THE ARCTIC SEAS CI imatology, Oceanography, Geology, and Biology

Edited by

Yvonne Herman

IOm51 VAN NOSTRAND REINHOLD COMPANY ~ -----New York This work relates to Department of the Navy Grant NOOOI4-85- G-0252 issued by the Office of Naval Research. The United States Government has a royalty-free license throughout the world in all copyrightable material contained herein. Copyright © 1989 by Van Nostrand Reinhold Softcover reprint of the hardcover 1st edition 1989

Library of Congress Catalog Card Number 88-33800 ISBN-13 :978-1-4612-8022-4 e-ISBN-13: 978-1-4613-0677-1 DOI: 10.1007/978-1-4613-0677-1

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Library of Congress Cataloging in Publication Data The Arctic Seas. Includes index. 1. Oceanography-Arctic Ocean. 2. Geology-ArctiC Ocean. 1. Herman, Yvonne. GC401.A76 1989 551.46'8 88-33800 ISBN-13: 978-1-4612-8022-4 For Anyu Contents

Preface / vii Contributors / ix

1. The Present Climate of the Arctic Ocean and Possible Past and Future States 1 R. G. Barry 2. Arctic Ice-Ocean Dynamics 47 W. D. Hibler III 3. Chemical Oceanography of the Arctic Ocean 93 L. Anderson and D. Dyrssen 4. Polar Marine Ecosystem Evolution 115 M. J. Dunbar 5. Arctic Sea-Ice Biota 123 R. A. Horner 6. Primary Production, Chlorophyll, Light, and Nutrients Beneath the Arctic Sea Ice 147 O. G. N. Andersen 7. Arctic Ocean Phytoplankton 193 B. R. Heimdal 8. Foraminifera and Pteropoda Beneath the Arctic Sea Ice: New Distributions 223 Y. Herman and O. G. N. Andersen 9. Ecology of Arctic Ocean Cryopelagic Fauna 235 I. A. Mel'nikov 10. Evolution of Arctic Ecosystems During the Neogene Period 257 A. N. Golikov and o. A. Scarlato 11. Distributional Patterns of Echinoderms in the Eurasian Sector of the Arctic Ocean 281 N. A. Anisimova 12. Marine Bivalvia of the Arctic Ocean 303 V. V. Fedyakov and A. D. Naumov

13. Arctic Ocean Gastropod Prosobranchs 325 A. N. Golikov 14. Arctic Ocean Bryozoa 341 V. I. Gontar and N. V. Denisenko 15. Arctic Ocean Mysids (Crustacea, ): Evolution, Composition, and Distribution 373 V. V. Petryashov

v vi Contents

16. Hydrozoa of the Eurasian Arctic Seas 397 S. D. Stepanjants 17. Arctic Ocean Cumacea 431 S. V. Vassilenko 18. Quarternary Calcareous Nannofossil Biostratigraphy: The Eastern Arctic Ocean Record 445 C. Card 19. Arctic Ocean Radiolarians 461 S. B. Kruglikova 20. Diatoms in Arctic Shallow Seas Sediments 481 E. I. Polyakova 21. Ecology of Recent Foraminifera on the Canadian Continental Shelf of the Arctic Ocean 497 C. Vilks 22. Thorium and Uranium Isotopes in Arctic Sediments 571 B. L. K. Somayajulu, P. Sharma, and Y. Herman 23. Late Neogene Arctic Paleoceanography: Micropaleontology, Stable Isotopes, and Chronology 581 Y. Herman, f. K. Osmond, and B. L. K. Somayajulu 24. Sediment Composition and Sedimentary Processes in the Arctic Ocean 657 D. A. Darby, A. S. Naidu, T. C. Mowatt, and C. A. Jones 25. Late Cenozoic Stratigraphy and Paleoceanography of the Barents Sea 721 V. S. Zarkhidze and Yu. C. Samoilovich 26. The Last Glaciation of Eurasia 729 A. A. Velitchko, L. L. Isayeva, D. B. Oreshkin, and M. A. Faustova 27. Geological and Paleoclimatic Evolution of the Arctic During Late Cenozoic Time 759 I. D. Danilov 28. Organic Geochemistry of Barents Sea Sediments 761 A. N. Belyaeva, A. 1. Daniushevskaya, and E. A. Romankevich 29. Physiography and Bathymetry of the Arctic Ocean Seafloor 797 J. R. Weber 30. Tectonic History of the Arctic Region from the Ordovician Through the Cretaceous 829 L. P. Zonenshain and L. M. Napatov

Index / 863 15: Arctic Ocean Mysids (Crustacea, Mysidacea): Evolution, Composition, and Distribution

V. v. Petryashov

INTRODUCTION ment of present-day water circulation in the Northern Hemisphere. Forty of mysids belonging to 20 genera, 2 families, and 1 suborder have been described from the Arctic Ocean. PREVIOUS STUDIES These species belong to 5 biogeographic groups. There is a decline in the number Studies of the Arctic Ocean Mysidacea (re• of species northward and eastward of the stricted to the area north of the Wyville• adjacent Atlantic boreal waters (from 27 to Thomson Ridge, the southern coast of the 4 or 3 species) and a slight increase in the Baffin Bay, the eastern part of the Hudson area of mixing of Arctic and Pacific waters Strait, and the Bering Strait) commenced (up to 5 species). Atlantic boreal and am• in the first half of the nineteenth century. phiboreal species are predominant in the Ross (1835) mentioned (= Mysis) Norwegian Sea, the southwestern part of flexuosus in his work on marine inverte• the Barents Sea, and the Greenland Sea; brates collected during his second voyage Pacific boreal species dominate in the to the Arctic regions. Further studies of Chukchi Sea off the Alaskan coast, and bo• mysids from this area were carried out in real-Arctic and Arctic species dominate in the second half of the nineteenth century other areas of the Arctic Ocean. Species by Walker (1862), Jarzynsky (1870), Sars that belong to different biogeographic (1870), and Miers (1877). These investiga• groups occur in specific water masses. The tions dealing with mysids collected mainly distributional patterns of the recent mysid in the Norwegian, Barents, and White seas fauna were established at the onset of the were followed by a study of Mysidacea off Postglacial transgression with the develop- the northwestern and northern coast of Alaska (Murdoch, 1885a,b). Simultane• Note: This chapter has been reviewed, edited, and re• ously intensive studies of mysids from written by the Editor, Yvonne Herman. the Norwegian, Greenland, and Barents

