Translation Series No.1375

Translation Series No. 1375

Bioebenoses and biomass of benthos of the Newfoundland-Labrador region.

By Ki1N. Nesis

Original title: Biotsenozy i biomassa bentosa N'yufaund- • .lendskogo-Labradorskogo raiona..

From: Trudy Vsesoyuznogo Nauchno-Issledovatel'skogo •Instituta Morskogo Rybnogo Khozyaistva Okeanografii (eNIRO), 57: 453-489, 1965.

Translated by the Translation Bureau(AM) Foreign Languages Division Department of the Secretary of State of Canada

Fisheries Research Board of Canada • Biological Station , st. John's, Nfld 1970

75 pages typescript 'r OEPARTMENT OF THE SECRETARY OF STATE SECRÉTARIAT D'ÉTAT TRANSLATION BUREAU BUREAU DES TRADUCTIONS FOREIGN LANGUAGES DIVISION DES LANGUES DIVISION ° CANADA ÉTRANGÈRES

TRANSLATED FROM - TR,ADUCTION DE INTO - EN

Russian English

'AUTHOR - AUTEUR Nesis K.N.

. TITLE IN ENGLISH - TITRE ; ANGLAIS Biocoenoses and biomas of benthos of the Newfolindland-Labradoriregion

Title . in foreign_iangnage---(tranalitarate_foreisn -ottantatere) Biotsenozy i biomassa bentosa N i yufaundlendSkogo-Labradorskogoraiona.

, .ReF5RENCE IN FOREIGN ANGUA2E (NAME OF BOOK OR PUBLICATION) IN FULL. TRANSLITERATE FOREIGN CFiA,IRACTERS. • REFERENCE' EN LANGUE ETRANGERE (NOM DU LIVRE OU PUBLICATION), AU COMPLET. TRANSCRIRE EN CARACTERES PHONETIQUEL •. Trudylesesoyuznogo nauchno-iàsledovaterskogo instituta morskogo — rybnogo khozyaistva i okeanogràfii

- :REFEREN CE IN ENGLISH - RÉFÉRENCE EN ANGLAIS • Trudy of the 40.1-Union Scientific-Research Instituteof Marine

. Fiseriés and Oceanography.

PUBLISH ER ÉDITEUR PAGE,NUMBERS IN ORIGINAL DATE OF PUBLICATION NUMEROS DES PAGES DANS DATE DE PUBLICATION . L'ORIGINAL

YE.tR ISSUE.NO . 36 VOLUME ANNEE NUMERO PLACE OF PUBLICATION NUMBER OF TYPED PAGES LIEU DE PUBLICATION NOMBRE DE PAG.ES DACTY LOGRAPHIEES 1965 5 7

REQUEr IN G• DEPA RTMENT Fisheries Research Board TRANSLATION BUREAU NO. 2591 MIN ISTERE•CLIENT NOTRE DOSSIER Pi°

ERANCH OR DIVISION Biological Station TRANSLATOR (INITIALS) •14.• DIRECTION OU DIVISION TRADUCTEUR (INITIALES) Mr. T.K. Pitt PERSON etEQUESTING St. John's, Nfld. DATE C.OMPLETE p 2 Feb 1970' DEMANDE PAR . ACHEVE LE

YO UR NUM BER 769-18-14 UNED:TI:D. DIT:AFT TRANSLATIop VOTRE DOSSIER N° • On:y inforination TRADLIC,i-i'ON NON REVISU . DAT E OF R EQUEST 1209.69 DATE DE LA DEMANDE ...swlement •

.FLD 69A CV' DEPARTMENT OF THE SECRETARY OF STATE SECRÉTARIAT D'ÉTAT r TRANSLATION BUREAU BUREAU DES TRADUCTIONS FOREIGN LANGUAGES DIVISION DIVISION .DES LANGUES ÉTRANGÈRES

CANADA

YOUR NO. DEPARTMENT DIVISION/BRANCH CITY VOTRE N0 ■ Amirriptc DIVISION/DIRECTION VILLE 769-18-14 Fisheries Research iological Station St.John's, Nfld. Board

OU R NO. LANSUAOE TRANSLATOR (INITIALS) DATE NOTRE No LANGUI TRADUCTEUR (INITIALES) 2951 Russian A .M. FEB - 5 1970

UNEDITED DRAFT TRANSLATION Only i or inIorrnatien TRADUCTION NON REVIS4 Information sot.demere

.BIOCOENOSES AND BIOMASS OF BENTHOS ,OF THE•NEWFOUNDLAND LABRADOR REGION K. N. Nesis

The shores of. Newfoundland and Labrador were the first parts of the American continent to be seen and visited by Europeans, the Norsemen of Iceland (Bjarne Herjulfson in 985 and Leif Ericson around the year 1000). The discovery of the fishing banks of Newfoundland was, from an economic aspect, the most important result of the voyages of Cabot in 1497 and Cortereal in 1500. The ' exploitation of the riches of the sea preceded.by decades the colonization of the shores. Thus it is, that for several centuries, tens of millions of people have been fed by the cod and haddock of Newfoundland. The abundance of fish in the area is a consequence of the abundance of fish foods, •the chlef of which are the invert,ebrate- animals. Despite *Translator's note: Numbers in the margin refer to the page of the original text.

SOS-100..10.»3 4 '

all this, the region of Newfoundland and Labrador has long escaped the attention of marine biologists. Major Zuropean expeditions, ("Challenger", "Princess Alice", "Irondel", •"Mikhaelt Sars" and others), visited the Newfoundland region only in passing. For this reason, existing faunal records in the works of Brunel (1961a), La Rocque (1953), Packard (1867), Verrill (1885) and Whiteaves (1901) encompass sufficiently fully only the fauna of the coastal areas. Only on the benthos of Georges Bank has there been a short paper, by P. Wigley (1961). The fish industry requires knowled ,_?.e of the productivity of a body of water and, in particular, the cond- ition. of the food base for commercial fish. It is essential also to have detailed knowledge of the hydrological regime, on which the distribution of fish depends to a large exteüt. In the solution of the latter problem, the hydrobiologist can render considerable aid to the hydrologist. The study of the regime of a body of water through the method of biological indicators, constitutes one of the most important problems facing marine biology (Nesis, 1962a). V. Shelford et al (1935) noted. that organisms., and more especially, ecological associations, are the best indicators of hydrological conditions. The author is grateful to I.K. Avilov, A.A. Georgiev, A.A. Elizarov, G.P. Lakharov, O.A. Popova, 3

A.I. Postalakio, V.D. Rvachev, V.P. Serebryakov, I.N. Sidorenko and K.P. Yanulov or the collection of material, L454 for their aid during work at sea and for their advice and consultations. N.M. Miloslavskaya and A.A. Neiman kindy examined the manuscript and submitted valuable directions and remarks. During the identification of fauna the author made use of the dvice of Z.I. Baranova, A.N. Golikovi.E.N. Gruzov, Professor E.F. Guriyanova, G.B. Zevina, Professor A. V. Deanov, V.M. Koltun, O.G. Kusakin,,N.B. Lomakina, D.V. Ntu- mov, F.A. Pasternak, O.A. Skarlato, Ya.I. Starobogatov, Prof,. essor A.A. Strelkov, Professor P.V. Ushakov, V.V. Khlebovich and other fellow workers. MATERIAL AND METHODS . The Soviet bottom fishing industry exploits primarily the regions of Newfoundland and Labrador, northward to 56 . a lesser extent the waters of Nova Scotia and North, to Georges Bank, and makes practically no use of the Western Greenland rogina,For this reason our benthos studies were limited to the Newfoundland-Labrador region. quantitative data were derived by us. from the fourteenth (June - September, 1959) and the seventeenth (June- September, 1960) voyages of the expeditionary ship "Sevas- topol" of the N.M. Knipovich Polar Scientific-Research Ins- titute of Marine Fisheries and Oceanography. 4

P Qualitative sampleS from the Sigsbee trawle and fromoommercial-trawls were taken in 1954, and in 1958- 1960 during the voyages of the expeditionary ships "Sevast- . opol","Odessa", and "Novorossiisk", b'y A.D. Starostin, K.P. Yanulov, A.A. Georgiev, I.N. Sidorenko, G.P. -Zakharov and others (Nesis, 1962b). The equipment used to Collect the quantitative samples wap the weighted bottom scoop "0cean-50"; with a grab area of 0.25 JO. Altogether, 163'samples were taken, including 106 Samples collected during the fourteenth •crUise • and 57 during the seventeenth (fig. 1). As a rule, at each station, one s bottom sample was taken, which was then washed through. two'sieves (the bottom one with a.1 mm mesh). The samples were fixed in 4% formalin and weighed by ordinary methods. The'bottoms of the banks from 45 to 1500 metres in depth were studied. The bottom scoop 9 0 cean-50" normally functioned. very well on ailt :and silt-sand. bottoms. Since stones and boulders (ice 'deposits) are frequently encountered at all depths in the Newfoundland region, the bottom scoop did not always obtain a fully useful sample, even frora soft bottOms; stones-were sometimes stuck between the jaws. The bottom scoop worked even worse where the bottom consisted of firm sand and shells. • • ' On stony:groUnd the bottom scoop, at the very best, wauld deltver only a few'stones, so that na quantitative . analysis of the fauna could even be considered. For the determination of the mean biomass we utilized all the quantitative samples. The bottom samples, however, were not all of equal value. The distribution of the observed biomass magnitndes by stations gives a typical logarithmic curve (fig. 2), the greatest number of stations had a biomass of lees than 25 g/m2 , and very few had a high biomass. If however, we consider only those stations at which the bott= scoop took the ground soil well or satisfactorily, we get m different curve, somewhat similar in its left side to the curve of the Poisson distribution. There is a sharp (more than one and one half times) increase in the magnitude of the mean biastass, from 157 to 260 g/m2 . Eliminated •are biomass values of less than 10 g/m2 . At depths of less than 1 km the latter were observed only in those samples for which the bottom scoop functioned poorly. At only very feet stations was the bottom soil well /456 taken and therefore we can only give a sketch of the distribution of the benthos biomass, separating out the areas with a biomass of less than 100 ene and more than l'kg/m2 . The bottom relief, the nature of the bottom and the hydrological conditions of the Newfoundland and Labrador regions have been described in detail (Avilov, Elizarov, 1962; Litvin, Rvachev, 1962; Commercial reports, 1962), and in our work we will not touch upon these questions. .

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; 401

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DISTRIBUTION OF THE BIOMASS • • The highest benthos biomass in our samples occurred . on the plateau of the Grand Bank and equalled 4.6 kg/m2 of which more than 95% cOnsisted of the bivalve mollusc. Meso- desmà arctatum (infauna). In .trawl catches in the same area, masses of epifauna were found, sea mussels, cucmaria, many starfishet4;.crabs.and large, deep-burrowing molluscs, not taken by the bottom scoop. It can be considered that the overall biomass in this area exceeds 5 kg/m2 . Such a high biomass, distant from the coast (the Grand Bank plateau area is more than 400 km from the coast of Newfoundland), is observed extremely rarely, and the biomass of infauna even close

to the coast seldom exceeds 1 . - 1.5 kg/m2 (Brotskaya and Zenkevich, 1939). The biomass of infauna of over 2 kg/m2, in Terpeniya Bay, consists of Leda Dernula - 2.5 kg/m2 (Skalkin, 1960; Gur'yanova and Kobyakova,1955), off Iceland - Cylprina islandica 2.5 kg/m2 (Einarsson, 1941), on the littoral of the White Sea - nft arenaria - 2.8 kg/m2 (Pavshtiks, 1949), in the mouth of the Mashigin on the north island of Novaya Zemlya - Saxicava ârctica, plya truncata and Cardium ciliatum 3.6 kg/m2 .(Brotskaya and Zenkevich, 1939), in the North Sea - /457

- 3.9 kg/m2 , Myp - 3.4 kg/m2 , Cardium - 2.5 kg/m2 (idagmeier, 1951). The high biomass on the Grand Bank is derived from Echinarachnius parma. In Our samples, the maximum biomass of this species equalled 1.2 kg/m 2 , a value in the same order as off western Kamchatka and in the Sakhalinskiy and Karag- Lankly Bays(Gordeeva, 1951; •Pasternak, 1957;. Lus and Kuz- netsov, 1961). A high biomass of benthos is found in the north- , western part of the Atlantic, on the heights of the banks of the outer reaches of the Shelf (fig. 3). On the slopes Fig. 3, Distribution of theseneral - benthos biomaàé ' In the Newfoundland-Labrador 'reeon (our data) end on Georges BarikÀfrciiirWiey,1961Y in g/. À Labrador, B - North Newfbitndland and the north-western slope • of the Grand Newfoundland Bank, C - The Grand Bank, • except for the - north-western slope, D - Flemish Ca i5 Bank, È Banks of St. Pierre, Green,. Misaine, Bancitiereau and Càbot Strait, F - Georges Bank. - : 1 - leàs than 100,2 - 1001000,3 - more than 1000. 10 of the banks it is lower. This distribution of the biomasa /458 of the benthos is linked to the distribution of the production of phytoplankton since, beyond the limits of a narrow coastal belt, phytoplankton constitute practically the only primary source of food matter for bottom fauna. The phytobenthos can play a part as a food soùroe for bottom animals only on St. Piérre Bank where, at depths of 45 to 65 mstres, we ene., ountered undergrowths of Ptilota and other red algae. The detritus produced by the littoral macrophytes of Newfoundland, is carried away by the coastal stream of the Labrador Current. The amount of sargasso and storm-torn coastal seaweed that is swept onto the Grand Bank is not great. The link between the regions of high benthos biomass and the regions of high phytoplankton production was described by M.C. Idel'son (1934); and it was shown that the eastern coast of Murman was richer in benthos in those areas where there was a higher production of phytoplankton (Miloslavskaya, 1961). Evidently, the differences in the benthos biomass on individual banks must also be tied to the distribution of phytoplankton production. The average biohass of benthos at depths up to 300 mstres (Table 1) reaches a maximum value on the Grand Newfoundland Bank, but off Labrador and on Flsmish Cap it is minimal. There is still a lack of precise data on the annual production of phytoplankton in the north-western part of'the 11

