The hadal community, an introduction

The hadal community, an introduction

The hadal* community, an introduction TORBEN WOLFF (Received 15 October 1958) Abstract~(l) During the last ten years the Danish Galathea and the Russ...

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The hadal* community, an introduction TORBEN WOLFF

(Received 15 October 1958) Abstract~(l) During the last ten years the Danish Galathea and the Russian Vitjaz Expeditions have made biological investigations in thirteen deep-sea trenches at depths exceeding 6000 m. Due to the complicated configuration of the trenches trawling is often very difficult and can only be undertaken after careful echo sounding. (2) Environmental features including extent, sediments, pressure, oxygen, salinity, temperature, and food supply were examined. (3) The quantitative composition of the bottom fauna seems to vary considerably from one trench to another, depending on the abundance of food supply in the photic zone above. Quantitative bottom samples indicate a biomass as high as 10-40 g/m 2 in certain trenches. (4) In eight of the trenches a total of 250-310 species have been found. The decrease in the number of species with increased depths within these eight trenches is smaller than might be expected (probably at the most 50 per cent). Dominant groups are actinians, polychaetes, isopods, amphipods, gastropods, lamellibranchs, and holothurians. Considering the total number of species in each group, the most important are : Pogonophora, echiuroidean worms, holothurians, and isopods. Decapod crustaceans, brachiopods, and turbellarians have no hadal representatives, and other groups (including fishes and bryozoans) are very insignificant in the hadal zone. (5) So far 127 species (and subspecies) from depths exceeding 6000 m have been identified. Of these 74 (58 per cent) are endemic to these depths. The upper limit of the hadal zone might better be set at 6800-7000 m resulting in 62 endemic species (43 per cent). As thus defined, fourteen genera and two families (of actinians and pelagic amphipods) seem to be restricted to the hadal zone. (6) Nine exclusively hadal species are found in more than one trench (two of them in three trenches), but most of these are close to one another geographically. (7) The communities in the hadal zone are listed in Appendix 3. Several species (especially of holothurians and Pogonophora) must occur in great numbers. (8) Both the benthic and pelagic hadal animals are without pigmentation and are blind, in probably all crustacean groups living at great depths a remarkable gigantism has been found, probably due to the effect of the hydrostatic pressure on the metabolism. (9) The hadal fauna is certainly derived from the abyssal zone. Endemic families, genera and perhaps also some species may be considered as relics of a preglacial abyssal and hadal fauna, but the main invasion of eurybathic species came after the onset of the first glaciation. (10) Very little is so far known of the hadopelagic fauna. The few hauls with closing nets seem, however, to indicate a high percentage of endemism. INTRODUCTION T i m i n v e n t i o n o f t h e e c h o - s o u n d e r s o m e 35 y e a r s a g o g r e a t l y i n c r e a s e d o u r k n o w l e d g e of the actual depth and the geographical distribution of the maximum deeps of the oceans. T h e g r e a t m a j o r i t y o f these d e e p s o c c u r in t h e Pacific, are m o r e o r less e l o n g a t e d t r e n c h e s , a n d l o c a t e d close to s m a l l o c e a n i c islands o r ( m o r e rarely) l a r g e r islands o r c o n t i n e n t s (cf. BRUUN, 1957, Fig. 2). *The following terms, which are essentially synonymous, have been suggested for the zone between about 6000-7000 m and the maximum depths of I1,000 m : Hadal and hadopelagic (BRUON, 1956a), "super-ozeanische Tiefen ' (ZENKEVICH, 1954), and ultra.abyssal and ultra-abyssopelagic (Z~r,rKEvacH and other Russian authors in later publications). 95

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TORBFN WOLFF

For a long time nothing was known about the possible existence of the fauna at these maximum depths. Would any organism be able to withstand the great pressures there ? FONTAINE'S (1930) experiments on tolerance to pressure made him suggest that the lower limit of life might be found at a pressure of 700-800 atm (i.e. at depths between 7000 and 8000 m). Only within the last ten years has this question been answered by sampling the fauna from the greatest depths. In 1948 the Swedish Deep-Sea Expedition on the Albatross made a haul on the east slope of the Puerto Rico Trench in the Western Atlantic at depths between 7625 and 7900 m (NYBELIN, 1951), 1600-1900 m deeper than that made at 6035 m in 1901 on a cruise of the Princesse-Alice. Then, in 1951, the Danish Galathea Deep-Sea Expedition first proved that life exists even at the greatest depths with five successful hauls between 9800 and 10,200 m in the Philippine Trench (BRuUN, 1951). Later, four more trenches were investigated (BRUUN, 1953 ; 1956b, WOLFF, 1953). Starting with a single haul in 1949 (at 8100 m in the KurileKamtchatka Trench), the Soviet research vessel Vitjaz chiefly between 1953 and 1958 carried out very extensive surveys in several trenches. The results of these are difficult to find, and almost all are published only in the Russian language. In addition to the papers on various animal groups listed below (Appendix 1), there are several very useful reports on trawling in the Kurile-Kamtchatka Trench (ZENKEVICH, et aL, 1954, 1955 ; ZENKEVICH, 1954). In more recent results I am only aware of two papers : BIRSTEIN,et aL (1958) on trawling in 1957 in the Mariana Trench, and BELJAEV, et al. (1958) on trawling in the Tonga Trench, also in 1957. All mention of other work carried out by the Vitjaz since 1955, cited in this paper, is derived from two sources : (1) Location, depth, temperature and station numbers for animals from depths exceeding 6000m, chiefly from the Trudy Instituta Okeanologii. (2) Diagrams, illustrations and preserved animals in the Russian deep-sea exhibit during the International Congress of Zoology in London in 1958. I take this opportunity to thank the Russian scientists who arranged this display and allowed me to copy and publish their preliminary details for those interested in deep-sea biology. These sources of information have made it possible to make a first attempt at a general survey of animal life in the deep-sea trenches and of the hadal community. TRAWLING TECHNIQUE In trawling at the greatest depths (KULLENBERG,1951 ; 1956), it is first necessary to know how much wire to pay out for a given depth at the desired speed of the gear along the bottom to make certain that the trawl just reaches the bottom. The length of wire can be calculated as a function of the wire angle on leaving the ship, the diameter, the resistance of the trawl to the water (depending on the type of gear), and the actual depth. It took 5-6 hrs to pay out the 12 km long steel wire, reaching bottom some 5 km behind the ship. Since most trenches are narrow and uneven in configuration, it is essential to have an echo-sounder recording continuously. Before any trawling can be undertaken there must be days of painstaking sounding to obtain a reliable survey of the bottom topography. Other difficulties may also be encountered. For instance, during the early trawling of the Galathea in the Philippine Trench all factors (wind, current, waves and swell) which served to force the ship off its course came from the east at fight angles to the north-south direction of the trench. Since the deepest part of the trench (roughly

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10 km) is a narrow plain only 1-3 km wide, it was necessary to head into the current to keep directly over the even bottom, while trawling north or south along the trench. The slopes were so steep that the trawl would be lost if it hit them. Fortunately, the west and east slopes were found to differ so that with the continuously recording echo-sounder, the position of the ship at any given moment could be l o c a t e d - and the course altered accordingly. There are similar problems everywhere in trawling in narrow trenches (i.e. BmSTEIN, et aL, 1958). THE ENVIRONMENT OF THE DEEP-SEA TRENCHES The following environmental features seem to be those of greatest importance for life in the trenches. Extent. Although depths exceeding 6000 m are distributed throughout the oceans, the total area amounts to only 1.2 per cent of the oceans, and is very insignificant compared to areas with depths between 3000 and 6000 m occupying 76 per cent of the oceans and more than half the surface of the globe. Sediments. So far there has been no thorough geological study of the bottom sediments in the various trenches, but, as in abyssal depths, they are a soft, fine, more or less clayey ooze with oxidized clay and diatom and radiolarian oozes as the main components. Moreover, the presence of large and small jagged stones of littoral origin (BRUUN, 1953, 1956b, p. 181) seems to be a characteristic feature of trenches in areas with frequent earthquakes and eruptions of submarine vulcanoes and are indicative of turbidity currents. These not only supply the deepest water masses with oxygen (and food .9) from above, but also may cause occasional catastrophes to the trench fauna, resulting from mud-slides. Together with stones the great abundance of waterlogged lumps of pumice in southern trenches such as the Kermadec Trench (WOLFF, 1953, Fig. 16) should also be mentioned as a substratum for attached animals such as hydroids, Stephanoscyphus and Scalpellum. Pressure. FONTAINE(1930) and several other physiologists have shown that increased pressure has a m a r k e d effect on the metabolism and ability to survive in littoral animals which are experimentally submitted to pressures of several hundred atmospheres. These experiments, and especially the demonstration (ZOBELL,1952 ; ZOBELL and MORITA, 1959) of barophilic bacteria in the deep-sea trenches seem to indicate that organisms from the greatest depths have special physiological adaptations to determine their upper limit of occurrence. Also the lack of certain groups of marine animals (i.e. Crustacea Decapoda) at greater depths is most probably due to the effect of the hydrostatic pressure. Oxygen. All measurements of the oxygen content in the trenches have shown sufficiently large amounts to support animal life. For instance, the Galathea found about 4.5 ml/1 in the Kermadec and Tonga Trenches (BRUNN and KIILERICH, 1955). It is not yet known, however, as to whether there is a decrease of oxygen in the water layers immediately above the bottom. Salinity. The slight differences in salinity so far found in the trenches (34.7%o + 0.1%o) is certainly without ecological significance. Temperature. The bottom temperature of the Aleutian, the Kurile-Kamtchatka, the Riu-Kiu, the Philippine, the Banda, the New Britain, the Kermadec and the Sunda Trenches (Appendix 3) vary between 1.2°C (6200 m in the Kermadec Trench) and 3.6°C (7300 m in the Banda Trench). There is an increase of temperature towards