373 374 V V Pe~yashov seas as well as from the Baffin Bay com• DISTRIBUTIONAL PATTERNS menced (Sars, 1885, 1886; Hansen, 1888, 1908; Aurivillius, 1896; Vanhoffen, 1897; Our results and literature data indicate Birula, 1897; Stebbing, 1900; Ohlin, 1901; that 40 species of mysids inhabit the Arctic Gerstaecker and Ortmann, 1901; Ort• Ocean; they belong to 1 suborder, 2 famil• mann, 1901). ies, and 20 genera. The number of species The results allowed Zimmer (1904, 1909) inhabiting the Arctic Ocean constitute 5 to draw the first conclusions concerning percent of the world fauna. the Arctic Mysidacea based on his study of Changes in species composition in dif• 32 species from the Arctic Ocean within ferent sectors of the Arctic Ocean were the geographical limits mentioned above. analyzed. The degree of similarity of fau• Studies by Linko (1907, 1908) off the north• nas in these areas was determined using ern Russian coast and by Stephensen the Chekanovsky-S0rensen index (S0- (1912, 1913, 1918, 1933, 1938) in the rensen, 1948): seas surrounding Greenland were subse• quently published. Investigations by Paul• 2' C' 100% IC5 = ----• son (1909), Kramp (1913), Bjorck (1916), D1 + D2 Schmitt (1919), Madsen (1936), Bertelsen (1937), Saemundsen (1937), and Dunbar and the degree of similarity of faunas was (1940) dealing with mysids or coastal determined using the Jaccard index (Jac• plankton containing Mysidacea from Ice• card, 1901): land, Greenland, and the North American coast should also be mentioned. During C·100% the last decades the Arctic Ocean mysids IJ = ----- D1 + D2 - C were studied by Holmquist (1949, 1958, 1959, 1963, 1982), Rusanova (1962, 1963, For the degree of inclusion of fauna of one 1971), and Kulikov (1980). In addition to area into the fauna of another region the the study of Arctic species, the investiga• index of Shimkevich (Simpson, 1943) was tion of this group in other regions was un• used: derway (Czerniavsky, 1887; Holt and Tat• tersall, 1905; Illig, 1930; Tattersall, 1951; C'100% Mauchline, 1980; Mauchline and Murano, 15z5 = D· 1977). However, to date the study of the mm Central Arctic Basin mysids has been inad• For these three equations, C is the number equate. of species in common, D1 and D2 are the numbers of species in the areas compared, and Dmin is the minimum number of spe• MATERIAL AND METHODS cies in the areas compared (Dmin :5 Dm.x). The results of our calculations are given The results presented in this chapter are in Table 15-1. The data obtained indicate based on our studies of Mysidacea from a rapid decline in the number of species the collection of the Zoological Institute of northward and eastward from the adjacent the USSR Academy of Sciences (710 sam• Atlantic boreal waters (from 27 to 4 to 3 ples containing 5,500 specimens were ana• species) and a slight increase in the area of lyzed). Most of these samples were col• mixing of Arctic and Pacific waters (up to lected with a Sigsbee dredge and a few 5 species). The mysids of the Norwegian were collected with a Djedy net. and southern regions of the Barents Sea Arctic Ocean Mysids 375 are of boreal character and differ from their II. Atlantic boreal species counterparts in other regions. The fauna of this area has close affinities with faunas of 1. Upper sublittoral ubiquitous boreal spe• the coast of eastern Greenland and the cies: , Praunus flexuosus, southern Baffin Bay (as far as 75°NL); P. inermis (Plate 15-1, b), and Schistomysis there is a great similarity between the fau• ornata, present in the temperate waters nas of these areas and the mysids from the of the North Atlantic from the English continental slope off the Central Arctic Ba• Channel in the south to Murman and sin from Spitzbergen and up to the north• the southern Novaya Zemlya Island to ern Laptev Sea. The fauna of most of the the north (Zimmer, 1904, 1909; Linko, remaining area of the coastal Arctic region 1908). is rather monotonous. The only exception 2. Elittoral*-upper bathyal ubiquitous bo• is the Chukchi Sea: Its northwestern re• real species: Erythrops microps, E. serrata, gions support a fauna similar to that of the Pseudomma affine, Mysideis insignis, Mys• adjacent East Siberian Sea. Off of the idopsis didelphys, and Hemimysis abyssi• Alaskan coast the mysids closely resemble cola: in the boreal waters of the North the faunal composition of taxa from analo• Atlantic and Arctic Ocean from the En• gous coastal regions of the Siberian Sea glish Channel and Cape Cod to the and the Canadian Arctic Archipelago. south, up to the southern regions of Baffin Bay (up to 71-7S0NL), along the BIOGEOGRAPHIC DISTRIBUTION eastern coast of Greenland (up to 76- 78°NL), Nordkapp Cape, and West Murman to the north (Zimmer, 1904, The following biogeographical groups are 1909; Linko, 1908; Tattersall, 1951). recognized: 3. Bathyal ubiquitous boreal species: Ery• throps glacialis, Parerythrops spectabilis, I. Boreal-subtropical species and Michthyops parva. Their distribution is analogous to species of the previous 1. Atlantic upper sublittoral boreal-sub• biogeographic group (Vanh6ffen, 1897; tropical species: Hemimysis lamornae: in Zimmer, 1904; Stephensen, 1913; Tat• the Atlantic boreal and subtropical as well as in the Mediterranean-Lusitan• tersall, 1951). t ian areas (Zimmer, 1904). 4. Sublittoral high boreal. Species: Dacty• lamblyops sarsi, and Mysis mixta with dis• 2. Atlantic bathypelagic boreal-subtropical tributions similar to species of the above• species found in the adjacent Arctic sec• mentioned biogeographic groups; how• tor as well: Boreomysis tridens (Plate 15- ever, they penetrate southward only as I, a), found in the Atlantic boreal and far as the coast of southern Norway, subtropical Mediterranean-Lusitanian Faroes Islands, Massachusetts Bay, off areas and off the Greenland coast the eastern coast of Greenland in the (Stephensen, 1933; Tattersall, 1951). 3. Amphiboreal-subtropical species, found *Elittoral zone of Kjelman (1877) ranging from 55-70 in the Atlantic sector of the Arctic: Boreo• m to 120-180 m, bathyal zone of Appellof (1912) rang• mysis arctica and B. microps; in boreal and ing from 200 m to 2000 m (in Golikov, 1985a). subtropical waters of the Northern tHigh boreal species are boreal species occurring in Hemisphere, in the Atlantic sector of the the northern parts of the boreal region; in the Atlantic and Arctic they do not penetrate south of Massachu• Arctic, reported off Greenland and off setts Bay, Faroes Islands, Norway (off Bergen). Their Franz Josef Land (Zimmer, 1904, 1909; northern limits of distribution are the same as those Stephensen, 1913; Tattersall, 1951). of the above-mentioned biogeographic group. w ~

Table 15-1. Results of the Calculations of Biogeographic Indexes.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 1 14 37.5 24.24 22.58 30 50 37.5 44.44 28.57 50 31.85 20 11.76 5.0 13.33 64.29 57.14 50 42.86 61.54 75 66.66 100 80 71.43 75 40 14.29 100

2 54.55 19 43.75 34.38 22.22 45.45 17.39 34.78 9.52 31.82 18.18 9.52 9.09 4.0 10 9 73.68 57.89 42.85 76.92 50 66.66 50 70 57.14 50 40 14.29 66.66 3 39.02 60.87 27 88.88 37.67 25 12.9 21.88 0 15.63 9.68 0 0 0 0 8 14 100 78.57 61.54 50 58.33 0 50 42.86 0 0 0 0 4 36.84 57.89 94.12 24 40.75 27.59 14.29 24.14 0 17.24 10.71 0 0 0 0 7 11 24 78.57 61.54 50 58.33 0 50 42.86 0 0 0 0 5 42.86 36.36 53.66 57.89 14 42.11 46.67 44.44 12.5 33.33 10.53 12.5 11.76 5.0 13.33 6 6 11 11 61.54 87.5 66.66 50 60 28.57 50 40 14.29 66.66