Atlantic. However, studieS in which the distribution of the phytoplankton biomass is given e apparently, confirm our suppositions. During the period of biological spring the quantity of phytoplankton on the Grand Bank is very great, the volume of seston reaching 10 cim3 in 1 m3 . Off Labrador the seston volume is several times lower and on the Flemish Cap Bank, a significant florescence of phytoplankton has never been observed (Vladimirskaya, 1962; Movchan, 1962;, Bainbridge, Jones, 1962). The production of phytoplankton on Georges Bank is little distinguished from,the production of its surrounding oligotrophic, oceanic waters (Riley, 1941). The abundance of phytoplankton in Arctic and mixed waters is related to the high vertical stability of these waters. Weakly stratified Atlantic waters are poor in phytoplankton (Bainbridge, Jones, 1962), but it is precisely these oligotrophic (Yashnov, 1961) waters which dominate on the Flemish Cap Bank. They also periodically intrude onto Georges Bank, and in many ways, determine its hydrological regime (Bigelow, 1927; Colton, Temple, Honey, 1962). • The Arctic waters of the Labrador Current,dre tioh . in •phytoplankton, but the florescence of thiS phytoplankton occurs late, the length of the vegetation period is short, and therefore, the annual production is not great. The most favorable conditions for the development of phytoplankton occur in the waters of the Grand Bank. These waters are constituted as a result of the mixing of the heavily 'stratified 1 2

but relatively nutrient-poor Arctic waters of the Labrador Current with the waters of the continental slope, which are weakly stratified but rich in biogenous nutrient elements. The amount of light available on the Grand Bank is high, the florescence of the phytoplankton begins early and the vegetative period is reasonably long. On the eastern slope of the Flemish Cap Bank, at a depth of about 1 km, there is located a patch of high biomass. The biomass here reaches 2.1 kg/m2 , 98% of which consists of sponges (Craniella cranium, Thenea muricata and other Tetraxonida). Similar accumulations of Tetraxonida were discovered in the Kuril- Kamchatka Trench (Sokolova, 1960). ln even higher biomass of sponges (up to 3 - 4 kg/m2 ), at depths of 1.0 to 1.3 km was revealed by A.P. Kuznetsov (1960), in the Irminger Sea. All these accumulations of sponges exist under a zone of contact between Atlantic and sub-Arctic waters. Where the water masses mix, the density of the water increases and the mixed water descends to the bottom, bringing to the water layers adjacent to the bottom, a great quantitY of nuttlents from the productive upper layer. .The abundance /461, of food permits the rapid develdpment of sestonophagous sponges. The distribution pattern of the biomasS of benthos (see Table 1), demonstrates an even decrease of the biomass with dèpth. On the - Shelf, thé highest biomass of benthos is

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is nbted on the Grand Bank while at a depth of the upper bathyal.(200 - 500 m), in the northern Newfoundland region. In other areas the benthos biomass at these depths ie almost even, and approximately two times less'than it is off north-. ern Newfoundland.,

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The high biomass in the upper bathyal Zone .off the north of Newfoundland results, in the main, from the grouping of B;isaster_freçgilis - Ctenodiscils crisbatus (average. biomass, 102 g/m2 ). • The bOttom in this area, at most stations, was found to be Soft, silt and sandy silt. A. light, cyclonic turbulence . of the current-apparently favors" an increased production of phyto- - ;Plankton. Soft bottom Soils,at depth's of 200 500 metres p were 16

noted also on Flemish‘Cap Bank and in Cabot Strait (T.itvin and Rvachev, 1962), but these regions find themselves under the influence of Atlantic waters, with their relatively low productivity. Along the outer slopes of the Grand Bank, where the productivity of plankton is high, a strong current exists at depths of 200 to 500 metres, which does not permit the settling of small particles.,Here the bottom is firm and conp- osed of sand and silty sand. For the Barents Sea, the north-western part of the Bering Sea, The Kuril-Kamchatka Trench and Antarctic waters, the decrease of the biomass with depth follows a hyperbola (Belyaev and Ushakov, 1957, Belyaev, 1960). For the Grand Newfoundland Bank we drew a similar graph, utilizing logarithmic coordinates (in logarithmic coordinates a hyperbola expresses itself in a straight line) (fig. 4). Only the bio-

mass values for depths of less than 200 metres and more than , 500 metres correlatedwith this straight line, the biomuss Lita • at depths of 200 - 500 metres being far too low. The values

for the biomass for, depths of 200 - 300 metres, about 150 g/m2 , and for 300 - 500 metres, about 100 g/m2 , can be obt- 'ained by interpolation • (from the graph). Theseyalues are closer to the ones observed off northern Newfoundland. The sharp difference •in the biomass between the rich (Newfoundland and Nova Scotia) and poor (Labrador and Flemish Cap) regions, is noticeable to a depth of 500 metres, after 1 r

which it evens out. The average benthos biomass for the enlie area under study was 154 g/m 2 (the mean arithetical of all

the samples), a value of the same order es for the biomass of the Barents, Bering and Okhotsk Seas„The Grand Bank of Newfounl- land, in terms of the abundance of benthos, does not yield to the most prolific regions of the seas of the 2ar .!I'ast and can be considered as one of the most richly endowed parts of tl,e oceans of the world. In terms of biomass, the areas of Lab- rador and Flemish Cap Bank are more nearly like the poor regions of the Siberian seas (Zenkevich, 1947) or.those of tropical Africa (Buchanan, 1958; Longhursb; 1959). BIOCOENOSES The meanings of the terms "biocoenosis'; "complex" and "grouping", as appearing in the literature, differ quite widely, as various authors attribute to bhem their own various concepts of extent and status. e distinDilsh three levels of biocoenotic units, successively overlapping each other; biocoenosis, a group of biocoenoses and grounin. •The basic level of association is the biocoenosis and in fts definition we are in full agreement with V.P. Vorobtev (1949, page 155). The . unit of the highest level is the group of biocoenoses. This understanding is identical to the "iso-community" of Thorson (1957, page 104), and defines the aggregate of ecologically parallel biocoenoses, constituted in various maritime regions of the world by the various species of one and the same genus. 18

The unit of lowest rank is the grouping. This understandin is close to the concept of "complex" of V.P. Vorobtev (1949). We define as groupings the variations of a biocoenosis, developing on'various bottoms within the liMits of the one water mass. Finally, an association, identified by us on the basis only of qualitative sampling (in the absence of cuentitative), in which the interrelationship of the dominant forms is und: as a complex. ermined, we define In making deter:ninations as to the biocoenosis to which the population of a siven station mi Lt belong, we e guided by the .q.eneral complement of the fauna, the quantitativL interrelationships of the species in the bottom scoops and trawls, depth , nature of the botton and the temperature of the water. In the Newfoundland-Labrador reon we lound the following associations (fig. 5) (discussed in Lhe order of increasing depth of habitation). Biocoenosis Uesodesma arctabum - Cucunaria frondosa

lytilus &lulls, occupies the shallowest part of the Grand Bank (depth 45 - 50 metres; bottom, sand with &lens). 'ore was obs- . erved the highest biomass of benthos in the refcion (the mean biomass, without takini, into account lare forms of epifauna - about 1.7 tp 1.8 kg/m 2 ). According to the bottom scoop sample the dominzt form consisted of the bivalve mollusc 1,-esodesma arctatum (with a mean 'biomass of over 1.5 kg/m2 ); the molluscs 19

nuccinum meridionale, âipho (4oma1osipho) ventricosus and

( SLunl1Laislandica make up a mean biomass of from 5 to 20 g/m2 . To judge from the trawl samples, the area is also inhabited by masses of Cucumaria frondosa, clumps of large Eytilus edulis (one of the world's few sub-littoral populations of sea mussels, and the only one separated from the shore b such a great dist- ance), and many hydroids. The Mesodesma biocoenosis does not, apparently, have an analogue in any other part of the seas of the world, the benthos of which haà been sufficiently well studied. Grout) of biocoenoses of epifauna with Yodiola, peel Cucumaria t Balanus, Chlamys, Ophiopholis, Strongylocentrotus and others. Biocoenosis Cucumaria frondosa - Rhodophyta, discovered in the shallow waters of the St, Pierre Bank (45 - 55 metres; bottom composed of stone, shingle, shells and lithothamnium). Here only one bottom scoop station was made (biomass 590 g/m2 , including C. frondosa 483, Ophiopholis aculeata - 102, Strongylocentrotus droebachiensis - 15, bryozoans - 8) Trawl samples indicated that characteristic of the biocoenosis was an abundance of crustiform lithothamnium, Ptilota and other red • seaweed, many Hydroidea, Thelepus cincinnatus, Balanus crenatus and others: This blocoenosis, in the composition of its fauna, is close to the biocoenosis Thelepus cincinnatus - Chlamys islandica • •

/463 '!

Fg. 6. Bottom biocoenoses of the Newfoundland- • Labrador region. 1 - biocoenosis Cueumaria frondosa Rhodophyba; 2 - biocoenosis Mesodesma arctatum - Cucumaria frondosa - Menus edulis; :.3._ biocoenosis Echinarachnius parma Ammodytes americanus; 4 - biocoenosis Echinarachnius parma - Strongylocentrotus droebachiensis -Djeura sarsi; 5 - biocoenosis Astarte montagui - Nacoma caloarea; biocoenosis Ctenodiscus crispatus Actiniae;

7 • - biocoenosis Trochostoma turgidum Ctenodiscus crispatus Ophiura sarsi; 8 -biocoenosis Brisaster fragilis . - Onuphis opalina Astarte crenata whiteavesii; 9 - grouping of Brisaster fragilis -

Ctenodiscus crispatus; 10 - grouping of Brisaster fragilis - Otébiura sarsi; 11 - biocoenosis Brisaster fragilis - Astarte crenata •sulcatoides Ophiocten sericeum gracilis; 12 - grouping of Spangle Strongylocentrotuseoebachiensis •- Ophiopholis aculeataf 13 - biocoenosis Brisaster fragilis - Ctenodiscus crispatus AMphiura otteri - Pennatularia; 14 - biocoenosis Brisaster fragilis - Spongia; 15- Complex zones of mixed Labrador and subLArctic waters on the eastern slope of the Grand Bank; 16 - biocoenosis Thelepus cincinnatus - Chlamys islandica - Opiopholls aculeata; 17 -.grouping of Spongia Potamil1a neglecta Bryozoa; 18 - grouping of Spongia - .Astarte crenata whiteavesiii- Brisaster fragi1is; 19 - biocoenosis Spongia; 20- abyssal complex.

7,71 -,Ophiopholis aculeata and apparently blends directly into it, but the presence of red seaweeds determines its specificity. It is similar to the biocoenosis of the Ramose lithothamnium of the Murman coast, one of the variations of the Amphiboreal biocoenosis Modiolus modiolus (Zatsepin, 1962).Modio1us were also encountered on the St. Pierre Bank, but only in the form of empty shells. The biocoenosis C. frondosa has been identified on the Spitsbergen Bank (Iderson, 1930). A very similar grouping of Asterias amurensis - Tethvum aurantium, parts of which contain Ptilota •ectinata and other Rhodophyta and an abundance of Cucumaria japonica, has been eound off Southern Sakhalin (Gur' yanoira, 1956; Skalkin, 1960). Biocoenosis Thelepus cincinnatus - Chlamys islandica Ophiopholis aculeata flourishes amoungst biocoenoses in which E. parma predominates (depths 65 - 135, mean 80 metres), though on stony and not on sandy bottoms. The mean biomass of the biocoenosis is very low, about 33 g/m2 , but this is the result of the unsatisfactory functioning of the bottom scoop on this type of bottom. The composition of fauna in the trawl samples was reminiscent of biocoenoses in which E. panne predominates, but this sand dollar was encountered rarely, and there were few burrowing bivalve molluscs. An analgous biocoenosis with Balanus balanus, Chlotnra islandica, StronRvlocentrotus droebachensis and Ophiophol„is aculeata has been noted in the shallow water areas of the Barents Sea (Medvezhinskaya jranslator - Bear7, Nadezhdinskaa

