242, 544 Swallower, Hartel’s, 79, 539–540 Tadpole, Sea, 78, 343, 454, 456–458 Taillight Gulper, 75, 216, 217, 348 Tapirfish, Shortspine, 75, 207–208 Telescope, Arctic, 76, 331 Theologian Eelpout, 78, 483–484 Thorny Skate, 75, 190–192, 553 Threadfin Grenadier, 77, 340–341 Threadfin Snailfish, 78, 469–470 Threebeard Rockling, 77, 355–356 Threespine Stickleback, 77, 166, 372, 397–399, 400, 401, 423 Toothfish, Patagonian, 80 Trout Brook, 29, 57, 58, 64, 67, 76, 242, 246, 250, 285, 289–292, 423, 522, 544 Bull, 292 Lake, 19, 76, 166, 223, 246, 263, 278, 285, 297–299 Twohorn Sculpin, 78, 415–417 Twolip Pout, 78, 474–475 Variegated Snailfish, 78, 462–463 Veiled Anglemouth, 76, 302–303 Viperfish, Manylight, 76, 310–311 Warbonnet, Atlantic, 79, 518–519 Waryfish, Blackfin, 76, 316 White Barracudina, 76, 318–319 White Sea Eelpout, 79, 493–494 Whitefish Alaskan, 257 Broad, 58, 62, 65, 76, 166, 250, 257, 258, 260–262, 264, 295

Humpback, 257 Lake, 19, 57, 58, 76, 166, 257–259, 260, 299, 423, 522, 544 Round, 76, 277–279 Whiting, Blue, 77, 368, 375–376 Wingmax, Oarjaw, 237 Winter Skate, 458 Witch Flounder, 79, 549–551, 569 Wolf Eel Checkered, 78, 480 Common, 78, 482–483 Moray, 78, 481–482 Wolffish Atlantic, 79, 530, 532–534 Bering, ix, 79, 531, 537–538 Northern, 79, 452, 530–532 Spotted, 79, 530, 532, 534, 535–537 Yellowfin Sole, 559

French Common Names agone atlantique, 78, 438–439 aiguillat commun, 175 aiguillat noir, 75, 175–177 alépocéphale à grands yeux, 76, 236 alépocéphale de Madère, 76, 238 alépocéphale multirai, 76, 235 alépocéphale obscur, 76, 234 anguille à nez court, 75, 209 anguille égorgée bécue, 75, 210–211 antimore bleu, 77, 347–348 aphanope charbon, 79, 546 avaleur d’Hartel, 79, 539–540 avaleur feu-arrière, 75, 216 avocette ruban, 75, 212–213 baudroie d’Amérique, 77, 378–379 brosme, 77, 364–365 capelan, 76, 240–244 cariste barré, 78, 471–472 chaboisseau à dix-huit épines, 78, 420–421 chaboisseau à épines courtes, 78, 426–429 chaboisseau à quatre cornes, 78, 421–424 chaboisseau arctique, 78, 424–426 chaboisseau bronzé, 78, 419–420 chabot maculé, 78, 444–445 chauliode très-lumineux, 76, 310–311 chimère de profondeur, 75, 171–173 chimère d’Haeckel, 167 chimère-couteau, 75, 169–170 chimère-spatule, 75, 168–169 cisco arctique, 76, 254–256 cisco de lac, 76, 251–253 cisco sardinelle, 76, 262–264