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the bottom, due to adiabatic heating (BRUUN and KIILERICH, 1955, including bibliography). Food Supply. BRUUN (1956a, 1957) and KRISS (1959) have pointed out that food in the abyss should be considered as any energy stored in organic matter, which can be utilized by heterotrophic bacteria which thus provide an important source of food for many deep-sea animals. On account of the great scarcity of abyssopelagic and hadopelagic life (VINOGRADOV,1954), the amount of food available in the trenches is probably of the same order of magnitude as in abyssal depths. Both faunas are ultimately dependent on primary production in the photic zone above*. The presence in some trenches of great numbers of Pogonophora whose peculiar mode of extra-oral feeding probably requires a fairly high concentration of food, as well as the extraordinary gigantism of certain hadal animals (see p. 103) suggest that in many trenches the amount of food available may be even greater than at abyssal depths. THE BOTTOM FAUNA Quantitative Composition So far, very few quantitative bottom samplings have been made at depths exceeding 6000 m. In spite of twelve lowerings with a 0.2 m 2 Petersen grab during the Galathea Expedition, only seven were successful, with living animals in five. The deepest sample (the Philippine Trench, 10,120 m) contained only one specimen, a small holothurian, Myriotrochus bruuni (weighing 0.1 g). The richest samples were taken in the Banda Trench and contained the following species : 6580 m

1 Ancistrocyllis constricta (Polych.) 3 Tharyx multifilis ,, 1 Travisia profundi ,, 1 ? Owenia lobopygidiata ,, 1 Leptanthura hendili (Tanaid.) 2 Harpinia spiircki (Amph.) 2 Cuspidaria sp. (Lam.) 1 Ceraplectana trachyderma (Holoth.) 8 species, 12 specimens. Total weight : 2.2 g.

7270 m 2 Macrostylis hadalis (Isop.) 1 Harpinia spdrcki (Amph.) 2 Cuspidaria sp. (Lam.) 3 species, 5 specimens. Total weight : 2.5 g.

The Vitjaz also took bottom samples with a 0.25 m ~ modified Petersen grab in the hadal zone (Aleutian, Kurile-Kamtchatka and Japan Trenches). A rich sample from the Aleutian Trench at 7246 (7286?)m depth contained a large specimen of the echiurid worm Vitjazema aleutica, weighing 10.1 g, and other invetebrates and protozoans weighing 0.1 g. Other bottom samples yielded species of Macellicephala and Macellicephaloides, Jakobia birsteini and two species of Elpidia, one of which was E. glacialis (ZENKEVICHand FILATOVA,1958). During their deep-sea trawling, the Russians attached a special 'trawlograph' to the sledge trawl (Agassiz trawl) to measure the trawling distance which permits a rough estimate of the quantitative distribution of animals brought up in the trawl. *Examples of the~ food sources arc : dissolved or colloidal organic matter, the waterlogged plant debris (from hugo trunks of tr~s down to suspendod particles), tho excromvnt of animals living between the surface and the bottom, and the enormous number of chitinous integuments shod by many moulting animals, especially crustaceans.

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By means of this technique the biomass of both the zooplankton and the benthos of the Kurile-Kamtchatka Trench and the adjacent part of the Pacific has been estimated (ZENr~VlCH, et al. 1955, ZENKEVlCHand BmSTEIN, 1956). The biomass of the trench between 8330 and 9950 m was found to be 0.3 g/m 2.

Number of Species Based on all the literature known to me, on the data of the exhibit in London (mentioned above) and on the Galathea collections I have tried to make as comprehensive a list as possible of all animals so far recorded from depths exceeding 6000 m (Appendix 1). If we exclude all localities with only scattered records of single species (i.e. most Russian records from 1955 onwards), approximately the following numbers of species of the most important major groups were taken from the Kurile-Kamtchatka, Philippine, Banda, New Britain (Galathea records only), Kermadec, Sunda, Mariana and Tonga Trenches (in the two latter trenches the deepest trawling only) : No. of species Porifera : Coelenterata : Hydrozoa : Scyphozoa : Anthozoa : Nemertini : Nematodes : Polychaeta : Echiuroidea : Sipunculoidea : Priapuloidea : Crustacea : Tanaidacea : lsopoda : Amphipoda :

8-10 23-30 4 1-2 t8-23 1

1-3 50-55 7-9 4--5 1

58-65 3 28-30 20-25

No. of species Pycnogonida : Mollusca : Gastropoda : 15-20 Lamellibranchiata : 25-30 Bryozoa : Echinoderma : Asteroidea : 8-10 Ophiuroidea : 6- 7 Holothurioidea : 25-35 Pogonophora : Enteropneusta : Tunicata : Pisces Total

2 45-55

1 43-55

5 I

2 3-5 250--310

Vertical Distribution within the Hadal Zone There is less of a decrease in the number of benthic species within the hadal zone (Table 1) than might be expected, but, on the whole, this is very uniform from one group to another. It is only slight for the molluscs and greatest for the echinoderms. The reason for the latter may be the non-hadal status of most crinoids, sea-stars and brittle-stars. A somewhat larger decrease in the number of species seems to occur in the Kurile-Kamtehatka Trench (ZENKEVICH, 1955 ; ZENKF.VlCHand Bn~STEIN, 1956) than in the other trenches investigated. It should be noted that the difficulties in trawling successfully increase with increasing depth and that the risk of animals washing out of the bag or the bucket in the cod end of the trawl on the way up increases with greater depths.

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The maximal recorded depths of the various marine groups is given in Appendix 2. Table 1. Number o f species in each major group between 6000 and 10,700 m, for intervals o f 1000 m, based on the same data as in the preceding section Depth range (m)

60007000

70008000

No. of hauls

9

6

80(0)9000

9000I0000

1000010700

6

4

5

Number of spectes m 1-2 7-8 Porifera* 2 7 7 4 13-14 Coelenterata 7 16 17 6 27 Polychaeta 1 4--5 2 1 6-8 • Gephyrea ' 6-7 12-13 26 18 4 Crustacea 4 20 13 9-10 9 Mollusca 5 10-11 7 31 13 Echinoderma 3 3 Pogonophora I 5 2 1 6 Others 26-27 65-70 136-140 74 35 Total number Calculated number, if nine 47-49 111 98-105 79 136-140 hauls in all *BELJAEVet al. (1958) record a fragment of skeleton of a hexactennelid sponge from the Tonga Trench (10,687-10,415 m). Since the authors do not, however, include it in the list of the total number of species from this trawling I have also omitted it in Table 1 and in Appendix 2.

Dominant and Insignificant Animal Groups The dominant groups in the trenches are coelenterates (actinians), polychaetes, crustaceans (isopods and amphipods), molluscs (gastropods and lamellibranchs), and echinoderms (holothurians) (cf.list on p. 99 and Appendix I). If the number of species occurring deeper than 6000 m is considered as a function of the total number of species in each major group, the following four groups are the most important : Pogonophora, Echiuroidea, Holothurioidea and Isopoda. Russian scientists have estimated that the following percentages of the total number of species of these same groups are found deeper than 6000 m (Russian exhibit, 1958): Pogonophora 30 per cent, Echiouroidea 9 per cent, Holothurioidea 1.7 per cent, and Isopoda 1.4 per cent. Deeper than 8000 m the following percentages, respectively, 27, 4, 0.8 and 0.2, are found. It is also evident (list on p. 99, Appendix 1) that some major groups, which play an important role in littoral, bathyal and even abyssal depths, are totally missing or very insignificant in the hadal zone. Decapod crustaceans, brachipods and turbellarians have never been recorded from depths exceeding 5300 m and fishes, tunicates, cirripeds, bryozoans, and sponges have only a few representatives at depths greater than 6000 m. Exclusively Hadal Species and Endemism Although it is premature to draw any definite conclusions, it appears that the true hadal animals (ciphers in bold type in Column 1, Appendix 1 and WOLFF, 1959) only occasionally are found at depths of less than 6800-7000 m. At present, they only total 62 species* (49 per cent) of the 127 identified species (and subspecies) *When the Russian results are worked out, this number will probably be increased two or three times.