1F0 Pe71 Kaninskaya, and Gusinaya 27.00se7 Banks, Murman coast) (Brotskaya and Zenkevich, 1939; Pergament, 1957; ._atsepin, 1962). In analagous conditions in the Far East, for example, on the stony bottoms off the island of Paramushir, there are biocoenoses with a predominance of Ophiopholis aculeata, sponges, hydroids and bryozoans (Gurtyanova, 1956; Kuznetsov and Sokolova. 1961), which are similar in composition to our biocoenosis, but are distinguished by the absence of. Chlamys islandica, which is extremely rare in Far Eastern waters. Groun of biocoenoses of Echinarachnius parma. Biocoenosls E. parma Ammodytes americanus is distr;b- uted over the Grand Bank of Newfoundland and Green, St. Pierre, Misaine and Banquereau Banks at depths of 45 - 100 metres (average depth, 69 metres), on sandy bottoms. The mean biomass of the biocoenosis was 432 g/m2 , including the flat sand dollar E. parma which gave 334 g/m2 , the sand eel Af. americanus - 76 g/m2 , and the sea urchin S. droebachiensis - 12 g/m2 . The biocoenosis E. parma is widely distributed through the Pacific Ocean basin, from the northern part of the Bering Sea to the Sea of Japan (Makarov,. 1937; Gordeeva, 1948; Gurt yanova and Kobyakuva, 1955; Kobyakova, 1959a; Pasternak, 1957; Vinogradova, 1954; Kuznetsov, 1959a; Neiman, 1960; Savilov, 1961). For all the differences between the Pacific and Atlantic /465 fauna, the number of species common to the Pacific and Atlantic 23 biocoenoses of E. Parma is reasonably large. Common, besides widely distributed circumpolar species, are also species whose range is interrupted by the Arctic, including a number of Pacific-South Atlantic forms, Biocoenosis E. parma - Strongylocentrotus droebachiensia Ophiura sarsi is an inevitable associate of the biocoenosis E. _parma. It forms a bordering ring around it on all banks at depths bf 95 - 220 metres (average depth, 146 metres), on sandy, and much more rarely, on silty sand bottome. At a depth of 225 to 250 metres it undergoes a transition into the biocoenoses of the group Brisaster fragilis. The mean biomass of tii:e biocoenosis was 149 g/m2 , made up of 89 g/m2 of E. parma, 10 g/m2 of S. droebachiensis, 9 g/m2 of Ophiura sarsi and Astarte borealis; Ophio- pholis aculeata, Onuphis conchvlega, Bryozoa, Thpifms_p_insaue- atus and Macoma calcarea contributed a biomass of. 2 to 4 g/m2 . Encountered in the trawl samples of biocoenosis, especially on the north-eastern part of the Grand Bank, on Whale Bank and off Cape Breton Island (in the zone of influence of cold currents), were many cold water species, absent from the south- western part of the Grand Bank and Banquereau Bank: Sabina sept- emcarinata t Cardiu ciliatum, Solaater syrtensiR,' Gorgonocephalus eucnemis, arcticus, Stegophiura nodosa, and besides these, masses of 0, sarsi and S. droebachiensis. Here we also found a fairly full representation of species wjth a Pacific-South Atlantic distribution.(Nesis,1962). 24

In the lower sub-littoral of the basin of the Pacific, the biocoenosis E.,-parma is replaCed by a biocoenosis predominating in Ophiura sarsi (Vinogradova, 1954; Guriyanova, 1956; Pasternak, 1957, Neiman,1960; Kuznetsov and Sokolova, 1961; Savilov, 1961). The Atlantic biocoenosis is similar to it in many respects, but in the Northwestern Atlantic, E. parma definitely predominates over other forms. Group of biocoenoses of Macoma calcarea. Biocoenosis Astarte montagui - Macoma calcarea is dist., ributed through the zone of influence of the coastal stream of the Labrador Current, on Hamilton Bank, off the shores of Lab- rador and the northern part of Newfoundland and on the north-

western slope of the Grand Bank, at depths of 100 - . 235 metres and possibly less, and in the main, on sandy and stony-sand, more rarely on silty-sand or sandy-silt bottoms. We discovered a patch of this biocoenosis in a blind hollow, "Whale Trough", at depths of 90 - 120 metres. The mean biomass was about 54 g/m2 , predominating in Astarte montagui and Macoma calcarea (mean biomass of 3-5 g/m2 ), Buccinum terrae-novae, Ophiura sarsi and others. This biocoenosis is one of the variations of the circumpolar group of biocoenoses of Macoma calcarea, known in the waters of Greenland, Jan Mayen, Spitsbergen, Iceland, The Fae- roes, north coast of Norway, the central part of the Baltic Sea, the Barents, White,Kara, Chukotsk and Bering Seas and the Sea of Okhotsk, and the Arctic waters of the Canadian archipelago 25

(Brotskaya, 1930; Idellson, 1930; Brotskaya and Zenkevich, 1939; Makarov, 1937; Zatsepin, 1962; Ivanova, 1957; Filatova and Zenkevich, 1957; Vinogradova, 1954; Neiman, 1960; Lenkevich and Filatova, 1958; Savilov, 1961; Parat, Devillers, 1936; Daniel, Mankowski, 1951; Ellis, 1960; Bertelsen, 1937; Sparck, 1933; Thorson, 1933, 1934, 1957; Vibe, 1939, 1950). • Group of biocoenoses of Ctenodiscus • Biocoenosis Ctenodiscus crispatus - Actiniae is encountered in the internal parts of underwater depressions of the North Newfoundland Shelf, displacing there the grouping Brisaster fragilis Ctenodiscus crispatus. The definition of a.precise boundary line between these two associations is difficult since the disappearance of the warm water species (Brisaster frag-

ilia, Ybldia (MeRavoldia), thracaieformis, Hitpasteria phrygiana and others), along the course of depressions penetrating into the body of the shelf, takes place very slowly, consistent with the gradual weakening of the weak flow of sub-Arctic water, intruding into the underwater depressions from the direction of the continental slope. This biocoenosis is encountered at a depth of 165 metres •on a patch of firm clay on the north-western slobe of the Grand Bank and at depths of 233 - 288 metrea, on the silt and sandy- silt bottom to the east of North Newfoundland. The mean biomass was 83 g/m2 . Predominant were Actiniae, Ctenodiscus crispatus, (mean biomass of 19 -25 g/m2 ), Stylarioides plumosus (9 g/m2) £03 and Others. At the entrances to the fiords of Newfoundland, on silt bottoms p .the fauna becomes qualitatively sharply poorer, with etiniae e etenodiScusaendalus aridleolychetes being almbst the only representatives of the macrobenthos in the trawls. 'This biocoenosis is quite similar to . the biocoencisis ot the lower Arctic, lealdane gazai, of the Murman coast (Leibson, 1939; Zatsepin, 1962), which is also developed in underwater depressions and fiords. Analagous associations have been di.sc- overed in the blind fiords of Iceland and Sweden as well (Thor- / son, 1957). Biocenosis Trochostoma turLidum - Ctenodiscus cristatus

Ophiura oars; was noted in the underwater straits between the Grand Bank of Newfoundland, Green. Bank and St. Pierre Bank, and in the deeps of the south-western part of the Grand Bank, at depths of 108 - 165 metres (average 133 metres), and on silty- sand. The mean biomass of the biocoenosis was about 95 g/m2 , composed of holothurians Trochostoma turgidum - 55 g/m 2 , 0. crispatus - 15 g/m2 and O. sarsi - 11 g/m2 . Further, there were Phascolion strombi, eternaspis scutata, Dentalium eritale, Praxillura longissima, Nephtys logosetosa, Amphithrite cirrata (mean biomass of 1 2 g/m2 ) and others. Notable is the mixture of bathyal and shallow water speees: in the straits between the banks the flow of cold Labrador water has, to a significant extent, expended its •ri force, and up to the shallows corne the warm waters of the Atlantic and their bathyal fauna, Brisaster fragilis and oth- On a much larger scale the same phenomenon is observed in the western part of the Barents Sea, in the lestern Trough, where the warm water of the northern branch of the North Cape Currnt is sucked under the cold waters of the Bear Current. In the Western Trough Z.A. Filatova (1938) noted a biocoenosis si. ilar to ours, Molpadia sp.- gtenodiscus crispaius, in whir the elements of the various complexes are mixed in the sa.:.e ,way. In the Sea of Okhotsk a holothurian of the Trochosto.A. type, together with Ctenodiscus crispabus, enters into the composition of the biocoenosis Brisaster (Savilov, 1961). Group of biocoenoses of Brisaster. This group of biocoenoses was studied by us in greater detail than ail the others (more than half of the bottom scoop , samples were taken here), inasmuch as it is adapted precisely to those depths at which the main fisheries effort for ocean perch and cod is carried out. The biocoenoses of the group Brisaster are diStributed amphiborealy and 9re to be found in the Barents, Norwegian and Bering Seas and in the Sea of Okhotsk /467 as well as in the north-western part of the Pacific Ocean (Fil- atova,,1938; Zatsepin, 1962; Faiman, 1960; -enkevich and Filat- ova, 1958; Deryugin and Ivanov, 1937; Savilov, 1961; Kuznetsov and Sokolova, 1961; Guriyanova, 1956; Gurtyanova and Kobyakova, 1955), 28

In the Atlantic B. fragilis predominates and in the Pacifie, the closely related species, B. latifrons, and more rarely, B. townsendi. In the biocoenosis B. fragilis of the Barents Sea predominant are B. fragilis, Astarte crenata, O. crispatus, Trochostoma boreale, T. tomsoni, O L_sarsj, and in the general composition, it is reminiscent of our groupine B, fragilis - C. crispatus which thrives at the saine depths and on the same types of bottom. Similar in composition Ells , are the biocoenoses of the upper bathyal of the Sea of Okhtsk, with its predominance of B. latifrons, C. crispatus and holothurians of the Trochostoma type (Savilov, 1961). In the Northwestern Atlantic the group Brisaster de y in different water masses and accordingly, makes up ten-212p different associations. Biocoenosis Brisaster fragilis - Astarte crenata sulcatoides - Ophiocten sericeum racilis is developed on the •Flamish Cap Bank at depths of 237 - 630 metres (average,339 metres), on silty-sand, more rarely, on sandy-silt. The mean biomass was 33 g/m2, of which B. fragilis gave 11 g/m2 , •Hormatia digitata (one individual which was caught in the bottom scoop by chance) - 10 g/m 2 , Astarte crenate suloatoides - 6 g/m2 and others. Of epifauna there was a striking abundance of Fontaster and Ophiocten. At depths over 330 - 340 metres, the bank is ringed by a solid zone of warm water corals, feather stars, large starifishes, Diplopteraster multipes, Solaster earlii and Tremaster mirabilis, sea lilies, Hathrometra sarsi and other 29 inhabitants of the Atlantic bathyal (Nesis, 1962b). GroupinK of Spongia - Strongylocentrotus droebachiensis gphiopholis aculeata. At depths of 145 - 333 metres (more often at less than 200 metre$),on the sandy bottoms of the Flemish Cap Bank, there develop considerable numbers of sponges (Genius, Lnyle,e Tentorium semisuberites and others) and sea urchins, Strorzylo- Q.entrotus droebachiensis, often Onyphis conchylega, PotamIlla neglecta and other Arctico-boreal and lower Arctico-boreal animals. Boreal fauna is very scanty, though the basic back- ground is the same. The mean biomass was 11 g/m2 , of which sponges made up g/m2 and S. droebachiensis and O. aculeata 1 - 1.2 g/m2 . Biocoenosis Brisaster fragilis - Ctenodiscus crispatus - Ophiura sarsi occupies the outer parts of the shelf (depths of 200 - 500 metres), in the North Newfoundland and Labrador regions, where it forms three groupings, on heavily silted bottoms, on sand and on stony bottoms. Grouping of Brisaster fragilis - Ctenodiscus crispatus is developed in the region of North Newfolmdland, at depths of 270 - 467 metres (average, 333 Metres), on sandy-silt and silt (three quarters of all the stations), more rarely on silty- sand. The mean biomass of the association was 102 g/m2 , of B. fragilis made up 38 g/m2 , C. crispatus, 14 g/m2 . A mean biomass of from 3 •to 5 g/m2 was contributed by Onuphis opalina, Psolus Ihantapusi•Maldane sarsi, Lumbriconereis spp. and others. Group of Brisester fragilis - Ophiura sarsi is identified off the shores of Labrador, in the shallows of the North Newfopndland Bank and on the northern slope of the Grand Bank, at depths . of 205 - 385 metres (average, 300 metres), on sand (79% of all the stations) or silty-sand. The mean biomass of the association - 31 g/m 2 , of which B.fragilis made up 18 g/r. Ophiura sarsi, 2 g/m 2 , complex ascidians, Onuphis conc_hylma 4 and sponges, 0.5 - 1.5 g/m2 . bryozoans Gropping of Spongia - Petamilla neglecta - Bryozoa is encountered on Belle Isle Bank and on the slope of Hamilton Bank, at depths of 340 - 350 metres, on sand containing gravel and shingle. The mean biomass - 67 g/m2 , of which various sponges made up 37 g/m2 , Potamilla neglecta, 7 g/m2 , bryozoans, 5 g/m2 , Astarte crenatà whiteavesii, Leptychaster àrcticus, Ophiura sarsi and Onuphis conchylega, 1 - 3 g/m 2 . Characteristic of the trawl samples taken in these three groupings was the mixture of \warm water and cold water elements. Complex of the zone of mixture of Labraàor and Atlantic waters. On the eastern slope of the Grand Bank of Newfoundland' the grouping of Brisaster fragilis Ophiura sarsi undergoes a transition into the complex of the zone of the mixing of Lab- rador and Atlantic waters. Recretably, from here we derived only Qualitative samples. The feline is extremely varied e - with to 100 species of the benthos apPearing even in commercial trawls.• Predominant among the cold water and warm water species are the highly eurrthermic forms,as well as some stenotherric- warm water,species. Corals.and feather stars, et depths to 50c metres, are non-existant. In places, sub-littoral forms such as Echinarachnius parma and Cyrtodaria silinua, extend to uncol:m- only great depths (400 - 500 metres), so that E. parma is ecoee- tered quite frequently in the sanie samples with Brisaster fru ilis (this was also noted for the biocoenosis Trochostoma ture 4-- idum Ctenodiscus crispatus - Ophiura sarsi, but here the B. fragilis had risen to shallow depths). The fauna preserves this character right up to the very southern point ("tail") of the Grand Bank, and even a little further west, to 5i) - 52° W. Biocoenosis Brisaster Spongia is encountered on the south-western slope of the Grand Bank and or the southern slopes of Green and St. Pierre Banks, at depths of from 150 - 200 to 500 metres. Its upper boundary rises hipher than the boundary of the preceding complex on the eastern slope of the Grand Bank. This phenomenon is related to the gmergence of Atlantic waters onto the shallows, particularly noticeable on the southern slopes of Green and St. Pierre Banks. Rere there are verr few cold water elements and more warm water forms than on the eastern slope of the Grend Bank, and to them 3;; are . added such stenothermic-warm water forrs as feather stars, several corals, lilies and others. This biocoenosis is in man- ways similar to the population of the lower levels (deeper th8n 330 - 540 metres) of Flemish Cap Bank, *though somewhat poorer in warm water species (Nesis, 1962b and c). Bioco enosis Brisaster fragilis Ctenodiscus cris -natee Amphiura otteri Pennatularia is situated in the underweter depression of the Cabot Strait (the "Laurentian Trough"), e 4 A depth of over 250 metres. The bottoms consist of clay-silt, silt, and at the edges of the depression, sandy-silt. The :—ne biomass - about 35 g/m2 , B. fragilis, C. crispatus and A. ot:, each having a mean biomass of about 6 g/mL, Astarte crenate whiteavesii, Lumbriconereis spp. (mostly L. frailis), ketin ,e and Kophobelemnon (Eukophoelemnon) stelliferurq in the order of 1.5 - 3 g/m2 . The large number of warm water species and the absence cold water forms distinguishes this biocoenosis from the grouping Brisaster fragilis Ctenodiscus crispatus, and the general Qualitative poverty and lace of variety in the fauna is the result of their existence on a very soft bottom, unfavorable for a large nnmber of species from bioceenosis'B. fragilis - Onuphis opalina - Astarte crenatawhiteavesii. Biocoenosis Brisaster fragilis - Onuphis opelina - Astarte crenata whiteavesii is developed on the continental slope in the Labrador region, in the re--, ion of North Newfound- /469 land and on the Grand Bank, at depths of 5eC 145 metres Z4