corégone tschir, 76, 260–262 cotte blême, 78, 442–443 cotte de Sadko, 440 cotte polaire, 78, 440–442 cyclothone à petites dents, 76, 302–303 donzelle arctique, 77, 377 doux-rêve ouvre-boîte, 77, 382–383 dragon-boa, 76, 313–315 dragon-saumon à grands yeux, 76, 309 dragon-saumon de Richardson, 76, 308 dragon-saumon élancé, 76, 312–313 drague rouge-verte, 76, 311–312 épaule-criblée long nez, 76, 231 éperlan arc-en-ciel, 76, 246–247 éperlan du Pacifique, 76, 244–245 épinoche à neuf épines, 77, 399–401 épinoche à trois épines, 77, 397–399 esturgeon jaune, 75, 203–205 faux-trigle armé, 78, 429–430 faux-trigle aux grands yeux, 78, 431–432 faux-trigle bardé, 78, 433–434 flet étoilé, 79, 561–563 flétan atlantique, 79, 556–559 flétan du Groenland, 79, 568–571 football fine-lampe, 77, 380–381 garcette-goître, 75, 226–227 gonostome étincelé, 76, 304–305 grand corégone, 76, 257–259 grande argentine, 75, 224–225 grande hache d’argent, 76, 306–307 grande lycode, 78, 486–487 grandgousier pélican, 75, 217–218 grenadier à barbe courte, 77, 335–336 grenadier à barbillon court, 77, 336–337 grenadier à filaments, 77, 340–341 grenadier berglax, 77, 342–343 grenadier de Günther, 77, 337–338 grenadier de roche, 77, 338–340 grenadier du Grand Banc, 77, 344–345 grenadier roux, 77, 333–335 grenadier-scie, 77, 345–346 grosse poule de mer, 78, 449–450 guetteur à nageoire noire, 76, 316 gymnaste atlantique, 76, 239 hameçon atlantique, 77, 408–410 hameçon neigeux, 77, 412–413 hameçon rude, 77, 410–411 hareng atlantique, 75, 219–221 hareng du Pacifique, 75, 221–223 hoplostète orange, 77, 395–396

Index

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icèle à deux cornes, 78, 415–417 icèle spatulée, 78, 417–418 inconnu, 76, 300–301 laimargue atlantique, 75, 180–183 laimargue du Pacifique, 75, 183–184 lampe-voilière du nord, 76, 329–330 lamproie arctique, 75, 165–167 lançon du nord, 79, 541–543 lançon gourdeau, 79, 543–545 lanterne à joue pailletée, 76, 326 lanterne glaciaire, 76, 323–324 lanterne ponctuée, 76, 328–329 lanterne-bouée râtelière, 76, 327 lanterne-de-coin nord-atlantique, 77, 332 lanterne-joyau, 76, 324–325 lépidion à grands yeux, 77, 349–350 limace à filaments, 78, 469–470 limace à longues nageoires, 78, 455–456 limace à museau noir, 78, 467–468 limace atlantique, 78, 458–459 limace de Kido, 78, 454–455 limace des laminaires, 78, 464–465 limace gélatineuse, 78, 460–462 limace marbrée, 78, 462–463 limace nébuleuse, 8 limace noire, 78, 466–467 limace pote, 78, 468–469 limande carline, 79, 559–560 lompénie de Fabricius, 79, 522–523 lompénie naine, 79, 517–518 lompénie tachetée, 79, 520–521 lompénie-serpent, 79, 523–524 loquette d’Amérique, 79, 513–514 lotte, 77, 373–374 loup à tête large, 79, 530–532 loup atlantique, 79, 532–534 loup de Béring, 79, 537–538 loup tacheté, 79, 534–537 lussion à bec de canard, 76, 320 lussion à menton, 76, 321 lussion blanc, 76, 318–319 lycaspine à chevrons, 79, 510–511 lycode à arc, 79, 504–505 lycode à carreaux, 79, 509–510 lycode à deux lignes, 78, 487–488 lycode à oreilles, 79, 505–506 lycode à selles, 79, 496–497 lycode arctique, 79, 502–503 lycode commune, 78, 482–483 lycode d’Adolf, 78, 484–485 lycode de la mer Blanche, 79, 493–494 lycode de Laval, 78, 491–492 lycode de McAllister, 79, 495 lycode de Paamiut, 79, 497–499 lycode de Sars, 78, 483–484 lycode de Terre-Neuve, 79, 508–509 lycode glaciale, 78, 488–489 lycode murène, 78, 481–482