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of Metazoa (Appendix 1). If, however, the upper limit of the hadal zone is set at 6000 m (rather than 6800-7000 m), there are 74 such endemic species (58 per cent of the total number of species found deeper than 6000 m). The number of species which are exclusively hadal (deeper than 6800-7000 m), in the various major groups, already worked up, are as follows (with the percentage of all identified trench species in each group given in brackets) : Hydrozoa : Acfiniaria : Polychaeta :

2 5 8

(67%) (84%) (36%)

Echiuroidea : Tanaidacea : Isopoda : Amphipoda :

4 2 15 9

(80%) (67%) (68%) (56%)

Crinoidea : Asteroidea : ~ Ophiuroidea : Echinoidea : Holothurioidea : Pogonophora :

1 (10%)

8 3

(50%) (50%)

Earlier it was stressed that isopods (WOLFF, 1956a, p. 150) have a greater number of species restricted to the trenches than other groups of animals represented by quite a number of species in the hadal zone, such as polychaetes and echinoderms, i.e. are more stenobathic*. One, or probably two, endemic hadal families have been described (Galatheanthemidae Carlgren, 1956- Actiniaria- 2 species; Vitjazianidae Birstein & Vinogradov, 1955 - Amphipoda - probably exclusively hadopelagic, 1 speciest). Several genera (with the number of species in brackets) are endemic in the hadal zone, namely : Actiniaria

:

Galatheanthemum Carlgren, 1956 [2] Hadalanthus Carlgren, 1956 [1]

Crustacea

:

Parascaphocalanus Brodsky, 1955 [ 1] Zenkevichiella Brodsky, 1955 [1] Herpotanais Wolff, 1956 [ 1]

Polychaeta :

Macellicephaloides Ushakov, 1955 [3] Echiuroidea : Jakobia Zenkevich, 1958 [1] Vitjazema Zenkevich, 1958 [2]

Bathycallisoma Dahl, 1959 [1] Bathyschraderia Dahl, 1959 [1 ] Princaxelia Dahl, 1959 [1] Echinoderma :

Vitjazaster Beljaev, 1958 [1] Hadalothuria Hansen, 1956 [1] Pisces :

Pseudoliparis Andriashev, 1955 (subgen.) (2) Two copopod genera Parascaphocalanus and Zenkevitchiella are probably hadopelagic (see p. 124). *This general picture has not been altered by later Russian additions to our knowledge of the hadal fauna. #Taken twice with closing net, once between 6000 and 8500 m and once between 4200 and 7800 m; further information about the hadopelagic fauna is given below (p. 104).

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A few more genera are generally hadal, but have a single abyssal or abyssohadal species: Alomasoma Zenkevich, 1958, Hirondellea Chevreux, 1889, Bathyspinula Filatova, 1958 (subgen.), and Heptabrachia Ivanov, 1957.

Distribution in the Various Trenches A certain number of exclusively hadal species occur in more than one trench :

Macellicephala hadalis : Philippine, Bands and Kermadec Tr. Jakobia birsteini : Kurile-Kamtchatka and Japan Tr. Vitjazema ultraabyssalis : Kurile-Kamtchatka and Idzu-Bonin Tr. Storthyngura herculea : Aleutian, Kurile-Kamtchatka and Japan Tr. Storthyngura vitjazi : Kurile-Kamtchatka and Idzu-Bonin Tr. Bathycallisoma schellenbergi : Kermadec and ?Puerto Rico Tr. Hirondellea (Tetronychia) gigas : Kurile-Kamtchatka and Philippine Tr. Pardaliscoides longicaudatus : Philippine and Kermadec Tr. Myriotrochus bruuni : P~lippine and New Britain Tr. Careproctus amblystomopsis : Kurile-Kamtchatka and Japan Tr. Some of these trenches (Aleutian, Kurile-Kamtchatka, Japan and ldzu-Bonin Trenches) are rather close to one another and are connected at depths of about 6000 m (BIRSTEIN, 1958). Others, however, (Philippine and Kermadec Trenches) are widely separated. It is of utmost interest to ascertain, whether the number of hadal species common to two or more trenches will increase or not when the fauna of the other trenches becomes known. In considering the distribution o f hadal species in the various trenches (WOLFF, 1956a, p. 150) it was noted that isopods and amphipods had a considerably higher number of species in the Kermadec Trench than in all the other trenches investigated by the Galathea, whereas the opposite was the case with echinoderms and polychaetes. This dominance ofisopods and amphipods in the Kermadec Trench might be explained by the fact that most of the abyssal species from which the hadal species may have originated occur at high latitudes*. Thus, of the trenches considered, the Kermadec Trench is the most easily accessible from the antarctic region. This theory is supported by the recent record of no less than five species of isopods from the deepest part of of the neighbouring Tonga Trench (see Appendix 1).

The Hadal Community In describing a community it is most important to give a comprehensive list of the species in the particular locality and to ascertain which species act as predators, prey, food competitors, etc. Very little information of this sort is available from the hadal zone. Between 6000 and 11,700 m, only 57 successfult hauls (trawling and bottom sampling) have yet been made, and in the true hadal zone (beyond some 7000 m) the number is reduced to about 38. For completeness I have listed all trawls and bottom samples (Appendix 3) taken at depths greater than 6000 m, which are presently known to me. Data for many of the Russian trawls, etc. have been gathered from several Russian sources. In papers by various authors inconsistencies were found in *At any rate this appears to be true for the isopods. t" Successful" here means touching the bottom and bringing up one or more bottom-living animals.

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recording the depth and geographical position at the same station or stations. In such cases the most probable or most frequently used figures were entered in Appendix 3. Reference is made to all animals, so far recorded from each station. In all cases where sutficient data is available, the total number of species and specimens from the station is included. The number of Foraminifera species has been recorded by the Russians in a number of cases. These are subjoined to the number of metazoan species (Appendix 3). The number of specimens of Foraminifera, however, are not included. The number of metazoan species from each station varies cofisiderably with the depth and more especially from one trench to another. It is largest in the KurileKamtchatka Trench (up to 45) and the Kermadec Trench (up to 31), This almost certainly is due to an especially rich fauna in these two trenches, compared to the others investigated. It should, however, be borne in mind that owing to the bottom configuration trawling is, for instance, much more easily carried through in the Kermadec than in the New Britain Trench. The number of specimens also varies greatly. In some places the number especially of holothurians must be enormous compared to that of other animals. Thus, in the Sunda Trench at a depth of 7160 m the Galathea collected ca. 3000 Elpidia glacialis sundensis and at 8200m in the Kermadec Trench ca. 1800 E.g.kermadecensis. Similarly, the Vitjaz brought up from 9000 m in the Kurile-Kamtchatka Trench ca. 1000 Elpidia glacialis and almost 2000 specimens of the pogonophores Zenkevitchiana longissima and Spirobrachia beklemischevi. Two remarkable photographs were taken by H. E. EDGERTON at 7500 m in the Romanche Trench, (EDGERTON, et aL, 1957; COUSTEAU, 1958). They s h o w - i n addition to an easily recognisable brittle-star-bottom animals which probably can be identified as sea-anemones or stalked barnacles as well as amphipods or isopods.

Special Peculiarities of Hadal Animals Morphologically the hadal species show the same adaptations to life in eternal darkness as many abyssal and even bathyal (and cave-dwelling) species : greyish or whitish colours and in the hadal zone probably always total blindness. Gigantism in Crustacea was first noted for almost all hadal species of isopods collected by the Galathea (WOLFF, 1956a). These isopods are larger than any other representative of the genus to which they belong. This is also true for tanaids (WOLFF, 1956b), the gigantic hadal Herpotanais kirkegaardi being the largest tanaid ever recorded. The hadal mysid Amblyops magna is the largest species of the genus and a hadal species of Haplomesus (isopods) is two-three times longer than the abyssal representatives of this genus (ZENKEVICHand BIRSTE!N, 1956). Large-sized gammarid amphipods were recorded abysso- and hadopelagically (BmsTEXN and VINOGRADOV, 1958). A correlation was found between the depth and size of the hadal species of the isopod genus Storthyngura from the north-west Pacific, but not between their dimensions and temperature (BmSTEIN, 1957). The great variety in food habits and source of food in groups of crustaceans showing deep water gigantism indicate that also the food factor certainly can be excluded. It is therefore most probable that this characteristic phenomenon is caused by the effect of hydrostatic pressure on the metabolism, as suggested by ZENKEVICH and BIRSTEIN (1956) and BIRSTEIN (1957).

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TORBEN WoLff

Gigantism has not yet been reported among the hadal representatives of any animal groups other than Crustacea.