Group of biocoenoses of Spongia. Lower bathyal blocoenosis of sponges, is identified on the eastern slope of Flemish Cap Bank at a depth of 1050 - 1130 metres, on silt and sandy-silt. The mean biomass of the biocoenosis (according to two samples) was 1272 g/m2 . Sponges, mainly Tetraionida (Thenea muricata, Craniella cranium and others) a_d bryozoans, mainlY Retepora sp., determine 99 •5% of the mean biomass. Abyssal complex.We did not sample depths of over 100 metres, but on one of the stations, on the south-western part of the Grand Bank (44 14.6 N, 5359.3W), the ring-trawl toucheo the bottom at a depth of 2150 metres and brought up representatives of the abyssal complex. In the catch were four large ophiurae, Ophiomusium lymani, one each of the starfishes Porcellanaster caeruleus, Bathybiaster robustus and Pontaster foroipatus lmw., three individuals of the sea urchins Pour- talesta wandeli, •two holothurians,Yipsilothuria talismani talismani, one young representative of the feather stars ,,Pen- atula prolifera, one Dentalium solidum and several empty shells of this species, pieces of Scaphander, one example of Lumbriconereis sp. and several Phascolion. All the forms determined down to the species are characteristic of the Atlantic bathyal and abyss south of the Greenland-Canadian and Atlantic thresholds. FEEDING GROUPINGS Bottom invertebrates can be organized into several 35 feeding groups (Hunt, 1925 and others), E.P. (l'urpaev.a (194, 1953, 1954) showed that the role of the representatives of the individual feeding groupings in bottom biocoenoses is closely related to the nature of the sedimentary accumulation. Biocoen- oses, with a predominance of animals of the one feeding group, fall into zones, which change regularly with the increase of depth, throughout the whole extent of the ocean bottom, from the littoral to the ultra-abyss (Kuinetsov, 1962; Neiman, 1961a, Savilov, 1961, Sokolova, 1956,1960). We utilized, as being the most evolved, the classific- ation of bottom organisms in accordance with the character of /470 their feeding and mobility, proposed bYA.I. Savilov (1961), and established the following feeding groups: 1) sessile, attached sestonophages on firm bottoms; 2) sessile, attached sestonophages on loose bottom soils (unlike Savilov, we do not include in'this group a range of sabellid polychaetes suoh as Chone and Potamilla.and such sea lilies as Rhizôcrinus e since these are oriented to firm bottoms and reasonably strong bottom currents); 3) mobile sestonophages on firm bottoms; 4) mobile sestonophages on loose bottom soils; 5) species which gather detritus ("gatherers"); 6) forms which ingest 'bottom soils ("ingesters"); 7) preying, necrophagourand omnivorous forms. In placing a species in this or that feeding group (Table 2), we were guided by the data in the studies of Hunt (1925), . 4 -*

(averaqe 767 metres), mainly on sandy-silt ',Ind silt (about 80i0 of all the stations). On the Shelf, tl is iocoenosis is encountered only at a depth of 467 metres, in the depression separating the Hamilton and Belle isle Banks. It is absent from the deep (up to 400 - 500 metres) lonitudinal valleys separating the banks of the outer part of the Shelf from the shore. On the north-eastern slope of the Grand Bank, where th edge of the Shelf is found to be at a depth of about 275 metres (Heezen, Tharp, Ewing, 1959), the upper boundary of th's biocoenosis rises to 360 metres (at depths of 280 - 340 metre, here, the grouping B. fragilis - Ophiura saisi is noted). Th

mean biomass of the biocoenosis is about 45 vdm2, B. fragilis, O. opalina and A, crenata whiteavesii have a mean biomass of about 6 g/m2, Leda pernula, about 5 g/m2 , Lumbriconereis.sup., Heteromastqà filiformis, in the order of 1 - 3 g/m2 .

. P-rouping_of_Sponea -___4tarte crenata whiteavsii Brisaster fragilis is identified at the ve2y same depths (435 - 1480 metres) as for the basic biocoenosis, on silty-sand or sandy-silt, but with a heavy admixture of stones, gravel and shingle. The mean biomass was 54 g/m2 , of ielich various Spongia made up 16 g/m2 , A.c, whiteavesii, B. fragilis •and Didernalua, albidum, in the order of 3 - 5 g/m2 . The trawl samples of the last two associations contained many representatives of bathyal,warm aber fauna. 36

Brand (1927), Thorson . (194l), lbequest,(1949) 1 Ockelftann (1958),

Sokolova and Kuznetsov (1960), KUznetsov and Sokolova (1961), .

Savilov (1961), Sanders et al (Sanders, Mills, Hampson, 1962) - and Clark (1962). Table 2 The affiliation of the main species of bottom fauna of the Newfoundland region to •oogeogra hic.comulme. and feeding groups.

4;.Zoo- Fea- Taxonomic unit -geo. - -aff. - aff.. Phylum Spongia _ - SSF î.. Phylum Coelenterata _ - • . Class Hydrozoa _ _ Subclass Hydroidea , ' • - - SSF ' Olass Anthazoa SubolasS Octocorallia . _ _. Order Alcyonaria • - - • Eunephthya glomerata Verrill. - AB -.• P E. fructicosa (M . Sara) - AB - P ' • • Order Pennatularia - -. • Kophobelemnon (eukophobelemnon) - - • stelliferum (0.F. Muller). - BB - SSL • Subclass Hexacorallia . - _ . - Order'.Aetiniaria _ _ Hortanthia digitata (0.F. Muller) - AB - ,P . - - EewardSiidae g'. sp. . _ - P • . Order Zoantharla - 1! Order Ceriantharia - - P Phylum Nemertint. _ ..- • P Phylum Némathelminthes : _ - P Class Nematoda , -. - P,'G .Phylum Priapulida • ' ' - - Priapulus. caudatus Lam. - AB - - G 'Phylum Annelida

. Class Polychaeta . • • - •- - •Subclass Errantia _ _ Leànira tetraàona (Orsted) - BB - P ' . .Glycera capitata Orsted - AB ... G • • Goniada.maculata Orsted ... BS - P Ceratocephala loveni Malmgren - BB .4 P? .Nephthys paradoxa Malmgren - AB .4. P . • • • - AB à* P N. longosetosa Orsted N. Malmgreni'Theel _ 413 .. p • 'Eunice aff..orstedi Stimps. . ..... tB - P E.:aff. .benedicti Stimps.. ..f. BB. - p • 'Onuphis.conchylega M. Sars AB - G? 37 Table 2 .(Contid)

O. opalina Verill - LB - G? C. aff. quadricuspis Sans _ ? - G? Lumbriconereis spp. ? - I Subclass Sedsntaria ScOloplos armiger (0.F. Muller) - A3 - G Laonice cirrata (M. Sans) - AB - G Prionospio malmgreni Clap - LB - G Cirratulidae gg. Spp. ? - C Stylarioides plumosus (0.F. Muller)- AB - I Diplocirrus hirsutus (Hansen) - AB - Brada *illosa (Rathke) - AB - I • Eumenia crassa Orsted - AB - I Travisia forbesii Johnston - AB - I Ophelia limacina (Rathke) - AB - I Notemastus sp. - AB - I Hetermastus filiformis (Clap) - BB - Maldane sarsi Malmgren - AB - 1 • Praxillella gracilis (M. Sans) - AB - I P. praetermissa (Malmgren) - AB - I Praxillura longissima Arvidsson - AB - I Nicomache lumbricalis (O. Fabr.) - AB - I Owena fusiformis Delle Chiaje - AB - I • Sternaspis scutata (Ranzani) - AB - G Pectinaria (Cistenides) granulata (L) - AB - G Melinna cristata (M. Bars) - AB - G Amphicteis gunneri Sans) - AB - G Terebellides stroemi M. Sars - AB - G Pista cristata (0.F. Muller) - AB - G Amphitrite cirrata (O. F. Muller) - AB - Neoamphithrite affinis (Malmgren) - AB - P,G • Thelepus cincinnatus (O. Fabr.) - AB - $SF Potamilla neglecta (M. Sans) - AB - SSF Jasmineira schaudini Augener AM - SSL Chone duneri Malmgren - AB - $SF ' Ch. infundibuliformis Kroyer - AB - SSF • Spirorbis spp. - AB - $SF Class Archiannelida - aff. Polygordius sp. - BS - G Class Sipunculida • Golfingia aff, margaritacea (M.Sars) AB - I Phasoolion strombi (Montagu) _ AB - I Phylum Arthropoda Class Crustacea Order Cirripedia Balanus crenatus Brug. - AB - SSF Order Cumacea Diastylis goodsiri (Bell) - A - MSL 38

Order Amphipoda - - Eippomedon propinquus G.O. Sars - AB - I Amphiporeia lawrenciana Shoemaker - BS - G Haploops setosa Boeck - AB -, G Ampelisca eschrichti Kroyer - AB - G Casco bigelowi Shoemaker - BS - G Order Decapoda - - Hyas coarctarus Leach - LAB - P Pagurus bankensis Nesis, nom. nov. - - pro. P. pubescens Stimpson not Kroyer BS - P Phylum Mollusca - - Class Loricata - - Lepidopleurus asellus (Chemn.) - BS - G Tonicella mermorea (Fabr.) - LAB - P T. rubra (L.) - BS - P Class Gastropoda - - Natica clausa Brod. et Sow - AB - P Tachyrlynchus erosus erosus (Couth) - AB - MSL Arrhoges occidentalis (Beck) - AB - ML Buccinum meriodale Verkr. - BS - p B. terrae-novae Beck - AB - P B. aff. ciliatum (O. Fabr.) - A? - P B. aff. belcheri Reeve - A - P Sipho verkruzeni Kobelt - A - P S. ventricosus (Gray) - BS - P Class Scaphopoda - - • Dentalium entale L. - BB - G D. occidentale Stimpson - BB - P •Class Bivalvia - - Leda pernula (0.F. Muller) - AB - G - Yoldia myalis Couth - LB - G Y. hyperborea hyperborea (Lov,) - A - G Y. (Megayoldia thraciaeformis Storer- LAB - G Yoldiella intermedia (M. Sars) - A - G • Arca (Bathyarca) glacialis Gray - AB - MSL Limopsis minuta Philippi - BB - MSL • Chlamys islandica (0.F. Muller) - LAB - MSF Arvella glandula (Totten) . - BS - SSF •Astarte borealis (Chemn.) - A - MSL A. montagui (Dillwyn) - A - MSL A. elliptioa (Brown) - AB - KSL A. crenata whiteavesii Dall. - BD - MSL • A, crenata sulcatoides Nesis - BD - MSL A. aff. polaris pall. - A - MSL • Cyprina islandica L. - ES - MU, •Mesodesma arctatum Conrad - BS - MSL Spisula pulynyma polynyma Stimpson - LAB - MSL Gomphina fluctuosa (Gould) • - A - MSL Macoma calcarea (Chemn.) - AB - G Periploma fragilis (Totten) - A - MSL Cuspidaria glacialis G.O. Sars - A - P , Table 2 (Cont'd).