lycode pâle, 79, 499–500 lycode plume, 78, 489–490 lycode polaire, 79, 500–501 lycode quadrillée, 78, 480 lycode rose, 78, 492–493 lycode ventre-écaillé, 79, 506–507 maxailé aviron, 237 ménomini rond, 76, 277–279 merlan bleu, 77, 375–376 merluche à longues nageoires, 77, 356–357 mollasse atlantique, 79, 512 more délicat, 77, 348–349 morue franche, 77, 367–371 motelle à quatre barbillons, 77, 351–353 mustèle arctique à trois barbillons, 77, 355–356 mustèle argentée, 77, 353–354 myxine du nord, 75, 162–164 navaga jaune, 77, 365–367 ogac, 77, 371–373 ogre, 77, 392–394 omble chevalier, 67, 76, 283–289 omble de fontaine, 76, 289–292 omble malma, 76, 292–297 pailona, 75, 178–180 pêcheur à deux massettes, 77, 386–387 petite limace de mer, 78, 456–458 petite poule de mer arctique, 78, 450–452 petite poule de mer atlantique, 78, 452–453 petite poule de mer douce, 78, 446–447 petite poule de mer McAlpine, 78, 447–448 pharaon, 76, 317–318 plie à grande bouche, 8 plie arctique, 79, 564–566 plie canadienne, 79, 551–554 plie de Béring, 79, 555–556 plie grise, 79, 549–551 plie lisse, 79, 566–567 poisson-alligator arctique, 78, 436–438 poisson-alligator atlantique, 78, 435–436 quatre-lignes atlantique, 79, 519–520 raie à queue courte, 75, 189–190 raie à queue épineuse, 75, 194–195 raie bathyale, 75, 197–199 raie boréale, 75, 186–188 raie de Bigelow, 185 raie épineuse, 75, 190–192



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raie linon, 75, 201–202 raie molle, 75, 196–197 raie ronde, 75, 199–201 rêveur piquant, 77, 385 roussette de profondeur, 75, 174 sagre rude, 175 saïda franc, 77, 360–363 saïda imberbe, 77, 358–360 saumon atlantique, 76, 279–282 saumon chinook, 76, 275–277 saumon coho, 76, 270–272 saumon kéta, 76, 267–270 saumon rose, 76, 264–267 saumon rouge, 76, 272–274 sébaste acadien, 77, 402–404 sébaste atlantique, 77, 404–406 sébaste orangé, 77, 406–407 serrivomer trapu, 75, 214–215 sigouine rubanée, 79, 528–529 stichée arctique, 79, 524–526 stromatée à fossettes, 79, 547–548 tact géant de Vanhoeffen, 77, 388–389 tapir à grandes écailles, 75, 206–207 tapir à petites épines, 75, 207–208 télescope arctique, 76, 331 terrassier à six lignes, 79, 515–516 tête-à-crète robuste, 77, 390–391 touladi, 76, 297–299 toupet marbré, 79, 518–519 tricorne arctique, 78, 413–415 tube-épaule apode, 76, 232–233 tube-épaule de Maul, 75, 229 tube-épaule grosse tête, 75, 228 tube-épaule petites écailles, 75, 230 ulvaire deux-lignes, 79, 526–527 unernak à deux lèvres, 78, 474–475 unernak aurore, 78, 476–478 unernak caméléon, 78, 478–479 unernak de Barsukov, 78, 473–474 unernak de Knipowitsch, 78, 475–476

Indigenous Names Aalisangar, 367 Aanaaksiiq, 260 Aanaaliq, 260, 277 Aanaarliq, 260 Aanak, 289 Aaqaksaaq, 537 Adglernaq, 191 Akkulliakitsok, 524 Akoak, 530 Akoaksaluk, 530 Akuhaauk, 502 Amaatuuq, 264

Amagiak, 240, 254, 262 Amajak, 541 Amerikap Suluppaagaa, 402 Âna, 289 Anâtlik, 289 Anaakłiiq, 260 Anaakliq, 260 Anadleq, 257 Anâdlerk, 257 Anahik, 257 Anah’lih’, 260 Anahluk, 277 Anakheek, 257 Anaklek, 260 Anaklik, 260 Ananaaklik, 260 Anaqkik, 260 Anaqklik, 260 An-ark-hlirk, 260 Angmaggeuck, 240 Angmagiak, 240 Angmalook, 283 Angnaklin, 260 Angusatdluk, 449 Aniak, 283 Aniaq, 67, 283 Anma, 254, 262 Anmagiak, 251, 254 Anmagiak Piqquaqtitaq, 254 Anmaglak, 262 Ânna, 289 Anokik, 289 Anuk, 289 Aopalayâk, 283 Aoparktulâyoq, 283 Armagiak, 251, 262 Arnaqsleq, 251 Arnardluk, 449 Atihkamàkw, 257 Atihkamekw, 257 Atlantikup Kanajua, 408 Aupalijaat, 283 Axmagiaq, 240 Chehluk, 373 Col-lic-puk, 297 Coogjannernak, 478 Dalts’an, 257 Dalts’in, 257 Dhik’ii, 292 Eekallûk, 283 Eekalook, 283 Eetooknuit, 244 Ehohok, 297 Ehok, 297 Ekalluak, 360 Ekalluk, 67, 283 Ekaludjuaq, 180 Ekalugak, 360 Ekalugssûp Piarâ, 180