Origin of the Hadal Fauna There is little doubt that hadal animals are derived from the general abyssal fauna which have many features in common. Since the relatively sudden onset of the Glacial Age must have resulted in a fundamental temperature change in abyssal and hadal depths of from some 10° to below 4 °, only the most eurythermic and eurybathic species could survive. The hadal fauna may have two origins : (1) taxons higher than subspecies or perhaps species and of exclusively hadal occurrence are relics of a preglacial abyssal and hadal fauna ; (2) a new invasion of sufficiently eurybathic species has taken place since the beginning of the first glaciation and is probably still going on (BRtmN, 1956a). The hadal representatives of the family Neotanaidae which are probably archaic and which are presently almost entirely abyssal and hadal (WOLFF, 1956b) are examples of the first type. kIadal species of some isopod genera (i.e. Macrostylis and Storthyngura) whose closest relatives live in cold water at rather high latitudes in both hemispheres, the most probable place of origin for an invasion of the hadal zone (WOLFF, 1956a), are examples of the second type. Within deep-sea Malacostraca, there is a dominance of ancient groups in the abyssal and hadal regions (BIRSrEIN, 1958). The ancient fauna with seemingly endemic genera and even families, it should be noted, are not in agreement with the generally accepted idea that most deep-sea trenches are recent, geologically speaking (BmSTEIN, 1958). They can, however, have migrated from the abyssal zone into the trenches after their formation and later have become extinct in the abyssal zone. THE HADOPELAGIC FAUNA Thus far, only the Vitjaz has operated closing nets below 6000 m. One haul made in 1953 in the Kurile-Kamtchatka Trench between 6000 and 8500 m revealed an astonishingly rich fauna : twenty species of copepods, four species of ostracods and five species of amphipods (Appendix 4). One of the amphipods is moreover the only known representative of a new, probably hadopelagic family, the Vitjazianidae. One or more hauls between 6000 and 8000 m were made in 1957 in the New Britain Trench (VINOGRADOV, 1958) and with species (probably new) of the genera Parargissa, Andaniexis and Hyperiopsis, fairly closely allied to hadopelagic species of the same genera from the Kurile-Kamtchatka Trench (BmSTEIN, 1958). Moreover, three abyssohadal hauls with closing nets have been made in the KurileKamtchatka Trench (4190-8050 m, 4200-7800 m, and 4295-6550 m) and the amphipods in them enumerated (BIRSTEIN and VINOGRADOV, 1955). According to ZENI~VIC~ and BIRSrmN (1956) the hadopelagic animals are consistently unpigmented while the abyssal plankton is usually characterized by an intense dark red colour. This seems to indicate vertical migration by the abyssopelagic, but not by hadopelagic animals which are completely isolated from the illuminated water layers above.

Acknowledgments--A considerable number of people have helped with this survey. These include Russian colleagues who at the exhibit in London gave me much help and advice for which I wish

The hadal community, an introduction

105

to thank them. I am also grateful to Professor J. A. BIRSTEIN,Moscow, who answered questions and helped with some references. The following Danish specialists have kindly supplied me with additional information: E. WESENBEgo-LUNO(echiurids), J. KmKEGAARD(polychaetes), J. Knudsen (molluscs), F. J. MADSEN (echinoderms) and A. KIILERICH(hydrography and echo sounding). Last but not least my thanks are due to the leader of the Galathea Expedition, Dr. A. F. BRUUN for fruitful discussions on many aspects of the hadal fauna.

Zoological Museum University of Copenhagen Copenhagen, Denmark REFERENCES ANDRIASHEV A. P. (1955) On a new Liparid fish from a depth of over 7000 m. Trud. Inst. Okeanol. 12, 340-344. In Russian. BELJAEV G. M., VINOGRAOOVAN. G. and F1LATOVAZ. A. (1958) Trawling at a depth of 10.5 km in the Tonga Trench. Dokl. Akad. Nauk SSSR 121, No. 1, 74-77. In Russian. B1RSTEIN J. A. (1957) Certain peculiarities of the ultra-abyssal fauna as exemplified by the genus Storthyngura (Crustacea Isopoda Asellota). Zool. Zh. 36, 961-985. In Russian; English Summary. BIRSY~IN J. A. (1958) Deep-sea Malacostraca of the north-western part of the Pacific Ocean, their distribution and relations. 15th Int. Congr. Zool. Sect. III, Paper 33, 1-4. BIRSTEIN J. A., SAVILOVA. I. and UDINZEV G. B. (1958) Trawling at the maximum depths of the ocean. Priroda 1958, No. 3, 70-71. In Russian. BmSTEIN J. A. and TCHINDONOVA J. G. (1958) The deep-sea mysids of the north-western part of the Pacific Ocean. Trud. Inst. Okeanol. 27, 258-355. In Russian. BIRSTEIN J. A. and V1NOGRADOV M. E. (1955) The pelagic gammarids of the KurileKamtehatka Trench. Trud. Inst. OkeanoL 12, 210-287. In Russian. BIRSTEIN J. A. and VINOGRADOV M. E. (1958) Pelagic gammarids from the north-western part of the Pacific Ocean. Trud. Inst. Okeanol. 27, 219-257. In Russian. BmSTEIN J. A., VrNOORADOV M. E. and TCHJNDONOVA J. G. (1954) Vertical zonation of plankton of the Kurile-Kamtchatka Trench. Dokl. Akad. Nauk SSSR 95, No. 2, 389-392 In Russian. BRODSKV, K. A. (1955) The fauna of Calanoida in the Kurile-Kamtchatka Trench. Trud. Inst. Okeanol. 12, 184-209. In Russian. BRUUN A. F. (1951) The Philippine Trench and its bottom fauna. Nature, Lond. 168, 692. BRUUN A. F. (1953) Dybhavets dyreliv. Galatheas jordomsejling, Copenhagen, 153-192. BRtJUN A. F. (1956a) The abyssal fauna : its ecology, distribution and origin. Nature, Lond. 177, 1105-1108. BRUUN A. F. (1956b) Animal life of the deep-sea bottom. The Galathea Deep Sea Expedition 1950-52, London, 149-195. BRUtrN A. F. (1957) Chapter 22. Deep Sea and Abyssal Depths. Geol. Soc. Amer. Mem. 67, No. 1, 641-672. BRUtJN A. F. and KHLERICH A. (1955) Characteristics of the water-masses of the Philippine, Kermadec, and Tonga Trenches. Deep-Sea Res., Suppl. to 3, 418-425. CARLGREN O. (1956) Actiniaria from depths exoeeding 6000 metres. Galathea Rep. 2, 9-16. COUSTEAU J. Y. (1958) Calypso explores an Undersea Canyon. Bobbing on 5½ miles of anchor line, the famed ~eseareh vessel probes the secrets of the Atlantic Romanche Trench. Nat. Geogr. Mag. 133, No. 3, 373-396. DAHL E. (1959) Amphipoda from depths exceeding 6000 metres. Galathea Rep. 1. In press. EDGERTON H. E., COUSTEAtJ J. Y., BROWN J. B. and HARTMAN R. H. (1957) Nylon rope for deep-sea instrumentation. Deep-Sea Res. 4, No. 4, 287. ELIASON A. (1951) Polyehaeta. Rep. Swedish Deep-Sea Exp. 2, (Zool. 11), 129-148. FAGE L. (1956) Les Pycnogonides du genre Nymphon. Galathea Rep. 2, 159-165. FILATOVA Z. A. (1958) Some new species of bivalve molluscs from the north-western part of the Pacific Ocean. Trud. Inst. Okeanol. 27, 204-207. In Russian. FONTAINE M. (1930) Reeherehes exl~rimentales sur les r6aetions des &res vivants aux fortes pressions. Ann. Inst. Ocdanogr. g, 1-100. GISL~N T. (1956) Crinoids from depths exceeding 6000 metres. Galathea Rep. 2, 61-62. HANSON, B. (1957) Holothurioidea from depths exceeding 6000 metres. Galathea Rep. 2, 33-54. IVANOV A. V. (1952) New Pogonophora from the far eastern seas. ZooL Zh. 31, No. 3, 372-391. In Russian.