6 59.26 62.5 40 43.24 59.26 13 40 66.66 13.33 64.29 33.33 13.33 12.5 5.26 14.29 8 10 8 8 8 75 83.33 50 90 71.43 50 40 14.29 66.66 7 54.55 29.63 22.88 25 63.64 57.14 8 33.33 33.33 38.46 25 33.33 18.18 7.14 22.22 6 4 4 4 7 6 62.5 75 50 42.86 75 40 14.29 66.66

8 61.54 51.61 35.9 38.89 61.54 80 50 12 23.07 83.33 35.31 14.29 13.33 5.56 15.38 8 8 7 7 8 10 5 75 100 71.43 50 40 14.29 66.66 9 44.44 17.39 0 0 22.22 23.53 50 37.5 4 27.27 57.14 60 28.57 10 75 4 2 0 0 2 2 3 3 75 100 75 50 25 100 10 66.66 48.28 27.03 29.41 50 78.26 55.55 90.91 42.86 10 41.66 16.67 15.38 6.25 30 8 7 5 5 6 9 5 10 3 71.43 50 40 14.29 100

11 47.52 30.77 17.65 19.35 19.05 50 40 52.63 72.72 58.82 7 37.5 20 7.69 42.86 5 4 3 3 2 5 3 5 4 5 75 40 14.29 100 12 33.33 17.39 0 0 22.22 23.53 50 25 75 28.57 54.55 4 50 22.22 40 3 2 0 0 2 2 3 2 3 2 3 75 50 66.66

13 21.05 16.67 0 0 21.05 22.22 30.75 23.53 44.44 26.66 28.57 66.66 5 33.33 33.33 2 2 0 0 2 2 2 2 2 2 2 3 60 66.66 14 9.52 7.69 0 0 9.52 10 13.33 10.53 18.18 11.76 14.29 36.36 50 7 11.11 1 1 0 0 1 1 1 1 1 1 1 2 3 33.33 15 35.29 18.18 0 0 23.53 25 36.36 26.67 85.71 46.15 60 57.14 50 20 3 3 2 0 0 2 2 2 2 3 3 3 2 2 1 Arranged Diagonally: Number of Mysid Species. Lower Left Corner Above Chekanovsky-S0rensen Coefficient: Number of Species the Regions Have in Common (bottom). Upper Right Corner: Jaccard Coefficient (top). Shimkevich-Simpson Coefficient (bottom). Regions: 1, Baffin Bay; 2, eastern Greenland; 3, Norwegian Sea, region of Tronheim; 4, Norwegian Sea, region of Lofoten Islands; 5, southwestern Barents Sea; 6, northern Barents Sea; 7, Barents Sea southwest of the Novaya Zemlya; 8, northern Kara Sea and Novaya Zemlya Kara Sea trough; 9, southeastern Kara Sea and Laptev Sea; 10, northern Laptev Sea; 11, East Siberian Sea; 12, Chukchi Sea, east of Wrangel Island; 13, Chukchi Sea area off Point Barrow; 14, Bering Sea, Anadyr Bay; 15, Straits of the Canadian Arctic Archipelago.

w ~ 378 V V Petrvashov

southern parts of the boreal region; in V. Boreal-Arctic species the Atlantic and Arctic from the English Channel, the southern coast of Ireland, 1. Sublittoral upper bathyal boreal-Arctic and Cape Cod (eastern coast of the species: Meterythrops robusta, Pseudomma United States) to the south; up to No• truncatum (Plate 15-1, e) and Stilomysis vaya Zemlya, the western coast of Spit• grandis (Plate 15-1, f), present in the zbergen, , southern parts of the temperate waters of the Atlantic, Pa• Baffin Bay, and in the Intermediate Wa• cific, and Arctic oceans (Zimmer, 1904, ter Mass of the Greenland Sea to the 1909; Linko, 1908; Stephensen, 1913; north; (up to 72°N) (Zimmer, 1904; Tattersall, 1951); in the Arctic mainly in Linko, 1908; Tattersall, 1951). the Intermediate Water Mass from 5. Bathypelagic high boreal species dis• Spitzbergen up to the northern sector tributed in the sector adjacent to the of the Laptev Sea, seldom penetrating Arctic: Boreomysis nobilis is present from the Arctic surface water. the Faroes Islands, Iceland, eastern 2. Sublittoral upper bathyal Atlantic bo• coast of Canada up to Greenland (up to real-Arctic species: Erythrops abyssorum, 77°NL off the eastern coast and up to E. erythrophthalma (Plate 15-1, g), Parery• 75°NL in Baffin Bay) off Spitzbergen throps obesa, Pseudomma roseum, and Ps. (Hansen, 1888; VanhOffen, 1897; Ohlin, frigidum have the same distribution as 1901; Zimmer, 1904, 1909) and Franz Jo• the previous biogeographic groups, but sef Land. not recorded from the Pacific Ocean 6. Meso-bathypelagic amphiboreal spe• (Zimmer, 1904, 1909; Linko, 1908; cies: Amblyops abbreviata (Plate 15-1, c), Stephensen, 1913; Tattersall, 1951). present in the temperate waters of the 3. Bathypelagic Atlantic boreal-Arctic spe• Atlantic, Pacific, and Arctic Ocean but cies: Amblyopsoides ohlinii, found in the absent from the Arctic water masses northwestern sector of the Atlantic (Zimmer, 1904, 1909; Linko, 1908; Tat• Ocean and in the Norwegian Sea (Zim• tersall, 1951). mer, 1904; Stephensen, 1913; Tattersall, 1951). III. Sublittoral Pacific boreal species pene• 4. Sublittoral circumpolar Arctic-boreal trating into the Chukchi and Beaufort species: Mysis oculata (Plate 15-1, h) and Seas: Neomysis rayii, Exacanthomysis arcto• M. litoralis, found in the Arctic Ocean pacifica, and Xenacanthomysis pseudoma• and adjacent temperate waters of the cropsis (Murdoch, 1885a,b; Holmquist, Atlantic and Pacific oceans (Zimmer, 1982). 1904, 1909; Linko, 1908; Tattersall, 1951; Holmquist, 1982). IV. Bathyal-pseudoabyssal* bipolar spe• 5. Estuarine-freshwater circumpolar Arc• cies: Birsteiniamysis inermis (Plate 15-1, d), tic-boreal species: Mysis relicta, found observed in the Antarctic region and Sub• in estuaries of rivers flowing into the antarctic, in the North Pacific, North At• Arctic Ocean and also in a number of lantic, and Arctic (Zimmer, 1904; Steph• rivers and lakes of northern , ensen, 1913; Tattersall, 1951). northern Asia, and North America (Linko, 1908).