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Ekalukpik, 283 Ekalupik, 283 Ekaluppik, 283 Eqalugaq, 360 Eqaluk, 283 Eqalukjuaq, 180 Eqalussuaq, 180 Eqalussuaq Qernertoq, 175 Equaluaq, 360 Equaluk, 283 Erlakukpik, 283 Evitaruk, 283, 292 Geo Sahba, 267 Higaq, 300 Higayuriaq, 297 Hiwiterro, 283 Holili-gah, 240 Hutdaun, 522 Idlôk, 297 Ihkaluk, 283 Ihok, 297 Iituuq, 221 Ikalopik, 283 Ikalukpik, 67, 283 Ilhuagnik, 244 Iloruleqqortooq, 364 Iluuraq, 67, 297 Imaluunniit, 375 Imaviup Kanajua, 412 Imingoak, 342 Ingmingoak, 342 Ingminniset, 342 Iqaluakpak, 283, 292, 297 Iqaluarak, 244 Iqalugruaq, 267 Iqalujjuaq, 180 Iqaluk, 283 Iqalukpiaryuk, 283 Iqalukpiik, 283 Iqalukpiit, 283 Iqalukpik, 283 Iqalukuak, 180 Iqaluppik, 283 Iqaluqpik, 292 Iqaluruaq, 267 Iqalusaaq, 262 Iqaluupik, 283 Iriqpaligaurat, 262 Irkaluk, 283 Isiuralittaaq, 297 Issittup Tarraleqisaava, 186 Itok, 360 Ivatarak, 67, 283 Ivik, 162 Ivisaruk, 283 Ivitaaruq, 283, 292 Ivitagok, 283 Ivitaroq, 283 Ivitaruk, 283

616

Jikuktok, 257 Kaïtilik, 283 Kakidlautidlik, 399 Kakilahaq, 399 Kakilasak, 399 Kakilaychok, 397 Kakilisak, 397 Kakilishek, 397, 399 Kakilusuk, 397, 399 Kakiva, 399 Kakiviaktok, 257 Kakiviartût, 257 Kakkiviartoq, 257 Kaktak, 254 Kaleralik, 568 Kaloarpok, 67, 283 Kalushak, 262 Kanajoq Kapinartulik Nalinginnaq, 415 Kanajoq Nalinginnaq, 426 Kanajorlak, 438 Kanajuk, 421, 424, 426 Kanatik, 194 Kanayuk, 408, 421, 424, 426 Kanayuq, 421 Kaneeok, 421 Kan-ny-yoke, 421 Kanyok, 421 Kapahilik, 262 Kapihilik, 257 Kapihillik, 254 Kapisalik, 323 Kapisalingoak, 323 Kapisilik, 219, 251, 254, 257, 277, 323 Kaporniangaq, 283 Kashkanamesh, 240 Katilautik, 397 Kausriluk, 260 Kavasilik, 260 Kaviselik, 251, 257 Kavisilak, 219 Kavisilaq, 221 Kavisilik, 251, 257, 260, 279 Keblernak, 323 Keki-yuak-tuk, 257 Kipinartulik, 397 Ko le le kuk, 240 Koupjhaun-ohuk, 478 Kraaktak, 254, 262 Krolleliprark, 221 Kugrauna, 496 Kugsaunak, 478, 528 Kùkamàs, 297 Kùkamàsh, 297 Kùkamàw, 297 Kùkamesh, 297 Kukilik, 175 Kukumes, 297 Kuliligak, 240 Kumaliq, 279