106

TORBEN WOLFF

IVANOV A. V. (1957) Neue Pogonophora aus dem nordwestlichen Teil des Stillen Ozeans. Zool. J. 85, (4/5), 431-500. KmKEGAARDJ. B. (1956) Benthic Polycheata from depths exceeding 6000 metres. Galathea Rep. 2, 63-78. K.RAMP P. L. (1956) Hydroids from depths exceeding 6000 metres. Galathea Rep. 2, 17-20. K.RAMP P. L. (1959) Stephanoscyphus (Scyphozoa) Galathea Rep. 1 173-185. In press. KRms A. E. (1959) Microbiology and the chief problems in the Black Sea, Deep-Sea Res. 5, No. 3, 193-200. KULLENBERG B. (1951) On the shape and the length of the cable during a deep-sea trawling. Rep. Swedish Deep-Sea Exp. 2, (Zool. 2), 29-44. KULLENBERG B. (1956) The technique of trawling. I n : Galathea Deep Sea Expedition 1950-52, London, 112-118. First published in Danish (1953). KOEHLER R. (1909) Echinodermes provenant des campagnes du yacht Princesse-Alice (Ast6ries, Ophiures, Echinides et Crinoides). Rds. Camp. Sci., Monaco 34, 1-317. MADSEN F. J. (1953) Holothurioidea. Rep. Swedish Deep-Sea Exp. 2, (Zool. 12), 151-i73, G6teborg 1955. MADSEN F. J. (1956) The Echinoidea, Asteroidea, and Ophiuroidea at depths exceeding 6000 metres. Galathea Rep. 2, 23-32. MILLAR R. H. (1959) Ascidiaeea 1, 189-209. In press. MURINA V. V. (1957) Abyssal Sipunculids (genus Phaseolion Th6el) of the north-western part of the Pacific, collected by the Vitjaz Expeditions in 1950-55. Zool. Zh. 36, (12), 1777-1791. In Russian. NORDENSTAMilk. (1955) A new isopod from the deep sea. Rep. Swedish Deep-Sea Exp. 2, (Zool. 16), 205-212. NYBELIN O. (1951) Introduction and Station List. Rep. Swedish Deep-Sea Exp. 2, (Zool. I), 3-28. PASTERNAK F. A. (1958) The deep-sea Antipatharia of the Kurile-Kamtchatka Trench. Trud. Inst. Okeanol. 27, 180-191. In Russian. RASST. S. (1958a) Deep-sea fishes of the Northern Pacific and far-eastern seas. 15th lnternat. Congr. Zool., Sect. 111, Paper 34, I-2. RAss T. S. (1958b) Fishes of the greatest depths. Priroda 47, No. 7, 107-108. ROULE L. (1913) Notice pr61iminaire sur Grimaldichthys profundissimus nov. gen., nov. sp. Poisson abyssal recueilli b. 6.035 m~tres de profondeur dans l'Oc6an Atlantique par S.A.S. le Prince de Monaco. Bull. Inst. Ocdanogr., Monaco No. 261, 1-8. SCHELLENaERG A. (1955) Amphipoda. Rep. Swedish Deep-Sea Exp. 2, (Zool. 14) 181-195. SPXRCK R. (1956) The density of animals on the ocean floor. In : The Galathea Deep Sea Expedition 1950-1952, London, 196-201. First published in Danish (1953). STSCHEDRINAZ. G. (1958) The Forminifera fauna of the Kurile-Kamtchatka Trench. Trud. Inst. Okeanol. 27, 161-179. In Russian. TARASOVN. I. and ZEWNA G. B. (1957) Cirripedia Thoracica from the seas of SSSR. Fauna SSSR., Rakoobraznie 6, No. 7, 1-268. In Russian. USHAKOVP. V. (1950) Polychaetes from the Sea of Okhotsk. Isslyed. Daljnevost. Morei SSSR No. 2, 140-234. USHAKOV P. V. (1952) Investigations on the deep-sea fauna. Priroda 1952, No. 6, 100-102. In Russian. USHAKOV P. V. (1953) New genus of polychaetes of the family Phyllodoeidae (Polychaeta). Trud. Inst. Okeanol. 13, 20%209. In Russian. USHAKOV P. V. (1955) Aphroditidae (Polychaeta) of the Kurile-Kamtehatka Trench. Trud. Inst. Okeanol. 12, 311-321. In Russian. VJNOGRAOOV M. E. (1954) The vertical distribution of the biorrmss of zooplankton in the Kurile-Kamtchatka Trench. Dokl. Akad. Nauk SSSR 96, No, 3, 637-640. In Russian. VINOGRADOV M. E. (1958) On the vertical distribution of d,;~e.;ea plankton in the west part of the Pacific Ocean. 15th Int. Congr. Zool., Sect. III, Paper 31, 1-3. WOLFF T. (1953) Galatheas undersogelser i dybdegravene. Naturens Verden 37, 121-149. WOLFF T. (1956a) Isopoda from depths exceeding 6000 metres. Galathea Rep. 2, 85-157. WOLFF T. (1956b) Crustacea Tanaidacea from depths exceeding 6000 metres. Galathea Rep. 2, 187-241. WOLFF T. (1959) La faune hadale ou faune des profondeurs sup6rieures ~t 6000-7000 m~tres La Terre et la Vie, Paris, 106. In press. ZENKEVICH L. A. (1954) Erforsehungen der Tiefseefauna im nordwestlichen Toil des Stillen Ozeans. In : On the Distribution and Origin of the Deep Sea Fauna. Un. Int. Sci. Biol., B, No. 16, 72-85.

The hadal community, an introduction

107

ZENKEVICH L. A. (1958a) The deep-sea echiurids of the north-western part of the Pacific Ocean, 2 part. Trud. Inst. Okeanol. 27, 192-203. ZENKEVICH L. A. (1958b) Certain aspects of ocean depths studies. Paper at CSAGI (Comit6 Sp6ciai de l'Ann6e G~ophysique lnternationale) Meeting in Moscow July-August 1958, 1-8. (Mimeo.). ZENKEVlCH L. A. and BIRSTEIN J. A. (1956) Studies of the deep water fauna and related problems. Deep-Sea Res. 4, No. 1, 54-64. ZENKEVICH L. A., BIRSTEIN J. A. and BELJAEV G. M. (1954) Exploration of the fauna of the Kurile Kamtchatka Trench. Priroda 1954, No. 2, 61-74. In Russian. ZENKEVtCH L. A., BIRSTEIN J. A. and BELJAEV G. M. (1955) Investigations of the bottom fauna of the Kurile-Kamtchatka Trench. Trud. Inst. Okeanol. 12, 345-381. In Russian. ZENKEVICH L. A. and FmATOVA Z. A. (1958) General characteristic of the quantitative distribution of the bottom fauna in the north-western part of the Pacific Ocean. Trud. Inst. Okeanol. 27, 154-160. In Russian. ZoBELL C. E. (1952) Bacterial Life at the Bottom of the Philippine Trench. Science 115, 507-508. ZOBELL C. E. and MORtrA R. Y. (1959) Deep-Sea Bacteria. Galathea Rep. 1, 139-154. In press.

1

1

4 I

many < 50

6861)'-'7230 8100

9000[9050

6860"7230 9000-9050 6860-7230 7210-7230 6860-9050 7210-7230 8100

9995-10,002 10,687-10,415 6860-7230 8100 7210-7230 6860 8100 6860-8100 8100 7210-7230 8100 6861) 7210-7230 9000-9050 7210-7230

Depth (m) (in trench)

6620[7000 Kermadee Sunda 6730 6860-8660 Kur:KamL I 10,687-10,415 Tonga [ 6860

Kermadec Tonga Kur.-Kamt.

Trench(es)

BRUNN, 1'953 J, ZENK. et at., 1955 BELIAEVet al., 1958

Exhibit 1958 BEJ-JAEVet at., 1958 STSCHEDRINA, 1958

Recorded by

Vit.

Gat,

Vit.

Research vessel

-

!

--

--

-

(3) (I) (i)

(1) -

(3)

--

Total no. o f localities

--

In one trench Depth range only [ outside trenches

*As far as yet unidentified species are concerned, the lowest taxon (genus, family, etc.), which was recorded, has been given in the table. Sometimes only the number of a higher taxon (for instance Porifera) was recorded from a certain locality; in such cases the number of species is given after the name of the group.

1 FORAMINIFERA I (5 spp,) 2 II (4 spp.) 3 Ammobaculites agglutinans (d'Orbigny) 4 Ammodlseus sp. 5 Astrorlza sp. 6 Cyclammina sp. 7 Glomospira gordialls (Jones & Parker) 8 Haplophragmoides glomeratus Brady 9 Haplophragmoides nltidus (Go~s) 10 Haplophragmoides sphaeriloculus (Cushman) lI Haplophragmoides sp. 12 Hormostna globuttfera Brady 13 Hyperammina elongata Brady 14 Hyperammina sp. 15 Millollna sp. 16 Nodosinum sp. 17 Recurvoides sp. I 18 Recurvoides sp. II 19 Reophax bactllarts Brady 20 Reophax bllocularls Flint 21 Reophax nodulosus Brady 22 Reophax sp. I 23 Reophax sp. II 24 Saccammina difflugiformis Brady 25 Thurammina compressa Brady 26 Thurammlna papillata Brady 27 Tolypammina sp~ 28 Trochammlna sp. 29 Trochaminella bullata H6glund 30 PORIFERA I (3-4 spp.) 31 ,, II (?) 32 llI (3-4 spp.) 33 ,, IV (1 sp.) (?) 34 Hyalonema sp.

No. o f specimens in trenches

List of bottom animals from depths exceeding 6000 m, their number and depth range in the trenches, depth range in other localities and total number of records. Species printed in bold types indicate those known exclusively from depths exceeding 6000 m (in a few cases 5850 m). The exclusively hadal species and subspecies (from depths exceeding 6800-7000 m) also have their cipher (to the extreme left) printed in bold type*.