Phylum Bryozoa - - SSF Phylum Brachiopoda Terebratulina septentr ionalis Couth.- AB - SSF Phylum Echinodermata Class Crinoidea leg Rhizocrinus lofotensis M. - Sars - BB - SSE Class Asteroidea Ctenodiscus crispatus (Retz.) - AB - Leptychaster arcticus (M.Sars) - BB - I Pontaster teniuspinus hebitus Sladen- BB - Class Orhiuroidea WM, Ophiura sarsi Lutken - AB - G O. robusta (Aires) AB - G Ophiocten sericeum gracilis (G.O.Sars) BB p Ophiopholis aculeata (L.) - LAB MSF Ophiocantha bidentata (Retz.) - AB - G Amphiura otteri Ljungman - BB - A. fragilis Verrill - BB - G Class Echinoidea Echinarachnius parma (Lamarck) : LAB MSL Strongylocentrotus droebachiensis - (0.F. Muller) - AB - P Brisaster fragilis (Dub. et Mûr.) - BB - Class Holothurioidea - - Psolus phantapus (L.) Struss - AB - MSF Cucumaria frondosa Gunn • - LAB - MSY Stereoderma unisemita (Stimpson) - BS - SSF Trochostoma turgidum (Verrill) - BB - I . Hedingia albicans (Theel) - BB - I Phylum Chordata 1.1 Glass Ascidiacea Monascidia gg. spp. ? SSF Synascidia gg. spp. - ? - $SF Didemnium albidum (-Verrill) - AB - SSF Class Pisces Ammodytes americanus De Key - BS P Legend: A ..arctiq; BB , - boreal bathyal; ,BS - boreal sub-littoral; . AB - arctico-boreal; LB - lower arctico-boreal; SSF - sessile -sestonophage, firm bottoms; MSF - - mobile sestonophage, firm bottomâ;• SSL - sessile séstohophage, loose bottoms; MSL - mobile sestonoPhage, loose bottoms; G - gatherer of detritus; 1 - ingéster . of bottom soils; . P - preying, necrophagous and omnivorous forms _ 40

We determinadthe membership of a biocoenosis in a /473 feeding grouping in the following manner: we combined the data on the feed type of the le'ading form of the biocoenosis with the data on the dominance of this or that feeding grouping amongst the whole group of the leading and characteristic species of the biocoenosis (Table 3). On the chart showing the distribution of feeding ;roupu (fig. 6), we combined the feedine groupings of the sessile and mobile sestonophages of the firm bottoms, inasmuch as they aro similar ecologically (Turpaeva, 1954, 1957; Savilov, 1961). A.A. Neiman (1961a), on the basis of his own materiu'L and material available in the literature, constructed a pa-U- ern of the distribution of benthos on shelves of various.types. On wide, platform-like shelves, the upper sub-littoral is occ- upied b3/ a zone of filter-sesbonophages, followed deeper by zones in which predominant are detritus :,:eltberars and bottom soil ingesters; on the edge of the shelf there is again a zone of filterers, and on the continental slope, a zone of ingesters. In our area, we would have to designate as beine of the wide, platform type, the shelf off Labrador and the northern part of Nawfoundland. Observed here is the following pattern of the distribution of feeding zonas (see fig. 8); at the very shore, overgrowth fauna (sessile sestonophaees of firm bottoms); further out, a mixed zone of mobile sestonophages of loose bottoms and gatherers; and then a zone of ingesters. On 41

the outer edge of the shelf there is an incrIse in the ro i. of the gatherers (Ophiura sarsi) and nakjn , thr clrypearance are individual patches of sessile sestonor, s of firm sub- strata (Spongia Potamilla neRlecta - 731TozPa). Occupying the

predominant position on the slope is the zone of inee,ers. Ai of this is very reminiscent of the pattern deri.ved by 17eiman, except that not all the zones are present in a "pure form". The shelf of the eastern and soutrn rart of the Grand Bank is considerably narrower, the contleiental slope e. steep and cut into by a multitude of erater eanyons, ar' the area is tectonically active (well knovin is the ewfoundland earthquake of 1929). In genere, tHs region cohes close to being a geosynclinal •type of shelf with shallow slopes. For such shelvesà. Neiman'proposes the fellOwinçr. Pattern: the zone of the filterers of the upper sub-littoral is widened, the zone of the gatherers becomes narrower snfl is roved to a creater depth, and the zone of the ingesters is nerrovied to the point of complete disappearance.

• On the Grand Bank of 1:ewfoundland nd ;re e n and St. Pierre Banks we have, in the upper sub-littoral, a zone of sestonophages of firm bottoms or a zone of mobile sestonophaes of loose • bottoms thouqh with an increased role of Te sestonophages Of firm bottoms (cucumaria, sea mussels). Lyinc deeper is the Strongly defined zone of mobile, loose bottom sastonophages in the 1owet layers of which thare is a sllarp increase (from 4L, Role o the individual feed groupings in bottom biocoenoses /474 (in % of the overall biomass) . -Chara-Ind- . - Feed groupings -cter -ex of Biocoenoses -of bi,unif-- -SSF-SSL-MSF-MSL- G - - P -ocoen-ormity

OM. •••• - -cals Cucumaria-Rhodophyta - 16- - -958 25- SF - 7.1 Mesodesma-Cucumaria-Mytilus- - 7 - 97,8-q04-0,06- ao- MSL 7.4

Echinarachnius-AMmOdytes - 01- - - 11-7 7.7- 012- 0.1- 2011ele MSL - 4.7 Echinarachnius-Strongylo- centrotub-0. sarsi - 414- - - 2.8-6 9,1-120- I4- MSL - 3.3 Thelepus-Chlamys-Ophiopholis44,,3-.- -39i- ZZ- 44- - - 116- SF - 2.0 Astarte-montagui-Macoma • calcarea -11e- - 2 7.2-341.... 22,e-SSF:G- 1.0 Ctenodiscus crispatus- • Actiniae - - - •w-12.6-4u, 37,3- I - 2.3 Trochostoma*turgidum- Ctenodiscus-0. Sarsi - -16.5-80.1- 30- I - 4.8 •BrisastérAstarte . crénata-Ophiocten. 2Q- I.3-385- 39.ef I - 1.9 .SPongia-Strongylocentrotus, Ophiopholis -5%0- - -104- 6.5- 2 8- 2 3- SF ,- 2.3 Brlsaster fragilis- • Ctenodiscus crispatus - 411- - - 4P- 3.6-112-75.8- 2.5- I - 4.1 BrisaSter fragilis-- 'Ophlura:sarsi -. 94,7- - 12- 2.4-124-92.2- 2.1- I - 3.7 Spongia-Pot amilla7Bryozoa -78.1- 15- - 5,1- 9.3- 35- 25- SF - 4.4 Brisaster-Onuphis opalina- . Astarte crenata - C18- .36- I - 1.6 Spongia-Astarte crenata- Brisàster - -104- ?2-l9,9- 10.9- SF - 1.7 Brisaster-Ctenodiscus-

Amphiura otteri • - - - - -10.2-25.3-50.5- ap-.. I - 1.8 .Spongia -99.4 q2 - - 01- .SF - 7.8 Legend: SF - sestonophages of firm bottoms; remaini ng designations are the same as in Table 2.

*Of which AmmodyteS, 17..5% **Including one chance appearance of aflormathia, 32.0%. 0.2 to 12.7%) of the role of the detritus gathering forms. A zone, of gatherers on the shelf:is non-existant. On the upper , . , . • _ • of the continental slope there is an ablindange of .sessile, . . part . 43 firm bot.tom sestonophages, and deeper, a zone of ingesters. On the Flemish Cap Bank, the zone of sessile, firm bottom sestonophages undergoes a direct transition into a zone of ingesters (as in fig. 40 from the work of A.A. Neiman, 1961a.). The change of feeding zones on shelves Of various types in our area, is the sanie as in the Bering Sea and the Sea of Okhotsk, L 4 even though not all the zones appear in "pure form".

70 6 ss ,- • .60° ses so° 45° f de , • L.., 1 • • • I • (.• • •

4 ,-a *S •

" • • S011 — - Î.00 .500

?le rei

, :rçAet

1. 6: 61 4,

70 ° 65 50° se so° 4,5* • Fig.6distribution of feeding groupings ot benthos in' the NeWfoundland - Labrador region. 1 - sessile and mobile sestonophages of. firm bottoms; 2 Mobile sestonophages àf loose bottoms; 3 - mobi]e, loose bottom sestbno- phages and detritus gathering forms; 4 - soil ingesting forms... Let us return to the question of why the curve of the decrease of the biomass with depth differs from a hyperbola (see fig. 4). In the work of Odum and his co-authors (Odum, Cantlon, Kornicker, 1960), it was shown that the distribution of the members of any association in accordance with quantitative indicators (the number of species or the number of individuals), will have the form of a hyperbola (in logarithmic coordinates, the form of a straight line) only if the association is homogeneous. Where we have a mixture of two associations,the curve of the distribution of the,members of the association under

study will differ frbm a hyperbola (in logarthmic coordinates /476, the graph will have the form of a.broken line). Let us try to apply this situation to a region of the sea containing several biocoenoses. We can consider as a normal, that is a homogeneous •"association", a region where there is a regular change and full development, on the shelf and in the upper bathyal, of all the feeding zones (as it is observed in all areaswhere the decrease of the biomass with depth follows a hyperbola, the Barents S'a, •the north-western part of the Bering Sea, the waters of the Antarctic), that is, where there is a strong development and a high biomass of filterers, gatherers and ingesters and a corres- ponding change in the nature of the bottom of the shelf (the replacement of course grained firm ground by fine gfained soft groinad, an increased role of course grained ground on the bend of the shelf and in the upper bathyal, and then again its 45 displacement by fine grained ground on the continental slope ("Sovremennye osadki..."; "Modern sediments.", 1961). At tho same time, in regions with well defined and regular transition of trophic zones, but with a poor development of phytoplankton and a low primary production (the eastern part of the Bering Sea and the Labrador region), the belt of filterers of the upper sub-littoral has an abnormally low biomass, and the maximum biomass moves to the lower levels of the sub-littoral. In regions with a high primary production but a lowered intensity of modern sediment formation in the lower sub-littoral and the upper bathyal„ that • s, on geosynclinal shelves, the biomass at depths of 100 - 500 metres will be. unusually low ( the Grand Bank of Ngwoundland, and in part, the south-eastern part of the Bering Sea), and the zone of high biomass will change sharply to a zone of low biomass. In these areas the curve of the decrease of the biomass with depth will be the saine as for hetero- geneous associations, We do not have the data for a sufficiently full substantiation of the hypothesis presented above. The degree of the complexity of the feeding structure of the biocoenosis we characterize in the term "index of unif- 'ormity", or the dispersion of feeding groupings of a given biocoenosis related to the least of the values of dispersion for all biocoenoses (-q-) ; with the dispersions being calculated , crmin from the data in Table 3. The lower the magnitude of dispersion, the more varied is the structure of the biocoendsis. man idealiy

varied bioconosis all the feeding groupings have the sanie blomass and the dispersiin of the leeding groupings will be equal tu zero. The lowest dispersion, that is, the most varied structure, turned out to be for the biocoenosis Astarte montaomi

•acoma calcarea, the only biocoenosis wbich was related to a combined feeding group. It was accepted as the unit of measure- ment. The most uniform feeding structures were found to be in the biocoenoses Spongia, resodesma - Cucumaria - Myt,ilus and Cucumaria RhoeophTta, in which more than 95% of the biomass consisted of the representatives,of one feeding grouping. If we compare the mean biomass of the biocoenosis with the index of uniformity and the proportion of the dominant form in the overall biomass, we arrive at a very important conclusion: the higher the biomass of the biocoenosis, the more simple and uniform is its structure, and the greater the role that is played in the biocoenosis by one (dominant) species. Mean biomass of the Index of Biomass of the biocoenosis, g/m2 . uniformity dominant species

From-To Mean ( % of overall)

Under 50 30 2.0+0.32 31.4 • 50-200 92 3.4+0.50 200-1000 511 ' 5.9+1.2 78.2 Over 1000 1419 7.6+0.2 97.5 'alien the biomass of a single species constitutes more /477, than 90% of the overall biomass of a biocoenosis, it is on the way to being transformed into a monocoenosis. - Utilizing the data of V.I. Zatsepin (1962). Neiman (1980) and V.P. Vorob'ev (1949), we compared the mean biomass .of the biocoenoses with the proportion of the predominant species. It was discovered that in the biocoenoses of the Bar- ents and Bering Seas and in the Sea of Azov, that is, both in polymict and in oligomict biocoenoses, the role of the predominant species increases with the growth of the mean biomass.