Kurksaunak, 528 Kuxrauna, 496 Lepisuk, 449 Łuk dagaii, 260 Łuk digaii, 260 Łuk zheii, 260 Man-iktoe, 452 Màsimàkus, 289 Màsimàkush, 289 Màsimekush, 289 Màsimekw, 289 Matameku, 289 Mikiapic Kapisilik, 323 Nadalna, 555, 559 Najorpilik, 530 Namekush, 297 Nameo, 203 Namew, 203 Nataagnaq, 564 Nataarnaq, 556, 561, 564 Nataaznak, 561, 564 Natagnak, 399 Nat-ah-nuh, 568 Natanak, 555, 559 Natangnak, 561 Natarinaq, 564 Natarnak, 555, 568 Natarnaq, 555 Natarrnak, 555, 568 Nätarrnaq, 373 Nee-fitz-shak, 464 Nemeo, 203 Nemeryaq, 165 Nemew, 203 Netarnârak, 568 Niialingaq, 180 Nimaw, 203 Nipisa, 449 Nipi-sak, 464 Nipisarluk, 452 Nipishah, 464 Nulilighuk, 240 Nû-mug-û-shûk, 165 Nutilliajuk, 283 Nutilliq, 67, 283, 289 Nùtimĭwasù, 251 Nùtimĭweshish, 251 Nùtimĭwesù, 251 Ôarsuk, 371 Ogâ tsuk, 360 Ogac, 242, 319, 360, 367, 371, 423, 425, 427, 522, 525, 542, 544 Ogak, 365, 371 Ogaq, 360 Ogavik, 365 Oggak, 371 Oggaksuk, 371 Okeugnak, 277

Okôtak, 551 Oquutaq, 551 Osungnak, 277 Ovac, 360 Ovak, 367, 371 O-wuk, 371 Paiirluq, 267 Panmagrik, 240 Panmaksraq, 240 Pikoaktik, 221 Pi-kok-tok, 257 Pikuktung, 257 Pikuktuq, 257 Pikuktuuq, 257 Pingasunik, 397 Piqquaqtitaq, 221, 254 Pirkroartitak, 221 Putorugtoq, 541 Putorutorsôak, 321 Puyyaqiaq, 564 Qaaktak, 262 Qaaktaq, 221, 254 Qaantaq, 221 Qaatag, 254 Qaleralik, 568 Qaluaqpak, 283 Qaluaqpaq, 292 Qaluarak, 244 Qaluhaq, 221, 251, 254, 262 Qalukpik, 283, 292 Qalupiaq, 257 Qanirkuutuk, 426 Qanktaq, 254, 262 Qanktaq Siirgarq, 254 Qaqtak, 262 Qarlêk, 191 Qeeraaraq, 532 Qeeraasaq, 530 Qeeraq, 534 Qeerngaq Milagulaar, 534 Qelaluqaq, 257 Qoliiligaq, 240 Qorkshuyoq, 449 Qulilirraq, 240 Quvssaunaq, 528 Saama, 279 Saamakutaak, 279 Saamarug, 279 Saarullik, 367 Sahti Eda, 275 Saraudlik, 367 Saraudlirksoak, 367 Sârugdligaraq, 367 Sârugdlik, 367 Sâugdlik, 367 Savigunnaq, 277 Saviliursak, 353 Shalup-pau-gah, 517 Shii, 267

Index

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Shruh, 300 Shryuh, 300 Shulukpaoluk, 373 Shushashu, 283 Si-airryuk, 300 Sierak, 300 Sigaq, 300 Siiraq, 300 Siirgarq, 254, 300 Singayuriaq, 297 Siuryuktuuq, 365 Skalugsuak, 180, 182 Sruh, 300 Sulupavak, 502 Suluppaagaq Angisooq, 406 Suluppaagaq Itisoormiu, 404 Sùsàsù, 283 Suvaliviniq, 283 Tadlulik, 283 Tahiqmi Ihugaqyut, 297 Taqqalerisaaq, 194 Tarjaxfaq, 275 Teirark, 300 Tejernak, 522 Tiktaaliq, 373 Tiktabek, 373 Tiktailik, 373 Tiktalaq, 373 Tiktalerk, 300 Tiktalik, 373 Tinguttooq, 364 Tisuajuk, 283 Titaalirq, 373 Titale, 373 Titaliq, 373 Tittaalik, 373 Tivaqiq, 424 Treeluk, 254, 262 Tungujortoq, 530 Tupissut, 342 Uchùlipĭsh, 251 Ugak, 367, 371 Unernak, 473–476, 478 Uqsruqtuuq, 221 Utoqulaaq, 530 Utshashumek, 279 Utùlipĭ, 251 Utùlipĭsh, 251 Uugaatsuk, 371 Uugak, 367 Uugaq, 360, 371 Uugavik, 358, 360, 371 Uugayak, 371 Uuqaq, 365 Ûvak, 360, 371 Vit, 297