VI. Arctic species *Pseudoabyssal of Derjugin (1915) basins in the ba• thyal zone 2,000-5,000 m depth not connected with 1. Cryopelagic circumpolar species: Mysis the seafloor (in Golikov, 1985a). polaris (Holmquist, 1959; Kulikov, 1980). Arctic Ocean Mysids 379

2. Sublittoral upper bathyal species: Mich• Zemlya troughs of the Kara Sea. The re• thyops theeli, distributed in the adjacent gion from Wrangel Island and Beechey Atlantic sector of the Arctic Ocean from Point (North Alaska) to the north and to the eastern coast of Greenland (Ohlin, Prince of Wales Cape and Olyutorsky 1901) up to the northern Laptev Sea. Cape (North Kamchatka) to the south can 3. Bathypelagic species: Pseudomysis ab• be designated as an ecotone at the bound• yssi, observed in the Greenland and ary with the Pacific boreal fauna. Norwegian seas (Zimmer, 1904). The vertical distribution of mysids in the Arctic Ocean is presented in Figure 15- 3. Maximum number of species is ob• DISCUSSION AND CONCLUSIONS served in the Norwegian Sea, where spe• cies attain greatest diversity between 200 Distributional patterns of species belong• m and 400 m depth. Northward and east• ing to different biogeographic groups in ward the number of upper sublittoral, ba• the Arctic seas (Figs. 15-1 and 15-2) show thyal, and bathypelagic species declines. that Atlantic boreal and boreal-subtropical Thus, boreal and boreal-subtropical ba• species dominate the mysid fauna of the thyal and bathypelagic species can pene• Norwegian Sea, the southwestern areas of trate easily along the coast of east Green• the Barents and part of the Greenland seas land with the Intermediate Water Mass, (up to 76-78°NL); Pacific boreal species whereas boreal upper sublittoral species dominate the fauna of the Chukchi Sea, are not found here and the fauna of the northwestern Alaskan coast; boreal-Arctic coastal regions is represented by two species dominate the fauna of other parts sublittoral Arctic-boreal species. Bathyal of the Arctic Ocean. Arctic species do not and bathypelagic species are absent from dominate the mysids of other biogeo• the southwestern parts of the Barents Sea graphic groups in any sector of the Arctic and therefore the maximum number of Ocean, if the mysid fauna is regarded as a mysids occurs at 90-200 m depth. Like• whole and is not divided by separate water wise, maximum species diversity is ob• masses. served off the northwestern Alaskan coast Areas from the Labrador Peninsula up (Chukchi Sea) at 5-30 m depth. In the to 75°NL off the west coast and 78°NL off northern Laptev Sea, due to the predomi• the east coast of Greenland and from Nord• nance of elittoral-bathyal species, the kapp Cape up to the southwest coast of number of taxa is the highest at 200-400 m Novaya Zemlya and Franz Josef Land depth. should be considered as an ecotone, a The distribution of mysids at various transitionaL zone where mixing of boreal depths is related to the distribution of wa• and Arctic hydrobiocomplexes occur. * The ter masses in the Arctic Ocean. Thus the fauna of the latter, in the coastal regions, brackish-freshwater species Mysis relicta is dominated by Arctic and sublittoral Arc• occurs in water with temperatures not tic-boreal taxa belonging to the genus higher than 8°C or 9°C and salinities of 0- Mysis. Sublittoral upper bathyal and elit• 7%0. This species occurs in the Arctic toral upper bathyal species dominate the Ocean in estuaries at depths of 0-20 m; in fauna of the upper parts of the continental fresh water it was observed even at a slope from Spitzbergen up to the northern depth of 117 m (Lake Onega). The cryopel• sectors of the Laptev Sea and the Novaya agic species Mysis polaris is stenothermal (-1.7°C to + 1.1 0c) and stenohaline (29.47-34.42%0), living in the surface Arctic *hydrobiocomplex = hydroecosystem Water Mass and on the underside of ice. ~ % a 100 ......

' .. " .

I , 50 I \ .• ,. I ~ "/. : :..., I . \ l '·1 / ...... , .. .' " " , / \ I" \ --'" , ' . \\r;,,:,: . _ .. L .. _. \ I \ -11-11-11-11- ,,_ \ "'" - I'...... '" .-'-'-'-'-.-.- _:.,... '-, .,...... , I • • • . \ - ...... !"'. '~, ' ...... ,,_ .. - .. ~.. ~- .. - .. - .. - .. - .. - .. - \ - ,'- . I . ~-:-~._;,-== i, , I , I I 1\ I I J I : 700W 50° 30° 100W 100E 30° 50° 70° 90° 110° 130° 15001700E 1700W15001300 1100 90° 700W Figure 15-1. Number of mysid species belonging to different biogeographic groups in the Arctic Ocean. Abscissa: longitude; ordinate: percentage of species belonging to a certain biogeographic group out of the total number of species at a given site (%):

c -- - 1 boreal Atlantic species and amphiboreal species E··- .. - ..j bipolar species ,...... I boreal-Arctic species EII-II-IH boreal subtropical species

E· - . _. "'l Arctic species EI-I-H boreal Pacific species Arctic Ocean Mysids 381

%

20

15

10

5

OL-~~~~~~~--~-L~--L#'~I--~-L~--~~~--~·!V/~~~~-L~ 20 40 60 80 100 300 500 700 1000 2000 3000 m

Figure 15-2. Distribution of Arctic Ocean mysids at various depth zones. Abscissa: depth in meters (m); ordinate: number of species (%).

%

\ \ \ , . . . . .' . \ \ \: 50 :\ \ ------

20 40 60 80 100 300 500m

Figure 15-3. Vertical distribution of mysids off Nordkapp Cape. Abscissa: depth in meters (m); ordinate: percentage of species belonging to a biogeographic group out of the total number of species at the given site (%). Legend as in Figure 15-1. 382 V. V. Petryashov