A comparison of biomasses and the inJicies of uniformiy of biocoenoses, related to various feeding zones, leads to a second important conclusion: the biocoenoses of sesbonophages are richer and .more uniform than the biocoenoses of detrito- pbaaes. In the Northwestern Atlantic the meen biomass of sestonophage biocoenoses is 453 g/m 2 , and the rlean index of uniformity is 4.5 ± 0.8; for detritophage (gatherers and ingesters) biocoenoses the mean • biomass is 54 g/m2 (eirPht imes less), and the index of uniformity is 2.65 .4- 0.5 (1.7 tines less). In the Bar- ents and Bering Seas (Turpaeva, 1957; ,atsepin, 1962; Neiman, 1960, 1961a), the biocoenoses of sesbonopha-es are similarly riéher and more uniform than the biocoenoses of detritophages. The sestonophage biocoenoses flourish in zones of lowered intensity of sediment aCcumulation, where the vast majority of food particles are suspended in the layers of water above the bottom and the bottom is poor in organic matter. For this reason, food speializabion becomes a major.factor, and only one feeding group can develop; however the abundance of food leads to its riotous 40:

proliferation and the creation of a ridjpough non-diversifid biocoenosis. Characteristic of the zones of heavier accumula-Uon of sediment, besides the relative meagreness of the food, is the variety of the sources of food; organic:matter here is contained in tb.e layers of water adjacent to the bottom, on the surface of the'gimurid and in the depths of the soil. For this reason the competition between the animal forms for food has'brought about the development of feeding specializations, and a complication of the structures of biocoenoses, U.N. , Sokolova (1960) considers the approX‘imately ecual development . of the representatives of the detritophages sestohophaes and preying forms as being's characteristic feature Of zones of-more intense sediment accum ulation,-both in bathyal and.abyssal areas. A.un.iform biocoenosis, with a simple feed structure, , çan utilize rich reserves of food effectively, since the loss of- energy in a simple feed structure is not ;7;reat. However, this simplicity is accompanied a lowering of the stability of the biocoenosis, simple systems beincz less stable. As a result, •uch biecoenoses experience sharp.variationS in the biomass wlth the • onset - of unfavorable conditions. Poorer, but - diverSified biocoenoses s are significantly more stable. . As à result of our.deliberations.ue come.to the'conc- • . lusion that eXternall abiotic:factors,- the relief of the bottom, climate, the system of currents, the structure bf the water mass and the.degree of the . dynamicity of the water, determine the the evolutionary trend of sea bottom ecosystems. It eterri cL, whether this evolution will move towards bl e creation of an ecosystem, developed on the principle of the least mutual ini3r- dependence of its constituent members, stable but of relatively low productivity, or whether it will move toward the developun' of a system with a sharp predominance of one or a‘few species, unstable but hif2hly productive. The movement of the evolution of an ecosystem and itsiresults, are deterrlined by biotic factors, the interrelationships between the members of a given ecosysem and between.it and neighboring ecosystems. Inter-specific and intra-specific relationships, within the parameters of the eel- system, are the final deterMinants of the biocoenosis and, in particular, its feeding structure.

THE ZOOGEOGRAPII Y "OF '211E NORTM;EST E AT LANT I C AND THE

RELATIONSIIP OF THE BErnos To ATER MASSES /478, .The Northwestern Atlantic_is situated on th éi boundary between the Boreal. and Arctic regions. Part of the body of water studied by us belongs to the lower Arctic sub-region of the Arctic region, the sub-Arctic of Dunbar and Stephenson (Dunbar,. •

1054 and T.A.and A. Stephenson, 1954), and the ,other part„to, the American Sub,region of the Atlantic Boreal • egion (the >American.boreal,. which is the Acadian, or the Province of Nova Scotia .:Coomans, 1962) or- to the transitional sub-Arctic-bereal region (Bousfield, 1956). The aim of our zoogeographical analysis is . to accurately establish the 'boundary of the ArcticYand Boreal zones in the Northwestern Atlantic and the character Of 50

the fauna of , the transitional regio. For this we made use or the terminology of N. Hofsten (1915), and divided all animal forms.present in quantity and identified to the species (see Table 2), into groups: Arctic, Arctico-boreal. lower Arctico- boreal and boreal (sub-littoral and bathyal).

Arctic zonele 'Boreal zone • Ller Arctic Sub-Zone sub-Zone Arctico-Jporea species Arctic 1Lower Arctico-boreal species species -Y High Arctic Lower Arcticl species species Boreal species

The boundaries of the zones and sub-zones correlate well with the mid-yearisotherms: the boundary of the high Arctic and lower Arctic sub-zones with the eisotherm, the boundary of the Arctic and boreal zones with the 5° isotherm. In the bathyal, where the temperature of the water remains constant throughout the year, the eisotherm serves as the boundary for the Arctic and boreal zones (Blacker, 1957), for .the high Arctic -and lower Arctic sub-zones it is again, apparently, the eisotherm. The groups studied by us, as distinct from generally held views, characterize first of all, the relationship of the fauna to temperature, and not their range or place of origin : . 51 and so on. As a result of this kind of analysis we derive information on the landscape-geographical divisions of the body of . water under study, rather than on the genesis of its fauna. In the bottom scoop samples we discovered no high Arctic or lower Arctic species; the latter are probably few in number in any event. For the region as a whole, of particular significance are the Arctico-boreal species (see Table 4), which in general, is peculiar to those parts of the world's seas that are situated close to the boundary of the Arctic and boreal zones. Thus, In the Newfoundland region, as in the Barents Sea (Zatsepin, 1962), Arctico-boreal species predominate in the lower Arctic /479 biocoenoses. We consider these biocoenoses as being lower Arctic, and not Arctico-boreal, because the dominant species in them, even though encountered in boreal waters, are never dominant there, so that the distribution of these biocoenoses is limited by the Arctic. An increased role of Arctic species is observed in, besides the biocoenosis Astarte monta2,ui nacoma calcarea, the biocoenosis Echinarachnius parma - Strongylocentrotus droebachiensis Ophiura sarsi, located under the influence of the main stream of the Labrador Current and other cold currents. The results obtained permit the establishment of a boundary between the Arctic and boreal zones in the Northwestern Atlantic (Mize 7).

52 Table 4. The role of the representatives of separate zboeographical complexes in bottom biodoenoses, (in .% of overall biomass)

Zoogeographical -Charac- Biocoenoses complexes -teristic - AB - LAB- BB - BS -A ? -of bio- . Ma, -coenosis Cucumaria frondosa-Rhodophyta - - - 2.7-95.8- - - 0.1- 1.4- LAB Mesodesma-Cucumaria-Mytilus - - 7 0.1- - - - -99.7- 0.2- BS Echinarachnius-AmmOdytes 0.2- 3.4777.6- - -18.7- 0.1- LAB Echinarachnius-Strongylocent- rotus-0. sarsi . 6.3-24,6-65.1- - 0.1- 3.9- LAB Thelepus-ChlaMys 7Ophiopholis - 1.6-35.3-43.6- - - -19.5- LAB Astarte Montagui2MaCôma calc . area -22.7-65.2- 2.6- - - - - 9.5- LA CtenodisCus erispatus-Actiniae- 2.1-58.8- - - 1.5-39.6- - IA Troohostoilia tureidum-Ctenodis- cus70. sarsi' • - - 736.9- - -60,2- 0.7- 2.2- BB Brisaàter-Astarte Ophiocten . - - - - 0.6- BB Spongia-StrongylOcentrotus- Ophiopholis . - -14.2-10.4-18.4- - -57.0- BB Brisaster fragilis- Ctenodiscus crispatus 1.6-38.1- 2.4-52.2- - - 5.7- BB Brisaster . Ophiura sarsi - -19.3- - -70.0- - -10.7- BB Spongia-Potamilla-Bryozoa 1.1-20.1- - -11.2- - -67.7- BB Brisaster-OnuphiS opalina- Astarte crenate. 1.0-33.6- 0.7-59.3- - - 5.4- BD Spongia-Astarte crenata, Brisaster 3.5-21.3- 7 -31.2- - -44.0- BB Brisaster-Ctenodiscus-Amphiura otteri - -29.9- 1.7-61.4- - - 7.0- BB Spongia. - 047 - - 0.3- - -99.6- BD .Legend: A - Arctic; - :LA - lower Arctic; AB - Arctico-bôrear; LAB.- lower Arctico7boreal; BS boreal sub-littoral; BD - bathyal; 7 not determined to species. . .

The location of this bàundary in areas not studied by us, in the Gulf of St. Lawrence and off the coast of the Island of Newfoundland,- J.S . given in accordance with Bousfield e (1956 ). , 53

Brunel (1961b) and Hedgpeth (1957), and off Northern Labr a dor, by extrapolation, taking into acconnt bottom relief and hydrological data. As a matter of convention we show the boundary as a line, though it has the character of a fairly wide belt of mixed fauna. This belt is especially wide in the region of North Newfoundland where the boreal biocoenosis Brisaster fra- cilis - Ctenodiscus crispatus - Actiniae undergoes a smooth transition into the lower Arctic biocoenosis Ctenodiscus crispatus - Actiniae. To a significant extent, Arctic forms pene- trate the limits of lower Arctico-boreal biocoenoses (indicated by arrows in fig. 7. The lower Arctico-„boreal-srecies Echinarachnius parme, Cucumaria frondosa, Chlamys islandica and Ophiopholis aculeata are north-boreal in origin, and for this reason we,assign bio- /480 coenoses made up of these forms to the boreal zone. In this way we assign to the Arctic zone (the luwer Arctic sub-zone), the coastal regions of Labrador and North Newfoundland which are under the influence of •the coastal stream of the Labrador Current, and the remalning part of the region, to the boreal zone. On the Grand Bank of Yewfoundland, boreal species predominate at depths up to 50 metres, at the expense of sub- littoral forms, and beyond 225- 250 metres, at the expense of bathyal forms; and at intermediate depths, especially at depths /481 from 90 - 100 to 25- 250 metres, Arctic species are particularly, • • 54

* significant, while boreal snecles are almost non-existant. A completely analagous phenomenon is noted in the Bering Sea by A.A. Neiman (1961b) and off the -shores of Famchatka and the northern Mello Islands by A.P. Kuznetsov (1961). .

Fig. 7. The distribution of zoogew:raphioal complexes of bents in the Newfoundland-Labrador regio. 1 - lower Arctic biocoenoses; 2 - sub-littoral boreal biocoenoses; 3 - bathyal boreal biocoenoses.biocoenoses; 4 - lower Arctico-boreal biocoenoses; 5 - boundary of the and boreal zones (broken line for the boundary in accordance to the literature or extrapolation); 6 - region of the penetration of Arctic species into the limits of lower Arctico-boreal biocoenoses. 55 Among the sub-littoral boreal species, most are fol-ls which are distributeel only in the north-western part of the Atlantic: of the twelve boreal sub-littoral forms identified to species, shown in Table 2, such forms number nine (75). Among the bathyal boreal forms, forms endemic to the Northwestern Atlantic are considerably fewer: of 21 accurately identified

species (see Table 2), they included tIATO species, Onuphis opa- 1ina and Trochostoma turgidium (if the latter is not identical to some Eastern Atlantic trochostoma) and two sub-species of Astarte crenata. There are very few forms, endemic to the Northwestdrn Atlantic, among Arctico-borealforms (exnmples .of such are Arrhoges occidentalis and Sipho stimpsoni), and non among Arctic forms. In moving from the north to the south the difference in the shelf fauna of the western and eastern parts of the Atlantic increases, a fact which had already been noted by Loven in the middle of the last century. The endemic forms of the Northwestern Atlantic play a very small part in the benthos of the Newfoundland - Labrador Shelf, but their significance increases on the Grand Bank, and in the shallowest .parts of the Bank and off the shores of Nova Scotia they finally assume the predominant position. In the bathyal this phenomenon is expressed rather weakly. Here, to the north and to the south of the study area, the role of the forms endemic to the Northwestern Atlantic, is almost equal, and only off Nova Scotia do their numbers increase to any. extent. The reason for this is the smaller difference in the temperature regime between the northern and southern parts of the region at bathyal depths, compared to the sub-littoral. Sur- face temperatures off Central Labrador and off Nova Scotia differ, in summer, by 20° , but at a depth of 300 metres, only br 4° . The distribution of water masses in the Yorthwestern Atlantic has, until now, been studied rather poorly. Existing descriptions of the distribution of water masses were insuff- iciently detailed (L;amaev, 1960; Hachey, 1954). Vie had prev- iously published (Nesis, 1962b) some data on the distribution of water masses in the bottom layers of water for the upper bathyal region (on the basis of data on the distribution of warm water corals and feather stars). Having supplemented this with data for the sub-littoral, we present an overall picture of the distribution of water masses in the layers adjacent to the bottom (fig. 8). It will be understood that this can only be a preliminary and approximate representation. We distinguish the following types of water: 1) Arc- tic water of the coastal and main streams of the Labrador Current; 2) deep sub-Arctic waters of the Labrador circulation; 3) surface sub-Arctic waters of the Flemish Cap branch; 4) bank waters of the Newfoundland Banks and Nova Scotia; 5) modified deep waters of the continental slope; 6) deep 5'7 sub-Arctic waters mixed with the waters of the slope; 7) d-ei waters of the St. Lawrence Trough and Cabot Strait (they are distinguished from the waters of the south-western slope of the Grand Bank by a decreased level of water exchange in the bottom layers); 8) waters of the Cabot Current (freshened and relatively cold, especially in winter, coastal '-aters, flowing out of the Gulf of St. Lawrence); 9) deep and botbor waters of the North Atlantic.

On the basis of the data presented above, on the composition of. fauna and the zoogeographical affiliations of bottom associations, it is possible to establish accurately, the water mass in which one or another biocomnosis develops.

Both lower Arctic biocoenoses are found in Labrador /482, waters, as well as biocoenosis Echinarachnius parma Strongylo- centrotus droebachiensis - Ophiura sarsi of the Newfoundland regioh, with its characteristic increased role of Arctic species.