English and Latin Family Names Acipenseridae, 3, 4, 6, 75, 85, 93, 203–205. See also Sturgeons Agonidae, 7, 78, 82, 102, 108, 435–439 Alepisauridae, 6, 81 Alepocephalidae, 6, 76, 81, 105, 108–109, 233–239, 339 Ammodytidae, 7, 79, 83, 103, 109, 281, 404, 541–545. See also Sand Lances Anarhichadidae, 4, 7, 79, 103, 110, 473, 530–538. See also Wolffishes Anguillidae, 6, 81 Anoplogastridae, 7, 77, 102, 392–394 Argentines, 75, 104, 224–225 Argentinidae, 6, 75, 104, 224–225 Barbourisiidae, 7, 82 Barracudinas, 76, 104, 129, 317–321, 339, 343, 569. See also Paralepididae Black Swallowers, 79, 96, 539-540. See also Chiasmodontidae Bristlemouths, 76, 103, 120, 302–305, 339, 356. See also Gonostomatidae Butterfishes, 79, 98, 547-548. See also Stromateidae Bythitidae, 6, 77, 101, 376–377 Caristiidae, 7, 78, 82, 99, 471–472 Cat Sharks, 75, 93, 173–174. See also Scyliorhinidae Catostomidae, 85 Caulophrynidae, 6, 82 Ceratiidae, 7, 77, 82, 94, 386–387, 388 Cetomimidae, 7, 82 Cetorhinidae, 6, 81 Chiasmodontidae, 7, 79, 96, 539–540 Chimaeridae, 4, 6, 75, 81, 96, 171–173. See also Shortnose Chimaeras Clupeidae, 4, 6, 64, 75, 105, 111–112, 219–223, 358. See also Herrings Codlings, 77, 106, 125–126, 346–350. See also Moridae Cods, 77, 106, 117–119, 317, 323, 346, 358–376, 421. See also Gadidae Cottidae, ix, 3, 4, 7, 51, 64, 77–78, 82, 85, 107, 111–116, 339, 407–434. See also Sculpins Cryptacanthodidae, 80 Cutlassfishes, 79, 99, 545–546. See also Trichiuridae Cutthroat Eels, 75, 98, 148, 208–211. See also Synaphobranchidae Cyclopteridae, 7, 64, 78, 82, 97, 116–117, 446–453 Cyprinidae, 85



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Dalatiidae, 175, 178 Deep-sea Spiny Eels, 75, 100, 128, 205–208. See also Notacanthidae Diretmidae, 7, 82 Dragonfishes, 76, 102, 147–148, 307–315, 339. See also Stomiidae Dreamers, 77, 95, 128, 382–385. See also Oneirodidae Eelpouts, ix, 38, 78, 101, 149–155, 242, 244, 343, 368, 423, 427, 473–514. See also Zoarcidae Esocidae, 85 Etmopteridae, 4, 6, 75, 81, 93, 175–177, 178 Eurypharyngidae, 6, 75, 95, 217–218 Fathead Sculpins, 78, 107, 135, 440– 445. See also Psychrolutidae Footballfishes, 77, 95, 380–381. See also Himantolophidae Gadidae, 3, 4, 6, 64, 77, 82, 106, 117–119, 281, 351, 353, 355, 358–376. See also Cods Gasterosteidae, 3, 4, 7, 77, 85, 101, 119, 396–401. See also Sticklebacks Gigantactinidae, 7, 77, 94, 388–389 Gonostomatidae, 6, 39, 76, 81, 103, 120, 302–305, 338. See also Bristlemouths Goosefishes, 77, 94, 378–379. See also Lophiidae Grenadiers, 38, 65, 77, 106, 123–125, 333–346, 368, 531, 558. See also Macrouridae Gulpers, 75, 95, 217–218. See also Eurypharyngidae Gunnels, 79, 107, 515, 528–529. See also Pholidae Hagfishes, 3, 75, 92, 162–164. See also Myxinidae Halosauridae, 6, 81 Hemitripteridae, 7, 82 Herrings, 68, 75, 105, 111, 219–223, 270, 300, 358, 421, 427. See also Clupeidae Hexagrammidae, 7, 82 Himantolophidae, 6, 77, 95, 380–381 Hiodontidae, 85 Lamnidae, 6, 81 Lampreys, 3, 75, 92, 164–167, 203, 204, 222, 258, 261, 265, 268, 270, 300. See also Petromyzontidae Lampridae, 6, 81 Lantern Sharks, 75, 93, 175–177. See also Etmopteridae