The upper sublittoral species Exacantho• theeli (1.19°C to -1.80°C), Erythrops abys• mysis arctopacifica, Hemimysis lamornae, Neo• sorum (-1.77°-7.22°C), E. glacial is (0.9°- mysis integer, N. rayii, Praunus flexuosus, P. 1. 1°C), E. microps (10.7°C), Meterythrops ro• inermis, Schistomysis ornata, and Xenacan• busta (-1.77°-2°C), Pseudomma roseum thomysis pseudomacropsis are eurythermal (-1.77°-4.4°C), Ps. frigidum (0.6°-1.3°C), (-0.4° to + 10.0B°C) and euryhaline (23- Parerythrops spectabilis (0.3°-1.0°C) all are 32.66%0); some species occur even at river stenohaline (31.47-35%0) and were ob• mouths where salinities are much lower served in water saturated with oxygen than 23%0 and in the Arctic region where (72.B-109.B%). These taxa occur at 54-900 m they have been observed in the surface depth in the Atlantic boreal and a consider• North Atlantic and surface Pacific water able number have been found in the Inter• masses at depths of 0-51 m. The sublittoral mediate Atlantic-Arctic water masses. Mysis litoralis, M. oculata, and M. mixta are The meso-bathypelagic species Boreo• eurythermal (M. mixta 0.02°-13.6BoC and mysis arctica, B. nobilis, Amblyops abbreviata, M. oculata and M. litoralis -1.77°-5.4°C) and and Michthyops parva are also eurythermal euryhaline (6.2-34.93%0); they have been (-0.B2-5.5°C) and stenohaline (34.9- observed when the water was saturated 35.55%0), but inhabit depths of 140-1,250 with oxygen (up to 45-96.3 percent). M. m in the Atlantic boreal and Intermediate mixta occurs in the surface Atlantic Water Atlantic-Arctic water masses. The distri• Mass at 1-200 m depth while M. oculata bution of bathypelagic species Boreomysis and M. litoralis have been observed in sur• microps, B. tridens, Birsteiniamysis inermis, face Arctic water, estuarine-Arctic water Amblyopsoides ohlinii, and Pseudomysis ab• and even in cool surface Atlantic and Pa• yssi at various depths is more complex, al• cific water masses at 1-249 m depth. though all are stenothermal (+ 1.0° to The sublittoral upper bathyal Stilomysis -1.0°C) and stenohaline (34.9-34.99%0). grandis, Erythrops erythrophthalma, Pseu• B. microps and B. tridens occur in the Atlan• domma truncatum, Misidopis didelphys, and tic boreal and the Intermediate Atlantic• probably Dactylamblyops sarsi can be both Arctic water masses, while A. ohlinii is stenothermal (e.g., E. erythrophthalma and present in the Arctic deep-water mass at Ps. truncatum, -0.2° to -1.B2°C, but prob• depths of 400-2,000 m. B. inermis inhabits ably occur at higher temperatures) and eu• depths of 900-2,970 m within the Arctic rythermal (e.g., St. grandis -1.59° to Ocean, and P. abyssi lives in the deep-wa• +6.4°C); however, all are stenohaline ter mass exclusively at 1,406-2,700 m (31.15-35%0) and have been observed in depth. water saturated with oxygen (e.g., E. ery• It should be emphasized that in the In• throphthalma at 53.3-B3%, Ps. truncatum at termediate Atlantic-Arctic waters, which 32.3-93.6%0 saturation). These taxa are upwell in the Novaya Zemlya trough in found in the Arctic Ocean within the At• the Kara Sea, a number of species occur lantic boreal and intermediate Atlantic• at uncharacteristic depths. Thus Erythrops Arctic water masses at 1B-5BO m depth. abyssorum (with a depth habitat of 180-540 Very closely related to this group are m) rises here to 64 m and Michthyops theeli the elittoral upper bathyal species Ery• (119-575 m) rises here to 27 m. Analogous throps abyssorum, E. glacialis, E. microps, E. phenomenon has been observed in this re• serrata, Meterythrops robusta, Parerythrops gion for other bathyal invertebrates (Zen• obesa, P. spectabilis, Michthyops theeli, kevich, 1963). Pseudomma affine, Ps. roseum, Ps. frigidum, Information about the distribution of Mysideis ins ignis, and Hemimysis abyssicola. different water masses is provided by the In spite of the differences among these composition of different biogeographic species in their thermopathy Michthyops groups at different depths. Atlantic boreal Arctic Ocean Mysids 383 species (Fig. 15-3) dominate at depths shal• been observed here mainly within a lim• lower than 10 m in the southwestern Bar• ited depth range (10-30 m though some• ents Sea while boreal-Arctic species domi• times to depths of 200-240 m); the mysids nate at greater depths and the number of do not dominate species of other biogeo• Atlantic boreal species slightly increases at graphic groups. Atlantic boreal waters do depths of 400-600 m. In the southern areas not penetrate the Kara Sea beyond its of this sea, east of Kola Bay, boreal-Arctic southern sector; therefore, Atlantic boreal species dominated within the entire water species have not been observed in this sea. mass. This region is characteriied by a In the Kara Sea and further east along sharp decline in the numbers of boreal taxa the coastal regions of the Siberian Arctic in the area between the Varanger fjord and seas, the surface Arctic and estuarine-Arc• Kola Bay. tic water masses (except for the Chukchi Atlantic boreal species penetrate into Sea) at depths of 100-200 m are entirely the southern Barents Sea since the Atlantic dominated by sublittoral Arctic-boreal Boreal Water Mass extends in this area species and estuarine-freshwater Arctic• from the surface to 600 m depth. Sublit• boreal taxa. These groups are replaced by toral Arctic-boreal species occur in the cryopelagic circumpolar Arctic species cooler regions of this water mass (Nord• only in cryobiocenoses. The distributional kapp branch of the North Atlantic Drift) in patterns of mysids off east Greenland are the southwestern areas of the Barents Sea somewhat different (Fig. 15-4). Boreal• and further east. This mysid group has Arctic species occur in this area only to a

% 100 r - --, I I • , J I

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I · " f J , .., I I < ' It ,I ! o '/ // ao 20 40 60 80100 500700 1000 2000 3000 m Figure 15-4. Vertical distribution of mysids off the eastern Greenland coast (75°). Abscissa: depth in meters (m); ordinate: percentage of species of a biogeographic group of the total number of species at the given site (%). Legend as in Figure 15-1. 384 V. V. Petryashov depth of 200 m, in the surface water of the Laptev Sea (Fig. 15-5) in the uppermost Greenland Sea. In the Intermediate Water layers (0-5 m), and at depths of 5-40 m in Mass these taxa occur only in the upper• the summer period the fauna is domi• most layers, to a depth of 400 m. Atlantic nated by sublittoral Arctic-boreal species. boreal and amphiboreal-subtropical spe• The mysids of these two groups are typical cies found in the Arctic Ocean predomi• representatives of the surface Arctic Water nate in this water mass. Only two Atlantic Mass. Lower layers of this water mass and boreal species penetrate to depths greater the Intermediate Atlantic-Arctic Water than 600 m, into the cold Arctic Water Mass are dominated by sublittoral upper Mass and even then only in its upper lay• bathyal boreal-Arctic species. In the deep• ers (to a depth of 800 m). In the Greenland water mass of the Central Arctic Basin, at and Norwegian seas' basins at depths in depths in excess of 900 m, the bipolar Birs• excess of 1,400 m only the Arctic Pseudo• teiniamysis inermis has been found exclu• mysis abyssi, the boreal-Arctic Amblyop• sively. The mysid fauna within the Arctic so ides ohlinii, and the bipolar Birsteiniamysis Ocean is most peculiar in the southeastern inermis have been observed to date. parts of the Chukchi Sea where some Pa• The distributional patterns of mysids in cific boreal species penetrate (Fig. 15-6). different depth zones in the northern areas The latter dominate mysids off the north• of Kara and Laptev seas is quite unusual. western Alaskan coast from the Bering For example, the Arctic cryopelagic species Strait up to Cape Barrow at depths of 1-45 Mysis polaris occurs in the northwestern m. The great depths and regions not influ-

%

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50

:\ r·l· / 1: / .. . :\ -- --... • ....J I:

: \ I' ( , . I I " I 0 / • 20 40 60 80100300 700 10(fO 2000 3000 m Figure 15-5. Vertical distribution of mysids in the northwestern Laptev Sea. Abscissa: depth in meters (m); ordinate: percentage of species of a biogeographic group out of the total number of species at the given site ('Yo). Legend as in Figure 15-1. Arctic Ocean Mysids 385

% the species belonging to this genus may well have evolved in three directions: (1) •• '. ' ••••••• &" 100 by adaptation to conditions of the coastal Atlantic boreal waters (M. gaspensis, M. mixta, and M. stenolepis), (2) by adaptation to coastal Arctic regions (M. oculata, M. Ii• toralis, and M. reIicta), and (3) by adapta• tion to the epipelagic zone and later to the cryopelagic zone (M. polaris). The evolu• so tion of the cryopelagic species M. polaris is related to the onset of ice cover over the Arctic Ocean. Therefore, this species must have existed in the Arctic during the entire late Neogene ice age or at least during the Last Glaciation. Following the melting of ice sheets of the Last Glaciation along the o '---:-1 '="0-2-="0=---:3:'::0:-:'4'="0-'-S=--0::-:-1 ~00=-'--::2:-:0:-::0:--'-'-----I~ shallow Arctic seas and the subsequent in• 150 250 tensive freshening of coastal waters, the estuarine freshwater species M. relicta pen• Figure 15-6. Vertical distribution of mysids off the Northwestern Alaskan Coast. Abscissa: etrated the coastal zones. At the same depth in meters (m); ordinate: percentage time, M. oculata and M. litoralis penetrated of species of the biogeographic group out to great depths where salinities were ex• of the total number of species at a given site ceeding 6-8%0. With decreased freshwater (%). Legend as in Figure 15-1. discharge from the continent, these spe• cies became distributed in coastal waters also, and M. relicta must have formed iso• enced by the Pacific surface waters are lated populations in the river estuaries of dominated by sublittoral Arctic-boreal the Arctic seas. species. Mysids belonging to the genus Mysis (e.g., M. relicta), which originated in the period of intensive melting of glaciers and EVOLUTION OF THE MYSID FAUNA therefore strong freshening, were euryha• line and were able to penetrate from the In the view of this author, the formation of White Sea through the area of the recent the recent mysid fauna of the Arctic Ocean Onega and Ladoga lakes into the arm of is related to the termination of the Last the sea that existed at the locus of the pres• Glaciation (Wiirm-Valdai-Wisconsin Gla• ent Baltic Sea (Lavrova, 1960). When this ciation, 10-12 thousand years ago), the on• connection ceased, isolated populations of set of the Postglacial transgression, the es• M. relicta developed in the deepest lakes, tablishment of recent water circulation in the areas occupied by a former marine ba• the Northern Hemisphere, and the devel• sin. It is believed that at that time or dur• opment of Arctic Ocean water masses with ing a previous boreal transgression mysids their specific temperature-salinity struc• belonging to this genus penetrated from ture. the region south of the recent Ladoga and Each biogeographic group has its own Onega lakes through river systems or history of distribution within the study through Rybnoye Lake into the Volga Ba• area. The genus Mysis holds a particular sin and further into the Caspian Sea (Hog• place in the mysid fauna of the Arctic born, 1917; Berg, 1928) where the Mysis Ocean. In the Arctic and adjacent waters fauna (M. amhlyops, M. caspia, M. micro- 386 V V Petryashov lepis, and M. microphthalma) endemic to Greenland Sea's Intermediate Water Mass the Caspian Sea originated. at 200-500 m depth. The penetration of At• The Atlantic boreal and the genetically lantic boreal and boreal-Arctic (except Arc• related boreal-Arctic taxa (except for Arc• tic-boreal) species into Baffin Bay is via the tic-boreal) is the largest group, including Irminger Current and the relatively warm 36 species. Subsequent to the Last Glacia• West Greenland Current. tion, species belonging to these groups mi• The distribution of three Pacific boreal grated from the southwestern European species in the Chukchi Sea is related to the coast and the eastern coast of North Amer• development of the Postglacial transgres• ica northward. This distribution was en• sion and the opening of the Bering Strait hanced by the North Atlantic Drift and its (Monin, 1980; Morskaya Geomorphologiya, branches that reach far north. Thus, for ex• 1980). These events led to the penetration ample, with the Nordkapp branch of the of a warm current through the Bering North Atlantic Drift, 12 boreal and boreal• Strait that bifurcates in the Chukchi Sea. Arctic species reach into the middle Mur• The northeastern branch flows toward man region (Kola and Motoka fjord) and Point Barrow while the northwestern arm some taxa (Pseudomma truncatum, Stilomysis flows toward the region northeast of grandis, Praunus inermis, and Mysis mixta) Wrangel Island. This current influences a extend their range to Novaya Zemlya. restricted area, as compared to the influ• The West Spitzbergen branch of the ence of the North Atlantic Drift on the Arc• North Atlantic Drift has had a significant tic Ocean. This is due to the shallow influence on the regional faunal composi• depths of the Bering Strait, which limits tion. East of Spitzbergen this branch de• the volume of warm water carried through scends gradually under the surface Arctic the strait to 20,000 km3 as compared to the water and in the Arctic Ocean descends 150,000 km3 of Atlantic water carried into under the Intermediate Atlantic-Arctic the Norwegian Sea from the south (Zen• Water Mass. Near Spitzbergen it flows at kevich, 1963). a depth of 0-900 m, north of Franz Josef Of the two species Michthyops theeli and Land at a depth of 150-900 m, and north Pseudomysis abyssi endemic to the Arctic of Severnaya Zemlya at a depth of 200-900 Ocean, the former most probably is of bo• m (Zenkevich, 1963). With this narrow wa• real origin as suggested by its distribution, ter mass band, 9 elittoral upper bathyal which is similar to that of a number of and meso-bathypelagic species penetrate Atlantic boreal-Arctic species (Golikov, into the northern areas of the Barents, 1985b). The second species Pseudomysis ab• Kara, and Laptev seas including the No• yssi may be an Ice Age relict in the Green• vaya Zemlya trough of the Kara Sea. The land and Norwegian seas. The genus boundary of distribution of these species Pseudomysis is amphiboreal. The ances• coincides almost entirely with the 0.5°C tors of this taxon must have penetrated into isotherm at 300 m depth (Zenkevich, the North about 3 mya 1963). Some boreal and boreal-Arctic spe• from the Pacific Ocean along the Canadian cies belonging to the main elittoral upper coast during a period when the Arctic was bathyal and meso-bathypelagic zones oc• ice-free. The dispersal of other amphibo• cur off the eastern Greenland coast bathed real species and genera of mysids from the by the cool East Greenland Current. This North Pacific into the North Atlantic may occurrence is related to the formation of have been the same. Arctic Ocean Mysids 387

APPENDIX

Plate 15-1 Several characteristic mysids of the Arctic Ocean (after Zimmer, 1904): a, Boreomysis tridens; b, Praunus inermis; c, Amblyops abbreviata; d, Birsteiniamysis inermis; e, Pseudom• ma truncatum; f, Stilomysis grandis; g, Erythrops erythrophthalma; h, Mysis oculata.