Developing in the reP. ion where deep sub-Arctic maters predom-

inate is the biocoenosis Brisaster frailis Onuphis opalina - Astarte crenâta whiteavesii. Characte3 - istic of the area, where these two water masses mix are the qroupings Brisaster Ctenodiscts crispatus, Brisaster.- Ophiura sarsi, Spongia latamilla.z_nryonaand the complex of the mixed Labrador and sub-Arctic waters of the eastern slope of the Grand Bank, which

are characterized by the simultaneors existence of warm mater

and cold water species, accompanied b: a sharp impoverishment

of the warm water complex. 58

ar se ar se gr Fig. 8. Preliminary chart of the distribution of water types in the bottom layers according to data on the distribution of the benthos. . 1 - Arctic waters of the Labrador Current; 2 - deep sub-Arctic waters of the Labrador circulation; 3 - bank waters of the Newfoundland Bank 0 and Nova Scotia; 4 - deep sub-Arctic waters mixed with the waters of the slope; 5 - modified deep waters of the continental slope; 8 - deep waters of the St. Lawrence Trough and Cabot Strait; 7 - aters of the Cabot Current; 8 - surface sub-Arctic waters of the Flemish Cap branch; 9 - deep and bottom waters of the North Atlantic. Overlapping markings indicate areas in which there is a mixing of water masses of different oritYins. Oriented to the surface waters of the 71emish Cap •branch is the srouping Spongia- StronsTlocentrotus Ophio- pholis aculeata, in which there are no cold water forms, tiough few even of warm water forms. The lower reaches of the biocoenosis Brisaster - Astarte crenata sucatoides Ophiocten sericeum :racilis is tied to the deep sub-Arctic waters with an admiYiure of the waters of the slope, and the upper, to the zone where these waters mix with the waters of the -d'lemish Cap branch. It is possible that the waters of the slope mix with deep sub- Arctic waters also on the eastern slope of the Grand Bank, at depths of over 500 metres, but we have insufficient data on this area. The Arctic waters of Labrador do noh flow on to the Flemish Cap. Bank (Nesis, 1962 a,b,c). To this is related the sharp difference in the benthos between the Flemish Cap Bank and the nearer parts of the Grand Bank. The benthos of Flemish Cap Bank is deprived of both Arctc cold water forms and the more warm water lower Arctico-boreal forms, with a Pacific - North Atlantic distribution (these are linked to bank and Lab- rador waters which do not overflow Flemish Cap). The strait between Flemish Cap and the Grand Bank consbitutes an imapass- able obstacle not only for bottom fauna; but it presents a serious Lmpediment even for the distribution of fish: Flemish Cap Bank is inhabited by a special form of deepwater redfish, (Yànulov, 1962), a special population of cod (Templeman, 1962; • Postolakii, 1962), and capelin and sand eel, abundant on the Grand Bank, are totally lacking. The waters:which wash Fle.js Cap are oceanic. To their influence is linked the abundance•f globigerina in. the bottom sediments of rlemish Cap (globiger- , - inal sediments are noted only in ocean zones ("1,10dern sed-

iMents... 1961). . • . To the bank waters of the Newfoundland Banks is oriented the biocoenosis Mesodesma arctatum - Cucumaria frondosa edulis, and possiblY the biodoenosis Cucunaria frondosa _Ihyta; here there are no cold.water forms, and many boreal upper sul2-1ittore1 forms .(Nesis,' 19(52a,b). The biocoenosis Echinarachnius parma - Ammodytes ammericanus, in the Newfoundland region, develops in the zone where bank and labrador waters tix, and Off1.7ova Scotia, in bank Waters. To the modified waters of the sloPe is linked the biocoenosis Brisaster fragilis Spongia. These:waters,.together with those of Labrador, put tbeir staiu On te.biocoenosis Trgichostoma turgidum - Ctenodiscus crlspatus ..- OPhiUra saisi, and enter the shallows of the Grand Bank•frorn- he south west. .

In the region between - the "tail" of th Grand bank and hale Trough; . there are very few cold water.forp2.s. The waters pi' the •elope engender the distinct character of the upper bathyal com- . plex of the slopes of Sable Island Bank and Banquereali Bank (Nesis,'.1962b). The waters of the Cabot Current-influence the benthos off Cape Breton Island (at depths of - less than 10 0. metres) and, to a considerably lesser extent,.onthe easterh 61

and southern slopes of Bangereau Bank (at depths of 150 -

.200 T -etres). • The pattern of the vertical distribution of waters

of various types, accordin to , the distribution of

of bottom fauna.

The re;7, ion of Labrador and »cpr -b::.rn 7ewfound1and. Depth up to 200 - 25 0 metres - Labrador waters;

from 200 - 250 to 400 - 450 metres, on tbe slope end to

//pry maximum depth on. the shelf - a zone of rixed Labrador - and'sub-Arctic.waters;

over 400 -450.metres - deep sub-rctic waters of the Labrador circulation. Flemish Cap Bank Depth up to 200 metres - surface sub-Arctic ters of the Flemish Cap branch; .

from 200 to 330 - 34- metres - a zone \ilsre these waters

mix with lower water rliasses;

over 330 - 340 metres - deep vicers of the Labrador circulation, with an admixture of the waters of the slope. Grand Bank of rewpundlend Depth up to 50 - 60 metres bank waters;

from 50 - 60 to 90 - 100 metres - a zone of mi'xin of the bank and- Labrador waters; .

from 90 -100 to 225.-250 metres - Labrador waters;

from 225-2 5 0 to 340. 7 360 metres, on te north-eastern

slope, and to 500 metres or the ,east:',;rn slope.- a zone of

• • 62

• 'mixed Labrador and sub-Arctic waters; over 540 -560 metres (and accordingly, over 500 metres On the eastern slope) - deep sub-arctic waters. On the south-western (west of :51° - 52° W) slope of the Grand Bank, at depths of 70-100 to 150-200 metres, there is a zone in which bank waters mix with Labrador waters, and Atlantic waters fiowing into the area from the south-east, and over 150- 200 metres, the modified waters of the slope. Deep waters of the St. Lawrence Trough and Cabot Strait occupy the depths, from 250 metres to the bottom of the trough (470 metres). Abyssal fauna was encountered by us at a depth of 2150 metres, but in samples taken at a.depth of about 1500 metres we found practically no abyssal forms (we had no samples from depths of 1500 - 2150 metres). Inaamuch as the abyssal complex 'is linked to the deep and bottom waters of the North Atlantic, it can be considered that the upper boundary of this water mass, coinciding here with the zero hydrodynamic surface, is located, in this region, somewhere at a depth of about 1800 - 2000 metres, which agrees with the data of G. -Ye.. Shkudova (1962), derived from the analysis of hydrological data by means of 'mathematical calculation. Closely interwoven with the question of water masses is the question of the location of vertical faunal zones. On the basis of the data on the distribution of biocoenoses, the bound- •• arles of the vertidal zones in the study area, can be draWn.in 53

the following manner: the upper sub- littoral xtends to a

depth of 50 - 60 metres; the lower sub-littoral on the Labrador-

Newfoundland Shelf, to 200 - 250 metres; on Flemish Cap, to 200 metres; on the north-eastern and eastern slopes of the Grand Bank,

to 225 - 250 metres; on the south-western slope of the Grand Bank and off Nova Scotia, to 150 - 200 metres.Between the lower bound-

ary of tile sub.;Ittoral fauna and the upper boundary of the bath-

yal fauna there is a transitional level, the thickness of which

varies from 100 - 125 to 250 - 275 metres. The upper boundary of totally bathyal fauna is located, off Labrador and North Newfound-

land, at a depth of 400 - 450 metres, on Flemish Cap,at 300'- 340 /485

metres; on the north-eastern slopp of the Grand Bank, at 340 - 380 metres; and on the eastern, and evidently, on the south-western

slope of the Grand Bank, at 500 metres. The boundary between the bathyal and abyssal zones extends along the depth of about 1800 -

2000 metres. The boundaries of the vertical faunal zones in the

Newfoundland - Labrador region coincide fully with the boundaries . of the water masses, the upper sub-littoral fauna inhabits bank and coastal waters, the lower sub-littoral in Labrador waters, the bathyal, in Atlantic waters (sub-Arctic waters and the waters

' of the slope), the abyssal, in deep and bottom . waters. A.A. Neiman

(1961b), came to the same conclusion. The distribution of waters of different origins is related to bottom relief, so that the relief of the ocean bottom indirectly determines the location of

the vertical faunal zones.However, the location of the boundaries of the water masses, and accordingly of the vertical zones, does not always accurately coincide with the lines of the major variations in bottom relief. CONCLUSIONS The mean biomass of the benthos of the Newfoundland - Labrador region (154 g/m 2 ) is of the same order of magnitude as is the biomass of benthos of such high productivity areas as the Barents Sea, the Bering Sea and the Sea of Okhotsk. The shallow of the Grand Bank of Newfoundland, where the biomass reaches 5 kg/m2 , is one of the richest areas of all the seas. The distribution of the biomass is not even. The mean benthos biomass, at depths up to 300 metres, in the "rich" regions (Newfoundland and Nova Scotia) is more than ten times greater than in the "poor" regions (Labrador and Flemish Cap). The reason for this phenomenon is the difference in the annual production of phytoplankton. The highest benthos biomass is observed on the heights of the banks, which coincides with the optimum conditions for the development of phytoplankton in local bank waters. The lower annual production of phytoplankton in more Arctic regions (Labrador) and the more oceanic areas (Flemish Cap) results, in these regions, in a lower quantity of benthos biemass. Identified in our study area were twenty biocoenoses, complexes and groupingsof benthos. A regular change of biocoenoses is observed on the vertical. Thus, on moving from the top Grand Bank eastward, to the foot of the continental slope, six biocoenoses and complexes successively displace one another. In the system of biocoenoses there is also a distinct reflec- tion of latitudinal differences: in the shallows off Labrador the bioconosis astartae-macomae predominates, and on the Grand Bank, the biocoenoses of the echinarachnae group. Differences in the nature of the bottom soil determine the existence of variou s. groupings within one and the same biocoenosis. In the bottom biocoenoses of the Northwestern Atlantic sessile, firm bottom sestonophages, mobile, loose bottom sestonophages and ingesters predominate. The distribution pattern of the zones of predominance of the representatives of various feeding groupings reflects the peculiarities of the sedimental ' accumulation, and is closely related to bottom relief. It corr- elates well with similar patterns produced for the northern part of the Pacific Ocean by A.I. Savilov, A. A. Neiman and M.N. Sokolova, which apparently indicates a regulatory mech- anism of major significance. The Northwestern Atlantic is located on the boundary of the boreal and Arctic zones. The north-western part of the Grand Bank is the region where the Arctic zone extends furthest to the south, its southernmost . point being lopated 3efurther south than the northernmost point of the boreal zone, Ice Fiord in Spitsbergen. •Such a southern location of the boundary of the boreal and Arctic zones in the Northwestern Atlantic iS the result of the cooling influence of the Labrador Current. 66

Hydrological features exert a tremendous effect on the benthos. Each water mass and each type of wator has its own conforming biocoenosis or complex of bottom fauna. The distribution of benthos clearly identifies four main water masses in the Newfoundland - Labrador reion: waters which warm the shallows in summer, Arctic waters, warm deep waters of Atlantic origin and abyssal waters. Identifiable also are various types of water, variations of main water masses and regions of mixing of the various types of water. Altogether, nine types of water are distinguishable.

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Ivanova S.S. Kachestvennaya i kolichestvennaya kharakteristica bentosa Onzhskogo zaliva Belogo morya. (.ualitative rand_gmaaliz itative characteristics of the benthos of Onega Ba of the .,hite Sea), Mater. po kompleksn. izuch. Belogo morya, Materials on the comprehensive studies of the Ibite Sea), Vol. 1, Publishing House of the Academy of Science of the USSR, 1957. Idel'son I.S. Materialt po kolichesbvennomy uchetu donnol faunv Shpitsbergenskoi banki ( -Liaterials on the cuentitative count of the bottom fauna of thee Spitsbergen Bank), Tr. morsk. nauciiii..-1711771Trudy- of the Maritime Scientific Institute), 4, 3, 1930. Idelison M.S. Raspredelenie biomassy v yuzhnoi chasti Bar- entseva morya(The distribution of the biomass in the southern Part of the Barents Sea), Tr. Gos. okeanouraf, in-ta tTrudy of the State Oceanographic Institute), 3, 4, 1934. Kebyakova Z.I.0 nekotorykh razlichiyakh donnoi fauny severnykh I yuzhnykh ostrovov Kuril'skoi gryady (Several distinctive ' features of the bottom fauna of the northern and southern islands of the Kuril Ridge), Vestnik Leningr. yn-ta. (News of the Leningrad University), No. 15, 1959 a. Kobyakova Z.I. Bentos severnol chasti Tatarskogo proliva ego •znachenie dlya pitaniea - ryb(The benthos of the northern part of the Tatarskiy Strait and its significance in the feeding of fie, News of the Pacific ,Scientific-eseerch Institute of Marine Fisheries and OceanocereWVol. 47, 1959 b. Kuznetsov A.P. Raspredelenie donnei fauny b Kronotskom zalive (The distribution of bottom feuna in Kronotskiy Bay), Trudy of the Institute of Oceanology of the Academy of Science of the USSR, Vol. 36, 1959 a. Euznetsov A.P. Raspredelenie donnoi fauny u Severnikh Kuri11/487 skikh ostrovov (The distribution of bottom faune, off the Northern Kuril IslariT,-di Ditto, 1959 b. 69