Lanternfishes, 76–77, 103, 126–127, 310, 322–332, 339, 356, 368, 375. See also Myctophidae Linophrynidae, 7, 82 Liparidae, ix, 3, 4, 7, 23, 68, 78, 82, 97, 120–123, 454–470, 568, 569. See also Snailfishes Longnose Chimaeras, 75, 96, 138, 167–170, 171. See also Rhinochimaeridae Lophiidae, 6, 77, 82, 94, 378–379 Lumpfishes, 78, 97, 116–119, 446–453. See also Cyclopteridae Macrouridae, 4, 6, 77, 81, 106, 123–125, 333–346. See also Grenadiers Manefishes, 78, 99, 471–472. See also Caristiidae Marine Hatchetfishes, 76, 101, 305–307, 356. See also Sternoptychidae Melamphaidae, 7, 77, 82, 100, 390–391 Melanocetidae, 6, 82 Microstomatidae, 6, 75, 81, 104, 226–227 Moridae, 4, 6, 77, 106, 125–126, 346–350 Myctophidae, 6, 69, 76–77, 80, 81, 103, 126–127, 322–332, 339, 404. See also Lanternfishes Myxinidae, 4, 6, 75, 81, 92, 162–164. See also Hagfishes Nemichthyidae, 6, 75, 81, 98, 211–213 Notacanthidae, 6, 75, 100, 128, 205–208 Notosudidae, 6, 76, 104, 315–316 Nototheniidae, 7, 83 Ogrefishes, 77, 102, 392–394. See also Anoplogastridae Oneirodidae, 4, 6, 77, 82, 95, 128, 382–385, 388 Oreosomatidae, 7, 82 Osmeridae, 3, 4, 6, 76, 85, 105, 129, 240–247, 404. See also Smelts Paralepididae, 6, 76, 104, 129, 317–321, 339, 404. See also Barracudinas Pencilsmelts, 75, 104, 226–227. See also Microstomatidae Percidae, 85 Percopsidae, 85 Petromyzontidae, 3, 4, 6, 75, 81, 85, 92, 164–167. See also Lampreys Pholidae, 7, 79, 83, 107, 515, 528–529 Phosichthyidae, 6, 81 Phycid Hakes, 77, 106, 130–131, 351–357. See also Phycidae

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Phycidae, 6, 77, 81, 106, 130–131, 351–357 Platytroctidae, 6, 75–76, 81, 97, 131–132, 227–233, 339 Pleuronectidae, 4, 5, 7, 52, 64, 79, 83, 96, 132–134, 404, 549–571 Poachers, 78, 102, 108, 435–439. See also Agonidae Pricklebacks, 79, 107, 145–147, 368, 515–527, 541. See also Stichaeidae Psychrolutidae, 7, 78, 107, 135, 440–445 Rajidae, 4, 6, 75, 81, 92, 135–138, 178, 185–202. See also Skates Rhinochimaeridae, 4, 6, 75, 81, 96, 138, 167–170. See also Longnose Chimaeras Ridgeheads, 77, 100, 390–391. See also Melamphaidae Righteye Flounders, 79, 96, 132–134, 549–571. See also Pleuronectidae Rondeletiidae, 7, 82 Saccopharyngidae, 6, 75, 95, 215–216 Salmonidae, 3, 4, 6, 15, 43, 63, 67, 74, 76, 81, 85, 105, 139–143, 240, 248–301 Sand Lances, 79, 103, 109, 192, 242, 270, 275, 281, 286, 291, 368, 414, 419, 421, 423, 449, 541–545, 569. See also Ammodytidae Sawpalates, 75, 99, 213–215. See also Serrivomeridae Sciaenidae, 85 Scombridae, 7, 83 Scorpaenidae, 4, 5, 7, 77, 82, 107, 144, 401–407 Scorpionfishes, 77, 107, 144, 401–407. See also Scorpaenidae Sculpins, 25, 27, 32, 48, 58, 64, 77, 107, 111–116, 158, 192, 242, 270, 273, 286, 291, 295, 300, 339, 343, 364, 368, 374, 407–434, 528, 541, 558. See also Cottidae Scyliorhinidae, 4, 6, 75, 81, 93, 173–174 Seadevils, 77, 94, 386–387. See also Ceratiidae Serrivomeridae, 6, 75, 99, 213–215, 339 Shortnose Chimaeras, 75, 96, 167, 171–173. See also Chimaeridae Skates, 3, 65, 68–69, 75, 92, 135–138, 181, 185–202, 526, 533, 558, 559. See also Rajidae Sleeper Sharks, 75, 93, 144–145, 178–184. See also Somniosidae