Map 15-1 Northern limit of distribution of mysids in the Arctic Ocean.

1. I Atlantic upper sublittoral boreal-subtropical species. 2. E-- - - J Bathypelagic boreal and boreal-subtropical species: 6 Boreomysis arctica. 3. I Atlantic boreal and amphiboreal species: o Boreomysis nobilis. 4. E - - -1 Sublittoral Pacific boreal species. 5. E" -..=J Bathyal, pseudoabyssal bipolar species.

Map 15-2 Northern limits of distribution of mysids in the Arctic Ocean.

1. I··········· ·1 Northern limit of sublittoral, upper bathyal boreal-Arctic species. E------I Northeast limit of Erythrops abyssorum. 2. 6 Bathypelagic Atlantic boreal-Arctic species. 3. Southern limits of sublittoral circumpolar Arctic-boreal E'-'---I species. 4. • Cryopelagic circumpolar Arctic species . 5. r"-'-""-I Sublittoral, upper bathyal Arctic species. 6. 0 Bathypelagic Arctic species. a b

c d PLATE 15-1

388 389 Arctic Ocean Mysids

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Holmquist, Ch. 1949. Uber cventueIIe inter• REFERENCES mediare Formen zwischen Mysis oculata Fabr. und Mysis relicta Loven. K. Fysiografiska Aurivillius, C. W. S. 1896. Das Plancton der Baf• Siillskapets Handlingar, N. F. 60(10):3-26. fin's Bay und Davis Strait eine thiergeographi• Holmquist, Ch. 1958. On a new species of the sche Studie. Festschrift Lilljeborg, Uppsala, genus Mysis, with some notes on Mysis ocu• pp. 181-212, 1 plate. lata (0. Fabricius). Medd. Gmnland 159(4):3- Berg, L. S. 1928. 0 proiskhozhdenii severnykh 17. elementov v faune Kaspiya. Doklady AN SSSR Holmquist, Ch. 1959. Problems on Marine-Glacial 14(3):107-112. (In Russian.) Relicts, on Account of Investigations on the Ge• Bertelsen, E. 1937. Contributionals to the ani• nus Mysis. Lund, 270p. mals ecology of the fjords of Angmagssalik Holmquist, Ch. 1963. Some notes on Mysis re• and Kangerdlugssuaq in east Greenland. licta and its relatives in northern Alaska. Arc• Medd. Gmnland 108(3):1-58 and 2 plates. tic 16:109-128. Birula, A. 1897. Recherches sur la biologie et Holmquist, Ch. 1982. Mysidacea of the West zoogeographie, principalement des mers Coast of North America. Zoologische Jahr• Russes. III. Essai d'une faune des crustaces bucher 109(4):469-510. decapodes de la Mer Blanche et Mourmanne. Holt, E. W. L., and W. M. Tattersall. 1905. Schi• Ann. Mus. Zool. Acad. Imp. Sci., St. Petersburg zopodous Crustacea from the north-east At• 2:405-452 and 3 plates. lantic slope. Rept. Sea Inland Fisheries Ireland Bj6rck, W. 1916. Die Schizopoden des Eisfjords. pt. 2, app. 4, pp. 99-152, 11 plates. Zoologische Ergebnisse der Schwedischen Illig, G. 1930. Schizopoden der Deutschen Expedition nach Spitzbergen, 1908, Tell II. Tiefsee Expedition. In Chun, Wissenschaftliche Handl. K. Svenska Vetensk. Acad., Stockholm Erbegnisse Deutsche Tiefsee-Expedition auf dem 54(6):1-10 and 1map. Dampfer "Valdivia" 1898-1899. Jena, Gustav Czerniavsky, V. 1887. Monographia mysidarum Fischer, Bd. 22, pp. 399-625. imprimis imperii Rossici (marin., lacustr., et Jaccard, P. 1901. Distribution de la £lore alpine fIuviatum). Fasc. III. Trav. Soc. Imp. Nat. St. dans Ie Bassin de Dranses et dans quelques Petersburg 18:1-102 and 28 plates. regions voisines. Bull. Soc. Voudoise Sci. Na• Dunbar, M. J. 1940. On the size distribution tur. 37(140):221-272. and breeding cycles of four marine plank• Jarzynsky, Th. 1870. Praemissas Catalogus tonic from the Arctic. J. Ecol. crustaceorum decapodorum, inventorum, in 9:215-226, 5 figs. mari albo, et in mari glacialis ad litus mur• Gerstaecker, A., and A. E. Ortmann. 1901. Ma• manicum auno 1869 et 1870. Trudy Soc. Nat. lacostraca. In H. G. Bronn (ed.), Klassen und Petropol. 1(2):317. Ordnungen des Tierreichs. Leipzig, C. F. Win• Jashnov, V. A. 1948. Otryad Mysidacea. Oprede• ter'sche VerIagschandlung, Bd. 5, Abt. 1, Pt. litel Fauny i Flory Severnykh Morei SSSR pp. 2, pp. 602-686. 224-229. (In Russian.) Golikov, A. N. 1985a. Zonations and organis• Kramp, P. L. 1913. Schizopoda. In C. H. Osten• mic assemblages: Comments on the compre• feld (eds.), Resume des Observations sur Ie hensive review by Peres (1982). Mar. Ecol. Plankton des Mers Explorees par Ie Conseil Pen• Prog. Ser. 23:203-206. dant les Annees 1902-1908. 3 parts. A. F. Hest Golikov, A. N. 1985b. Biologiya. Istoriya prois• and Son, Copenhagen, pp. 539-556. hozhdeniya zhizni i biogeographicheskoe rai• Kulikov, A. C. 1980. K. ecologii dvukh vidov onirovanie. Geographiya Mirovogo Oceana-Sev• gammarid (Amphipoda, Gammaridea) i mis• ernyi I.edovityi i Juzhnyi Oceany, L. pp. 114- idy (Mysidacea) v kriopelagicheskom biot• 119. (In Russian.) zenose Centralnogo Arcticheskogo Basseina. Hansen, H. J. 1888. marine Biologia Centralnogo Arcticheskogo Basseina. pp. Groenlandiae occidentalis. Oversigt over det 111-118. (In Russian.) vestlige Groenlands fauna af malacostrake Lavrova, M. A. 1960. Chetvertichnaya Geologiya havkrebsdyr. Vidensk. Medde!. Naturhist. For. Kolskogo Poluostrova. - M. -L., 234p. (In Rus• Kjobenhaven 9(4):5-226. sian.) Hansen, H. J. 1908. Crustacea Malacostraca. I. Linko, A. 1907. Schizopodes de la mer glaciale Copenhagen, The Danish Ingolf Expedition du Nord Russe. Bull. Acad. Imp. Sci. St. Peters• 3(2):1-120, 4 figs., ~.plates, 1 map. burg 1(6):491-493. H6gbom, A. G. 1917. Uber die arktischen Elem• Linko, A. 1908. Schizopoda severnykh rus• ente in der Aralokaspischenfauna, ein tier• skikh moreL Mem. Acad. Imp. Sci. St. Peters• geographisches Problem. Bull. Geol. Inst. burg 18(8):1-76 and 5 figs. (In Russian.) Univ. Upsala 14:241-260, 2 figs. and 1 plate. Madsen, H. 1936. Investigations on the shore Arctic Ocean Mysids 395