• Kuznetsov A.P: Dannye po kolichestvennomu raspredeleniyu donnoi fauny lozha'Atlanticheskogo okeana (Data on the quantitative distMbution of lottoM fauna on the Atlantic'Ocean bed), Reports of the Academy of'Science of the USSR, Vol. 130, Nb. 6, 1960. . Kuznetsov A.P. Materialy po zoogeografil prikamchatskikh vod Tikhogo okeana (Materials on the zweeography of the waters of the Pacific Ocean off Kamchatka), Reports of the Academy of Science of the USSR, Vol. • 137, No. 2 1961. Kuznetsov A.P. Nekotorye zakonomernosti struktury i raspredeleniya morskikh biotsenozov na primere biotsenozov raiona Vost- ochnoi Kamchatki I Severnykh Kuriliskikh ostrovov ( A number of regulatory mechanisms for the structure and distribution of marine biocoenoses after theexample of the biocoenoses of the Eastern. Kamchatka and Northern Kuril Voprosy ekologii, (Problems of Ecology), Vol. 4, 1962. • Kuznetsov A.P., Sokolova M.N. 0 kharaktere pitaniya i raspredeleniya Ophiopholis aculsata (1,) (The feeding and distribution of Ophiopholis aculeata (L)), Trudy of the Institute of Oceanology of the Academy of Science of the USSR, Vol.46, 1961. Leibson R.G. Kolichestvennyi uchet donnoi fauny Motovskogo zaliva uantitative count of the bottom fauna of Motovsk Bau), Trudy of the All-Union Scientific-Research Institute of Marine Fisheries and OceanotiVol. 4, 1930. Litvin V.M., Rvachev V.D. Issledovaniya rel'efa dna i gruntov promyslovykh raionov Labradora i N'yufaundlenda (Research on the bottom relief and nature of the bottom in commercial areas of Labrador and Newfoundland). Sovetsk. rybokhoz. issledov. v severo-zapadnoi chasti Atlanticheskogo okeana (Soviet fisheries research in the north-wastern part of the Atlantic Ocean), Izdvo. "Rybnoe khozyaistvo"("Fishing Industry" Press), 1962. Lus Kuznetsov A.P. Materialy po kolichestvennomu uchetu donnoi fauny v Korfo-Karaginskom raione (Beringovo more) (Materials on the •uantitative count of the bottom fauna in the Korfo-Kara.tinsk rezion Barents ea , Trudy.of the Institute of Oceanology of the Academy of Science of the USSR, Vol. 46, 1961. Makarov V.V. Materialy po kolichestvennomy ucheteu donnoi fanny severnoi chasti Beringovo i yuzhnoi chasti Chukotskogo morel (Materials-on the quantitative study of the bottom fauna of the northern part of the Bering and the southern part of the Chukotsk Seas), Research on the ses of the USSR, No. 25, 1937. • Mamaev 0.1. 0 vodnykh masse1Severnoi Atlantiki i ikh vzaimo- deistvii (The water masses of the Northern Atlantic and their interaction), Tr. Morsk. gidrofizich. in-ta AN SSSR, (Trudy of the Marine Hydrophysical Institute of the Academy of Science of •

the USSR, Vol. 19, 1960. P.:iloslavskaya N.M. Nekotorye materialy o svyazyakh bentc:sa i planktona v pribrezh'e Vostochnogo Yurmana (Material on the relationship between benthos and plankton on the coast of 'i:ast, Murman), Sb. Gidrologich. i biologich. osobennosti pribrezhnf 7h vod Murmana, (Review of the Hydrological and Biological Pecul- iarities of the Coastal Waters of rurman) vurmansk, 1961. IZ:_ovehan Kolichestvennoe razvitie fitoplanktona b vodhkh N'yufaundlendskoi banki, Flemish-Kap i prilezhashchei aloTatori (Quantitative development of Theoplankton in the waters of the Newfoundland Bank Flemish Ca. and ad'acent bodies of wata7T, Sovetsk. rybokhoz. issled. v sev.-zap. chasti Atlantich. ofena, (Soviet Fishing Industry Research of the North-Western Part of the Atlantic Ocean), Moscow, 1962. Neiman A.A. Kolichestvennoe raspredelenie bentosa v vostochnoi chasti Beringovo morya (Quantitative distribution of benthôs in the western part of the Bering Sea), "Zoological Journal" Vol. 39, No. 9, 1960. Neiman A.A. Mekotorye zakondmernosti kolichestvennogo raspredeleniya bentosa v Beringovom more (Some of the governin', factors of the quantitative distribution of benthos in the Bering Sea), "Oceanology", Vol. 1, Yo. 2 1 1981 a. Neiman A.A. Vertikarnoe raspredelenie zoogeograficheskikh kompleksoV donnoi fauny shellfa i verkhnikh gorizontov sklona v vostochnoi chasti Beringovo morya (The vertical distribution of zoogeoeraphical complexes of the bottom fauna of the shelf and upper levels of the slope in the eastern part of the Bering Sea), "Oceanology" Vol. 1, No. 2, 1961 b. Nesis K.N. Donnaya fauna kak pokazatel' gidrologicheskogo rezhima morya (na primere severo-tsentral'nogo raiona Barentseva morya) (Bottom fauna as an indicator of the hydrolop,ical regime

of the sea (after, the example of the north-central re ion of the Barents Sea)), Nauchno-tekhnich. byull. PINRC, The Scientific- Technical Bulletin of the M.N. Knipovich Polar Scientific- Research Institute of Marine Fisheries and 0eeanogenek0, No.3 (13), 1960. Nesia K.N. Tikhookeanskie elementY v bentose , SeVero-Zapadnoi Atlantiki (Pacific-Ocean elements in the benthos Of the North!- western Atlantic), SoViet Fisheries Industry Research of Uorth4Settern_Part of the Atlantic - Ocaan, tiFishing :..Pi.ess: Moscow 1962 a. 71

Nesis K.N. Sovetskie issledovaniya bentosa N'yufaundlendsko ,lio- Labradorskogo promyslogo raiona (Soviet research of the benthos of the Newfoundland-Labradoe region), Ditto, 1962,b. Nesis K.N. Korally i morskie pertya - indikatory gidrologicheskogo rezhima (Corals and fdather stars as indicators of the hydrological regime , "Oceanology", Vol. 2, No. 4, 1962,c. Pavshtiks E.A. Kolichestvennyi uchet bentosa na fatsii mya- gkikh gruntov literali Gridinskoi guby (The quantitative study of benthos on the looàe bottom soils of the littoral of Grid- inskiy.Gulfr: Raboty morsk. biol. st . Rarelo-Finsk. Gos. un-ta., Papers of the Marine Biological Station of the Karelo-Finnish State University, No. 1, 1949. Pasternak F.A. Kblichestvennoe raspredelenie i faunicheskie gruppirovki bentosa Cakhalinskogo zaliva I prilegayushchikh uchastkov Okhotskogo morya (Quantitative distribution and faunal groupings of benthos in Sakhalin Bay ald the adjacent parts of the Sea of Okhotsk), Trudy of the Institute of Oceanology of t1-0 Academy of Sceince of the USSR, Vol. 25, 1957. Postolakii A.I. Biologiya trQski ralonov N'yufaundlenda i Labradora (The biology of cod of the Newfoundland and Labrador reoion), Review of Soviet Fisheries Research in the North- Western Part of the Atlantic Ocean, "Fishing Industry" Press, 1962. Promyslovoe opisanie raionov N'yufaundlenda i Labradpra. Sostavleno V.M. Litvinym i dr pod obshchei redaksiei K.G. Konstantinova i V.M. Maevskogo (ComnerciaJ. description the Newfoundland and Labrador. region. Compiled by V.M. Litvin et al under theSeneral- editorship of nG. Konstantinov and V.M. Maevsky), Murmansk Book Publishing House, 1962. • Savilov A.I. Ekologicheskaya kharakteristika.donnykh soo- bshchestv besPozvonochnykh Okhotskoo morya (Ecological chara-, cteristics of. bottom associationiof the-invertebrates. of the - - Sea of Okhotsk , Trudy of the Institute of •OceanolOgy of the Academy of Science of the USSR, Vol. 46, 1961. .

Stalkin T.A..Bentos zalivà Ti'erpeniya, ego znachénie y pit- • aniti raspredelenii zheltoperoi kambaly (The benthos of the Bay.of Terpeniya and.its significance in the 'feeding and distribution of yellow-finned sole), Publishing House of the. . PacificSéiàntifWResearch- Institute of Fisheries and Oceanograal* .Vol.-46, 1960 • ' Sovremeilnye osadki morei i okeanov (Modern sedimentation in • • the seaS and oceans), TrUdy of the Congress Of 24 -27 - May,-•1960, :Publishing Fouse of the'ACademy of Scienc of the USSR, 1961. '72

Sokolova 1.N. 0 zakonomernostyakh raspredeleniya glubok( odnogo bentosa. Vliyanie makrorel'efa i raspredeleniva vzbesi na pishcheve gruppirovki donnykh bespozvonochnykh (àoverniqg factors of 4e distribution of deep ter benthos. The influence of macrorelief and the disrtibution of suspensions on the feeding eroupings of bottom invertebrabes), Reports of the Academy of Science of the USSR, Vol. 110,4, 1953; Sokolova V.N. Raspredelenie gruppirovok (biotsenozov) donnoi fanny glubokovodnykh vpadin severo-zapadnoi chasti Tikhogo okeana (The distribution of ,roupings (biocoenoses) of the bottom fauna of the *deep depressions of the north-estern part of the Pacific Ocean), Trudy of the Institute of Oceanolonr of the Academy of Sciences of the USSR, Vol. 34, 1960. Sokolova r.N., Kuznetsov A.P. 0 kharaktere pitaniya i roll troficheskogo faktora v raspredelenii ploskogo ezha Echinarach- nius parma Laâ. (The characteristics of the feeding and the role of the tro .hic factor in the distribution of the flat sea urchin Echinarachnius parka Lam.), "Zoological Journal" Vol.39, No. 8, 1960. Turpaevau E.P. Pitanie nekotol2ykh donnykh bespozvonochneh Barentseva morya (The feeding of several bottom invertebrates of the Barents Sea), "Zoological Journal" -Vol. 27, No. 6, 1948. Turpaqva ,, U.P. Znachenie pishchebykh vzaimootnoshenii v strukture morskikh donnykh biotsenozov (The significance of food relationships in the structure of maritime bobtom bio- UU-W7W7-àeports of the Academy of Sciences of the ussn , Vol. 65, No.1, 1949. Turpaeva 7.P. Pitanie î pishchevye gruppirovki morskikh donnykh bespozvonochnykh (The feeding and feeding groupings of maritime bottom invertebrates), Trudy of the Institute of Oceanology of the Academy of Scieries of the USSR, Vol.7, 1953. Turpaeva T.pr morskikh donnykh biotsenozov î zavis- imostt ikh raspredelenlya ot abiotocheskikh faktorov sredy (Types of maritime bottom bioconoses end the dependence of ' their distribution on the abiotic fpctors of the environment), Ditto, Vol.11, 1954. Turpaeva E.P. Pishchevye vzaimootnosheniya mezhdu dominir- uyushchimi vidami v morskikh donnykh biotsenozakh (Food inter- relationshins between the dominatin_ s oc jas in marine bobtom biocoenoses , Ditto, Vol. 20, 1957. Filatova Z.A. Kolichestvennyi uchet donnol fauny yugo-zapadnoi chasti Barentseva morya (quantitative assessment 6f the bottom fauna of the south-westernartofthe3a. the Knipovich Polar Scientific-esearch Institute of Marine 2isheries and OceanogralibbNo. 3, 1938. '73

• Filatova Z.A., Zenkevich L.A. Kolichestvennoe raspredelenie donnoi fauny Karskogo morya (The quantitativedistribution of the bottom fauna of the Kara Sea ), Trudy of the A11-Union Hydrobiological Society, Vol.8, 1957. Shkudova G.Yà. P4lozhenie nulevoi dinamicheskoi poberkhnosti v severnoi chasti Atlanticheskogo okeana (The location of the zero dynamic surface in the northern part of the Atlantic Ocean), "Oceanology", Vol.2 • No.2, 1962. Yanulov K.P. 0 guppirovkakh okunya-kr-uvacha (Sebasbis mentella Travin) Ir-Labradorsko-NlyufaundlenàsoM raione (The groupinP; of the deep-water redfish Sebastes mentella Travin in the Labrador-Newfoundland region , Reviem of Soviet Fisheries Research in the Worth-Western Part of the Atlantic Ucean. "Fishing Industr Press, 1962. Yàshnov V.A. Plankton tropicheskoi oblasti Atlanticheskogo okeana (The plankton of the tropiegt1 region of the Atlantic Ocean),. Trudy of the Maritime Hydrophysical Institute, Vol.25, 1962.

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. . . mu . . . , . V'i b e C h r. -The. marine mammals and the marine fauna In • the •Thide.,distrkt . • (Narthwett Greenland) with . observations on ice; conditions in 1939-41. Ibid., 159, . . , . . W.hi t e a v es J. F:' Catalogue of the marine' invertebrata of eastern Canada. Geol. - Survey'Canada, Ottawa, 1901. • . . . , . ' . W i g1 e y R. L..Bentic fauna of Georges Bank. - 26-th. North ''American Wildlife and • Natùral Resources Cont. Wishington, D. C,, •1961..• - . , . • .. . . . , . . , . , - .