618

Slickheads, 76, 105, 108–109, 233–239, 339. See also Alepocephalidae Slimeheads, 77, 100, 394–396. See also Trachichthyidae Smelts, 76, 105, 129, 240–247, 275, 372, 421, 423. See also Osmeridae Snailfishes, 78, 97, 120–123, 368, 427, 446, 454–470. See also Liparidae Snipe Eels, 75, 98, 211–213. See also Nemichthyidae Somniosidae, 4, 6, 75, 93, 144–145, 178–184 Squalidae, 6, 81, 176 Sternoptychidae, 6, 76, 81, 101, 305–307. See also Marine Hatchetfishes Stichaeidae, 4, 7, 79, 83, 107, 145–147, 515–527. See also Pricklebacks Sticklebacks, 32, 77, 101, 119, 244, 368, 396–401, 419. See also Gasterosteidae Stomiidae, 6, 76, 81, 102, 147–148, 307–315, 339, 404 Stromateidae, 7, 79, 98, 547–548 Sturgeons, 3, 75, 93, 163, 203–205. See also Acipenseridae Swallowers, 75, 95, 215–216. See also Saccopharyngidae Synaphobranchidae, 4, 6, 75, 81, 98, 148, 208–211 Syngnathidae, 7, 82 Synodontidae, 6, 81 Trachichthyidae, 7, 77, 100, 394–396 Trachipteridae, 6, 81 Trichiuridae, 7, 79, 99, 545–546 Trouts and Salmons, 74, 76, 105, 139–143, 240, 248–301. See also Salmonidae Tubeshoulders, 75–76, 97, 131–132, 227–233, 339. See also Platytroctidae Viviparous Brotulas, 77, 101, 376–377. See also Bythitidae Waryfishes, 76, 104, 315–316. See also Notosudidae Whipnoses, 77, 94, 388–389. See also Gigantactinidae Wolffishes, 79, 103, 110, 530–538, 558. See also Anarhichadidae Zoarcidae, ix, 3, 4, 7, 8, 51, 78–79, 80, 83, 101, 149–155, 473–514. See also Eelpouts

French Family Names Alépocéphales, 76, 233–239 Anguilles égorgées, 75, 208–211 Argentines, 75, 224–225 Avaleurs, 75, 215–216 Baudroies, 77, 378–379 Caristes, 78, 471–472 Chabots, 77, 407–434 Chabots veloutés, 78, 440–445 Chimères, 75, 171–172 Chimères à long nez, 75, 167–170 Circés, 75, 227–233 Cyclothones, 76, 302–305

Rêveurs, 77, 382–385 Roussettes, 75, 173–174 Sabres de mer, 79, 545–546 Scorpènes, 77, 401–407 Serrivomers, 75, 213–215 Sigouines, 79, 528–529 Somniosidés, 75, 178–184 Stichées, 79, 515–527 Stromatées, 79, 547–548 Tacts géants, 77, 388–389 Truites et Saumons, 76, 248–301

Donzelles vivipares, 77, 376–377 Dragons à écailles, 76, 307–315 Éperlans, 76, 240–247 Épinoches, 77, 396–401 Esturgeons, 75, 203–205 Grandgousiers, 75, 217–218 Grands avaleurs, 79, 539–540 Grenadiers, 77, 333–346 Guetteurs, 76, 315–316 Haches d’argent, 76, 305–307 Harengs, 75, 219–223 Hoplites, 77, 394–396 Lamproies, 75, 164–167 Lançons, 79, 541–545 Limaces de mer, 78, 454–470 Lussions, 76, 317–321 Lycodes, 78–79, 473–514 Microbecs, 75, 226–227 Moros, 77, 346–350 Morues, 77, 358–376 Myxines, 75, 162–164 Ogres, 77, 392–394 Phycidés, 77, 351–357 Plies, 79, 549–571 Poissons-alligators, 78, 435–439 Poissons-avocettes, 75, 211–213 Poissons-football, 77, 380–381 Poissons-heaumes, 77, 390–391 Poissons-lanternes, 76–77, 322–332 Poissons-loups, 79, 530–538 Poissons-pêcheurs, 77, 386–387 Poissons-tapirs à épines, 75, 205–208 Poules de mer, 78, 446–453 Raies, 75, 185–202 Requins-lanternes, 75, 175–177

Index

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