APPENDIX

O

Z

Oo

Stephanoseyphus simplex Kirkptr., 1890

Ntephanoscyphus Sp~

41

42

[W[

72

67 68 69 70 71

Ampharetidae gen, I (2-3 spp.) Ampharetidae gen. II (1 sp.) Ampharetldae gen. III (1 sp.) Amphictei, s, raederi Pnoenkova, 1929

Ammotrypane galatheae Kirkegaard, 1956 Ammotrypane sp,

64 POLYCHAETA I (9-12 spp.) 65 II (I sp.) 66 ,, III (2 spp.)

ANTHOZOA 43 HEXACORALLIA (1 sp.) 44 ACTtNIARIA (2-5 spp.) 45 . Sp. 45 Bathydactylus kroghi Carlgren, 1956 47 Daontesia mlelehei Carlgren 1956 48 Galatheanthemum hadale Carlgren, 1956 49 Galatheanthemum prolundale Carlgren, 1956 50 Galatheanthemidae gen. I (1 sp.) 51 Galatheartthemidae gen. H (1 sp.) 52 Galatheanthemidae gem I l I (1 sp.) 53 Galatheaathemidae $en. IV (1 sp.) 54 Hadalanthus knadseni Carlgren, 1956 55 Paredwardsia lemehei Carlgren, 1956 56 CERIANTHARIA (2-3 spp.) 57 ANTIPATHARIA (1 sp.) Bathypathes patula Brook, 1889 58 59 OCTOCORALLIA (I sp,) 60 Umbellula sp. (1-2 spp.) 61 NEMERTINI ,(I sp.) 62 NEMATODES I (1 sp.) 63 ,, II

COELENTERATA HYDROZOA (1 sp.) Ag~aia(D galatheae Kramp, 1956 Aglaopheala tenuissima Bale, 1914 Cryptolarella (?) sp. Halisiphonia galatheae Kramp. 1956 SCYPHOZOA (1 sp.)

35 36 37 38 39 40

6 -e. 60 1 1 c. 20

2 1 1 12 7 -1 (ma/y

1

1 e. 10 13 12 1 50 c. 120 -2 --3 40 3 2 2

1

c. 160

----2

No, of specimen in trencba

Banda New Britain Kur.-Kamt. Sunda Kermadec Kur.-Kamt,

Kur.-Kamt. Tonga New ,, Britain

Kermadec Kur.-Kamt. Tonga Kermadec

Banda Philippine Kermadec Aleutian Kur.-Kamt. Japan Mariana Kermadee Sunda Kur.-Kamt. Sunda Kur.-Kamt.

,t

Kermadec

Kur.-Kamt,

Tonga Sunda Kermadee Kur,-Kamt. Kermadee Kur:Kamt. "Banda Kermadee

Trench(es)

APPENDIX

8610-8660 68609050 666O-677O 8210-8300 7290-7250 9820-10,210 5850-8300 7246 7210-7230 6156-6570 10,630-10,710 6660-6770 7160 6860-7230 7000-6900 8632-8779 8610-8660 6180-6730 7210-7230 I0,687-10,415 9995-10,002 6620-6730 6810-9050 10,68%10,415 8980-9043 8940 7290-7250 8980-9043 7210-8430 7000-6900 6660-6770 8100

7210-7230 7O00-6900 6660-6770 6860 8210-8300 686O 6490-6650 6180-7600 9995-1~002

Depth (m) (in trench)

1- - c o n t d ,

USWAKOV, I950

KIRKEGAARD, 1956 Exhibit, 1958 ZENK. et al., 1955 KmKEGAARD, 1956

ZENK. et al., 1955 BROUN, 1953 PASTERNAK,1958 ZENK. et aL, 1955 Bauur~, 1953 ZENK. et aL, 1955 BEUAEV et al., 1958 E~dbit,, 1958 BRUUN, 1953 Z~2qK. et al., 1955 BELIAEV et al., 1958 Exhibit. 1958

CARLGREN, 1956

Exhibit.,'1958 ZENK. et aL, 1955 Exhibit. 1958

CARLGREN, 1956 ,)

ZENK. et al., 1955

Exhibit., t958

) KI~Mp, 1959

ZENL et aL, 1955

~'aL ~;t.

&t

~t

Vit.

dk

Vit.

Gal, Vit.

d'~l. ~t.

),

Vit.

GaL

Vii. Gal, Vii.

Z~NK. et al., 1955 KRAMe, 1956

Vit, Gal.

Research vessel

BF,L1~.V et aL, 1958 IOtAMP, 1956

Recorded by

,,,,,,,

+

+ F + ÷

88-1366

1331-5303

9 (l) (2) (1) (1) (l) (I) (3) (1) (1) (1) (1) (I) (2) (I) (1) 2

1

(2)

1

1

3 6 (t) (I) (2) (1)

1 1

(t) (3) (I)

38

-430-5860

1

(1)

--

(1)

1

3

293-385

(1)

Total no. of localities

--

--

In one ] trench [ Depth range ____°nly t outside trenches

5"

f~

t-

O

~r

Kesun abyssorum Monro, 1930

Lumbriconereis sp.

Macellicephala abysslcola Fauvel, 1913

Maeellicephala hadalis Kirkegaard, 1956

Macellieephala mirabtlis MeIntosh, 1885 Macellieephala zenkeviehi Ushakov, 1955 MacellicephaIa sp. I Macelllcephala sp. II Macellicephaloides grandictrra Ushakov, 1955 Maeellicephaloides ven'ueosa Ushakov, 1055 Macellieephaloides vitia~ Ushak0v, 1955 Maldanella harai (Izuka, 1902) Maldanidae gen, (I sp.) Myriochele sp. Nephthys elamellata Eliason, 1951 Nephthyidae 8en. (1 sp.) Nereis proftmdi Kirkegaard, 1956 Nino#fusca Moore, 1911 Notomastus sp. Owenla lobopygidiata (?) Ushakov, 1950 Petaloproctns ctrratus (?) Monro, 1937 P01ynoidae 8en. (1 sp.) PotamUla sp. Poecllochaetus sp. Sabellidae sen. (1 sp.) Serpulidae gen. (1 sp.) Terebellides eurystethus Chamberlin, 19t9 Terebellidae gen. (1 sp,) Tharyx multO~lls Moore 1909 ~tavisia profurtdi Chamberlin, 1919

80

81

82

83

84 85 86 87 88 89

105 106 107 108 109

104

97 98 99 100 101 102 103

96

91 92 93 94 95

90

79

Ancistrocyllis constricta Southern, 1921 Cirratulidae (Cossura ?) Cirratulidae gen. (i sp.) Hesionidae (I sp.) llyphagus bythlneola Chamberiin, 1919 Jasmineira sp. I dasmineira sp. II

73 74 75 76 77 78

I I 2 3 5

37 10 4 9 18 5 toasty 4 I 3 I

c. 90 3 2 49

I

1

31 2 2 36 2

1

11 4

1 1

2

1

4

1

2

1

No. of specimens in trenches

Pto. Rico Banda

Tonga Kur.-Kamt. Tonga Kur.-Kamt. Kermadec

Banda

New Britain Banda Kermadec

Kermadec Kur.-Kamt. Kermadcc

Mariana Pto. Rico Kur.-Kamt.

Banda Kermade¢ Tonga New Britain Sunda Banda Kermade¢ Sunda Kermadec Kur.-Kamt. Aleutian Banda Kermadec Sunda Banda Kermadec Philippine Kur.-Kamt.

Trench(es)

APPENDIX

}

KIRK£GAARD, 1956 Exhibit. 1958

6580 9995-10,002 10,687-10,415 8980-9043 6730 7240-7290 6620-8300 6730 6960-8300 8100 7246 7290-7250 6620-7000 7160 7290-7250 6660-8300 10,150--10.210 7210-7230 8100 10,630--10,710 7625-7900 8100-9950 7210-7230 7210-8430 6620-6770 7210-7230 6180-8300 6180-7000 8980-9043 7290-7250 6620-7OOO 8210-8300 6490-6650 7240-7290 10,687-10,415 8100 10,687-10,415 7210-7230 6620-6730 6660-6770 7625-7900 6580 6490-7290 ELmSON, 1951 KIRKI~GAARD,1956

Gal.

Aih

Gal.

Vit.

Vit. Gal.

Exhibit. "1958 KIRKF~AARD, 1956

Exhibit., "1958 USHAKOV, 1952 Exhibit., 1958 ZENK. et oL, 1955 KIRKEGAARD, 1956

Vit. Gal.

dh

Vit.

21h.

i,

Vit.

dh.

Vit.

Gal.

Gal. Vit.