fauna of east Greenland with a survey of the iyu Vnutrennikh Vodoyomov Pribaltiki. Chapter shores of other Arctic regions. Medd. Gmn• I, Petrosavodsk, pp. 80-82. (In Russian.) land 100(8):1-79 and 17 figs. Saemundsson, B. 1937. Icelandic Malacostraca Mauchline, J. 1980. The biology of mysids and in the Museum of Reykjavik. Reykjavik, Visen• euphausiids. Adv. Marine Bioi. 18:681p. dafelag Islendinga 20:1-32 and 1 map. Mauchline, Land M. Murano. 1977. World list Sars, G. O. 1870. Carcinologiske Bidrag til of the Mysidacea, Crustacea. J. Tokyo Univ. Norges Fauna. I. Monographi over de ved Fish. 64:39-88. Norges Kyster forekommende Mysider. Pt. I. Miers, E. J. 1877. Report on the Crustacea col• Christiania K. Norske Videnskabsselskab Trond• lected by the naturalists of the Arctic expedi• hjem, pp. 1-64 and 5 plates. tion in 1875-1876. Ann. Mag. Nat. Hist. Sars, G. O. 1885. Crustacea I. In Den Norske 20(ser. 4):52-66 and 2 plates. Nordhavs Expedition, 1876-78. Christiania Monin, A. S. 1980. Popularnaya Istoriya Zemli. 14:1-280. M. 224p. (In Russian.) Sars, G. O. 1886. Crustacea II. In Den Norske Morskaya Geomorphologiya. 1980. Terminolog• Nordhavs Expedition, 1876-78. Christiania icheskii spravochnik. M. 279p. (In Russian.) pp. 1-96 and 1 map. Murdoch, J. 1885a. Description of seven new Schmitt, W. L. 1919. Schizopod . In species of Crustacea and one worm from Arc• Report of the Canadian Arctic Expedition 1913- tic Alaska. Proc. U.S. Natl. Mus. 7:518-522. 18, vol. 7. Ottawa, pp. 1-8. Murdoch, J. 1885b. Marine invertebrates. Exclu• Simpson, G. G. 1943. Mammals and the nature sive of mollusks). In Report of the International of continents. Am. ]. Sci. 241(1):1-31. Polar Expedition to Point Barrow, Alaska. Part 4, S0rensen, T. 1948. A new method of establish• Natural History, Washington, pp. 136-176 ing groups of equal amplitude in plant sociol• and 2 plates. ogy based on similarity of a species content Oceanographicheskaya Enciclopediya. 1974. Pod and its application to analysis of the vegeta• red. Rhodes W. Fairbridge, 631p. (In Rus• tion on Danish commons. Kgl. Dan. Viden• sian.) Translated from The Encyclopedia of skab. Selskab. BioI. Skr. 5(4):1-34. Oceanography, 1966, Reinhold Publ. Stebbing, T. R. R. 1900. Arctic Crustacea: Bruce Ohlin, A. 1901. Arctic Crustacea collected dur• collection. Ann. Mag. Nat. Hist. 5(ser. 7):1-16. ing the Swedish Arctic expeditions 1898, 1899 Stephensen, K. 1912. Report on the Malacos• and 1900 under the direction of A. G. Nath• traca Pycnogonida and some Entomostraca orst and G. Kolthoff. II. Decapoda, Schizo• collected by the Danmark expedition to north• poda. Handl. K. Svenska Vetensk, Bihang 27(pt. east Greenland 1906-1908. Medd. Gmnland 4):1-91. 45:501-630 and 5 plates. Ortmann, A. E. 1901. Crustacea and Pycnogon• Stephensen, K. 1913. Report on the Malacos• ida collected during the Princeton expedition traca collected by the "Tjalfe" expedition, to North Greenland. Proc. Acad. Nat. Sci. under the direction of Ad. S. Jensen, espe• 53:144-168. cially at W. Greenland. Vidensk. Medd. Dansk Paulson, o. 1909. Plancton investigations in the Naturhist. For. Kj0benhavn 64:57-134 and 36 waters round Iceland and in the North Atlan• figs. tic in 1904. Medd. Komm. Havunders0gelser, Stephensen, K. 1918. Gmnlands Krebsdyr og Copenhagen, ser. Plankton, 1(8):1-57 and 9 figs. Pycnogonider. Medd. Gmnland 22:479p. Ross, J. C. 1835. Marine invertebrate animals. Stephens en, K. 1933. The Godthaab expedition In Appendix to the Narrative of a Second Voyage 1928. Schizopoda. Medd. Gmnland 79(9):1-20 in Search of a North-west Passage and of a Resi• and 6 figs. dence in the Arctic Regions during the Years Stephensen, K. 1938. Euphausiacea, • 1829-1833 (Sir John Ross, captain). A. W. Web• cea, Cumacea and Nebaliacea. In A. S. ster, London, pp. 81-150. Jensen et al. (eds.), The Zoology of Iceland, Co• Rusanova, M. N. 1962. Materialy po biologii penhagen and Reykjavik, vol. 3, pt. 29. Levin massovykh vidov misid Belogo morya. In Gi• and Munksgaard, pp. 1-24 and 1 plate. drobiologicheskie issledovaniya. 3. Tartu, pp. Tattersall, W. M. 1951. A review of the Mysida• 136-140. (In Russian.) cea of the United States National Museum. Rusanova, M. N. 1963. Kratkie svedeniya po bi• Bull. U.S. Natl. Mus. 201:i-x, 1-292, 103 figs. ologii nekotorykh massovykh vidov bespoz• Vanh6ffen, E. 1897. Die fauna und flora Gmn• vonochnykh raiona mysa Kartesh. Materialy lands. In E. von Drygalski (ed.), Gmnland Ex• po Kompleksnomu Isucheniyu Belogo Morya pedition der Gesellschaft fUr Erkunde zu Berlin, 2:53-65. (In Russian.) 1891-1893, vol. 2. W. H. Kiihl, Berlin, pp. 1- Rusanova, M. N. 1971. Genesis misid Belogo 383, 30 figs., 8 plates, 1 map. morya. In Materialy XVI Conferencii po Isuchen- Walker, D. 1862. Notes on the zoology of the 396 V. V. Petryashov

last Arctic expedition under Capitain Sir F. L. fica, vol. 3. Gustav Fischer, Jena, pp. 415-491 M'Clintock, R. N. f. Roy. Dublin Soc. 3:61-77. and 172 figs. Zenkevitch, L. A. 1963. Biologiya Morei SSSR. Zimmer~ C. 1909. Die nordischen Schizopoden. (In Russian.) In K. Brandt and C. Apstein (eds.), Nordisches Zimmer, C. 1904. Die Arctischen Schizopoden. Plankton, vol. 12. Lipsius and Tischer, Kiel In Romer Fritz und Fritz Schaudinn, Fauna Arc- and Leipzig, pp. 1-172 and 384 figs.