Research vessel

i KIRK E~AARD, 1956 ZENK. et al., 1956 KIRKEGAARD, 1956

! USHAKOV, 1955

I US~KOV, 1955 , Exhibit., 1958 I ELIASON, 1951

] ZENK. et aL, 1955

USHAKOV, 1952 i Exhibit., 1958 KmKI~GA.4a~D,1956

"

KIRKEGAARD, 1956

~p

Recorded by

Depth (m) (in trench)

l---contd. trench only

~n one

975--4064

1-590

1410-4750

t10-1366 36-1046

4060

4255-4600

146--4700

13-1366

4255-4872

300-1470

3436

c. 5

7 6

(!)

4 (I) 3 5 (1) (1) (1) (1) (I) 3

1

2 13 (1) (3) 6 (1)

1

(1) (I) 3

1

12

(1)

(1) (2) 6

2

(1) (1) (l)

2

Total no. Depth range outside trenches of localites

O

X

Alomasoma nordpacifica Zenkevieh, 1958

Jakobia birsteini Zenkevich, 1958

Vitjazema aleutlca Zenkevich, 1958

115

116

117

Phascolion latenre Selenka, 1885

Scalpellum (Arcoscalpellum) japonicum Hock, 1883

Scalpcllum sp, CUMACEA Diastylis sp. TANAIDACEA Apseudes galatheae Wolff, 1956 Herpotanais kirkegaardi Wolff, 1956 Neotanais serratisplnoses hadalls Wolff •956 ISOPODA I (5 spp.) ,, II __ IIl Antarcturus ultraabyssai~ Birstein, n. sp. Bathyopsurus nybelini Nordensta~, 1955

126

127 128 129 130 131 132 133 134 135 136 137 138

CIRRIPEDIA

ARTHROPODA CRUSTACEA 124 COPEPODA HARPACTICOIDEA t25 OSTRACODA

121 Ptuzscolion pacificium Marina 1957 122 PRIAPULOIDEA (1 spO 123 Prlapulus sp.

120

118 Vitiazema ultraabyssalis Zenkevieh, 1958 119 SIPUNCULOIDEA (2-3 spp.)

Aloma~om ehaetifera II (1-3 Zenkevich, spp') 1958

113114

I10 Teavista sp. Ul Vitiazia dogieli Ushakov, 1953 112 ECHIUROIDEA I (1 sp.)

1

5 c. 10 3 8

4

2

i

1

160

c. 170 150

(,0 2

2

65

~.24

i

Ik

Trench(e,)

APPENDIX

Kermadec New Britain Kur.-Kamt. Pro. Rico

Ton~

Kur.-Kamt. Japan NW Pae. Kermadec New Britain Sunder Kermadee . ,,

Kermadee New Britain

7210-72307587

Ku't.-Kamt.

6860 6156--6207 6096 6620-7000 8006 7160 8928-9174 6660-6770 6960-7000 6960-8300 10,687-10,415 9995-10,002 8980-9043 7210-7230 7625-7900

9995-10,002 7657 (?)

7246 7587 8100 6156-6570 7246 7210-9950 9715-9735 6860 6156-6207 6860 6272-6282 6860 6156-6207

6860-8430 8100 10,150-10,210 6660-8300 7246 6860-8430

Depth (m) (in trench)

l--contd.

Ph~ppine Kermadec Aleutian Kur.-Kamt. Aleutian Japan Kur.-Kamt. Japan Aleutian Kur.-Kamt. Idzu-Bonin Kur.-Kamt. Japan Ku~.-Kamt. NW Pac. Kur.-Kamt. Japan

Kur,-Kamt.

vJ ,L

Recorded by

,

ZENK, et aL, 1955

,.

NORD~NST,~LM, 1955

BELJAEV et aL, 1958 Exhibit,, 1958

BRutm, 1953 Exhibit., 1958 BRUIm, 1953 Exhibit., 1958 WOLVF, 1956

I T A ~ 9 5 ~ and Z e r o ^

I

Exhibit., t958

Exhibit., 1958 ZENK. et aL, 1955

~ MMIIII~'' 19571957

!}

ZENKI~VlCH, 1958a

Zm'qK. et aL, 1955 U$~KOV, 1953 BRuin% 1953

,

Gal. vit. GaL

%

Vit.

Vii.

Research vessel

+

+

7

5500-5987

w

508-5500

5345

3568-5830

5560

520-4560

Depth range outside trenches

3

1

2 (1) (1) (1)

1 1

(3) (l) (1) (I)

I0

(i) (I)

(1) (1)

3

13

(1)

(l) 7

3

9

(3)

Total no. of localities

6"

8

8

D" o

Storthyngura chelata Birstein, 1957

Storthyngurafurcata Wolff, 1956

Storthyngura herculea Birstein, 1957

Storthyngura pulchra kermadecensis Wolff, n. subsp. Storthyngura tenuispinis kurilica Bit'stein, 1957 Storthyngura ten. tenuispinis Birstein, 1957

Storthyngura vitjazi Birstein, 1957

154

155

156

157 158

160

Bathyceradocus stephenseni Pirlot, 1934 Bathyschraderia magnifiea Dab.l, 1959 Eusirus bathybius Schellenberg, 1955 Harpinia spitrefli Dahl, 1959 Hirondellea dubia Dahl, 1959 Hirondellea (Tetronyehia) gigas, Bst. & Vinogr., 1955 Lepechinella wolffi Dahl, 1959 Metandania islandica Stephensen, 1925 Onesimoides cavimanus Pirlot, 1933 Orchomenella abyssorum (Stebbing, 1888)

167

Pardaliscoides Iongicaudatus Dahl, 1959

Phoxocephalidae (1 sp.)

174

175

170 171 172 173

169

167 168

165 166

Bathycallisoma schellenbergi Dahl, 1959

163

161 A M P H I P O D A I (5-7 spp.) 162 ,, II (1 sp.)

159

151 152 153

150

Eurycope galathene Wolff, 1956 Eurycope madseni Wolff, 1956 Eurycope menziesi Wolff, n. sp. Haplomesus gigantens Birstein, n. sp. Hydroniscus n. sp. Ilyaraehna kermadecensis Wolff, n. sp. llyarachna sp. Ischnomesus bruuni Wolff 1956 Ischnomeses spircki Wolff, 1956 Janirella macrura Birstein, n. sp. Leptanthura hendili Wolff, 1956 Macrmtylis galatheae Wolff, 1956 Maerostylis hadalis Wolff, 1956 Storthyngura benti Wolff, 1956 Storthyngura bicornis Birstein, 1957

139 140 141 142 143 144 145 146 147 148 149

~5-11 i I 3 2 3 1 4 1 1 J 1

c. 21 i I-~ I

!1

{ 15

2

1

11 13 7

('

~

3 2 28 150 3 13 7

1

2

I

7

---

1 1

2

No. o f specimens in trenches

Pto. Rico Banda Kermadec Philippine Kermadec Tonga

Kermadcc

Kur.-Kamt. Banda Philippine Banda Kermadec Japan Kur.-Kamt. Japan Kermadec Aleutian Kur.-Kamt. Japan Kermadec Kur.-Kamt. Aleutian Kur.-Kamt. Idzu-Bonin Kur.-Kamt. Tonga Kermadec Pto. Rico (?) Banda Kermadec Pro. Rico Banda Kermadec Philippine

Kur.-Kamt. Japan Kermadec Kur.-Kamt. Kermadec

Kermadec

Trench(es)

6960-7000

Depth (m) (in trench)

l--contd.

8330~8430 6475-6570 6660-7000 8330-8430 6960-7000 6660-7000 7210-8430 6580 9820-10,000 7270 5230-7000 6156-6207 686O 6475-6570 5850-6770 7246 7210-8100 6475-7280 6620-6730 7210-7230 7246 8330-8430 7305-7315 7210-8660 10,687-10,415 6960-7000 7625-7900 7290-7250 6960-7000 7625-7900 6580-7270 7640 10,020-10,190 6660-6770 7625-7900 6490-6650 8210-8300 9820-10,000 6180 10,687-10415

APPENDIX

Recorded by

Exhibit., 1958

"

BIRSrEIN, 1957

ZENK. et al., 1955 Exhibit., 1958 DAHL, 1959 SCHELLeNnFRO, 1955 DAHL, 1959

::

BELJ

V e t al., 1958

SCHELLENBERG, 1955 DAHL, 1959

SCtmLLENaERG, 1955 r DXHX, 1959

)

BIRSTEIN, 1957

BIRSTEIN, 1957

.. WOLFF, 1956

:)

/ I

Exhibit., 1958 WOLFF, 1956

Exhibit., 1958 I WOLFF, 1956

I

i; WOLFF, 1956

!

[

]

Vit.

Gal.

Ai~

Gal.

Gal. Alb. Gal.

,P

Gal. Vit.

Vit.

Gal.

Vit.

Vit. Gal.

Gal. Vit. Gal.

Vit.

Gal.

Research vessel

:

1318 1158 1414-4330

Also hadopelagic

2

÷

+ ÷ + + + +

1 1 1

+ + +

(1)

2

2 2 9

1

6

I

2

1 1

2

2

(4) (1)

5

+

3

1

5

I 1 1

2 2

1

(1)

2

(1)

I 1 1 1

Total no. o f localities

3

1165-1264

5345

I

L

i

i

m

L

Depth range outside trenches

+

÷ + + + ÷ + ÷ +

+ + + + + ÷

In one trench only

O

O ¢¢

tO

Kermadec

SOLENOGASTRES Neomenia (?) sp.

186

'*One egg capsule included.

207

GASTROPODA P R O S O B R A N C H I A I (6-10 spp.) I I (1 s p . ) I H (1-2 spp.) ,, I V (1 sp.) Admete sp. Buccin/dae ben. (1-3 spp.) Natacidae ben. (1-3 spp.) Odostomta sp. I Odostomla (?) sp. I I Trochiidae gen. I (1 sp.) Trochiidae gen. I I (1 sp.) L A M E L L I B R A N C H I A T A (3-5 spp.) Axinulus sp. I Axinulus sp. I I Bathynella flora (Dall, 1916) Cuspidarla sp. Glomus sp. ttyalopecten sp. KellieL'a sp. I (cf. pacifica E. A. Smith, 1885) Kelliella sp. I I Ledella ultraabyssalis Filatova, n. sp.

New Britain

MOLLUSCA LORICATA

185

187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206

Banda Kur.-Kamt.

PYCNOGONIDA Nymphonfemorale Fabe, 1956 Nymphon profundum Hilton, 1942

183 184

Kermadec

13 11 many

21 25

~lO

1

1

1

Kur.-Kamt. Tonga Kermadec Japan Banda Philippine Kur.-Kamt. Kermadec Sunda Kur.-Kamt.

6660~6770 8210-8300 6620-6730 5850-6770 8330-9050 10,687-10,415 9995-10,002 6156--6207 6580-7290 9820-10,190 8100 6620-8300 7000-6900 900O-9050

Kur.-Kamt.

{ 1o

1

7210-9050 8980-9043 10,687-10,415 9995-10,002 6660-6770 6860

Kur.-Kamt. New Britain Tonga Kerrnadec

152 several 7* 3

5660-6770

7657

~90-6650

7210-7230

Kur.-Kamt.

MYSIDACEA Amplyol~ m a g n a Birstein and Tchindonova, 1958

182

10,687-10,415 6620-8300 7160 6960-7000 6660~6770

Tonga Kermadec Sunda Kermadec

Depth (m) (in trench)

1--contd.

Pontobeneidae (?) (1 sp.) Prineaxelia uby~alis Dahl, 1959 Ritehotropis nemmin~i Dahl, 1959 Rachotropis (?) sp. Schisturellu galatheae Duhl, 1959 Tryphosa bruuni D a h l ,1959

French(es)

APPENDIX

176 177 178 179 180 181

No. of i specimens in trenches [

b

Exhibit., 1958

BRUUN, 1957 BRUUN, 1953 Z~NK. et al., 1955 BrtUUN, 1957

ZENK. et al., 1955 Exhibit., 1958

BaUUN, 1953

ZENK et al., 1955

BELJAEV et aL, 1958 Exhibit., 1958

Exhibit., 1958

Exhibit., 1958

FAGE, 1956 ZENK. et al., 1955

BIRSTEIN and TCHINDONOVA, 1958

BELJAEV etal., 1958 DAHL, 1959

Recorded by

]

i

i

i

i

i

,

Vit.

Vit. Gal.

Gal.

Vit.

Gal.

Gal. Vit.

Vit.

Gal.

Gal. Vit.

Vit.

Gal.

Vit.

Research vessel

+

+

+

+

In one trench only

i

i

L

m

m

2870

i

m

g)40-4330 3231

m

m

m

Depth range outside trenches

(2) (2) (l) (2) (1) (!)

2

(3) (1) (i) (l) (l) (1) (1) (1) (1) (1) (2) (3) (l) (1)

(1)

(l)

1

1

(l)

1

3

II)

Total no. ~f localities

O ¢:h r~

5"

5.

8

o ~r

2 2

ECHINODERMA C R I N O I D E A I (1 sp.) _ II (1 sp.) Bathycrinus australis (A. H. Clark, 1907) Bathycrinus sp. A S T E R O I D E A (1 sp.) Brisingidae gen. (2-3 spp.)

Eremieaster vicinus (Ludwig, 1907)

Freyella mortenseni Madsen, 1956 Hymenaster blegvadi Madsen, 1956 Hyphalaster sp.

229

230 231 232 233 234 235

236

237 238 239

3 54 3 38

(,

4

Kermadec

BRYOZOA Bugula sp.

228

N~0V Pac.

Idzu-Bonin Kur.-Kamt. Kermadec Kur.-Kemt. Mariana Kur.-Kamt. Sunda Kermadec Kttr.-Kamt. Kermadcc

Kur.-Kamt.

CEPHALOPODA Octopodidae gen. (1 sp.)

227

m

Sunda

SCAPHOPODA Siphonodentalium sp.

Kur.-Kamt. Riu-Kiu Japan N W Pac. Japan N W Pac. Banda Kermadec

Kermadec

Kermadec Aleutian I ~ Kur'-Kamt',,

Banda Kermadec

220

4 c. 50 125

1

{ 56

3

4 4 3

{ 390

m

221 222 223 224 225 226

Spinula (Bathysp.) oceanlca Filatova, 1958

219

i

;

Trench(es)

Spinula (Bathysp.) vitlazi Filatova, n. sp. Spinula (Spinula) calcar (Dall, 1908) Teredo sp. Thyastra sp. Xylohaga sp. I Xylophaga sp. II Yoldia sp.

Ledidae gen. (I sp.) Lucinidae gen. (1 sp.) Malletia sp. Nellonella subovata Verrill & Bush, 1897 Netlonella sp. I Nellonella sp. II Nucula sp. Nuculidae gen. (1 sp.) Peetinidae gen. (1 sp.) Propeamussium sp. Spinula (Bathyspinula) bogorovi Filatova, 1958

208 209 210 211 212 213 214 215 216 217 218

No. o f specimens in trenches

APPENDIX

9715-9735 6860 8210-8300 9000-9050 7585-7615 686O 7000--6900 6620-6730 6860 5850-6180 6660-6770 6272-6282

8210-8300

8100

7000-6900

6660~6770 6960-7000 6620-7000 6860-7230 6810 6156-6207 6272-6282 7587 6096 7290-7250 9995-10,002 6660-6770 7290-7250 8210-8300

6860-8430 721 0-7230 6960-8300 7246 7210-7230

Depth (m) (in trench)

1--contd.

Exhibit., 1958

ZENK. et al., 1955 MADSEN, 1956

Exhibit., 1958 ZENK. et aL, 1955 GISLC'N, 1956 ZENK. et al., 1955 Exhibit., 1958 ZENK. et aL, 1955 MADSEN in litt.

BRUUN, 1953

ZENK. et al., 1955

BRUUN, 1953

Exhibit" 1958 FILATOVA, 1958 BRUUN, 1957 Exhibit., 1958

BRUUN, 1957 ZENK. et aL, 1955 FILATOVA, 1958

Exhibit., 1958 ZENK. et aL, 1955

ZENK. et al., 1955 s,

Recorded by

Vii. Gal. ,, Vit.

Total no.

(1)

1

i

2

6

(1) (1) 5 (1) (1) (1)

(1)

(1)

(I)

6 (1) (1) (1) (1) (1)

16

(2) (I) (6) (1) (1) (1) (1) (1) (2) (2) (I)

of localities

!

5200-5800

2510-4640

4160-5790

4640-5876

3830-4540

L

Depth range outside trenches

, --

In one tr. only

o11. }

Gal. Vii.

Vit.

Gal.

Vit.

Gal. Vit. Gal.

Vit.

s,

,0 Gal.

Gal. Vii.

Vit.

Research vessel

O

Z

O ~e

Elpidia sp. Euphronides verrucosa Ludwig, 1894 Gephyrothuriidae gen. I (1 sp.) Gephyrothuriidae gen. I I (1 sp.) Hadalothuria wolffi Hansen, 1856 Mesothuria murrayl (Th6el, 1886) Myriotrochus bruuni Hansen, 1956

Myriotrochus sl~. P a r o r i z a grevei Hansen, 1956 Peniagone vedeli Hansen, 1956 Peniagone (? Scotoanassa) sp. Periamma naresi (Th~el, 1882) Pseudostlehopus villosus The,el, 1886 Pseudostichopus sp. Scotoplanes galatheae Hansen, 1956 Scotoplanes globosa Thin.el, 1879

260 261 262 263 264 265 266

267

270 271 272 273 274 275

268 269

Elpidia glacialis Th~el, 1876

ECHINOIDEA Portalesia sp. (?) ourorae Koehler, 1926 H O L O T H U R I O I D E A I (5-10 spp.) II (1 sp.) Apoda (I sp.) Benthodytes sanguinolenta Th~el, 1882 Ceraplectana trachyderma H. L. Clark, 1908 EIpidia glacialis kermadecensis Hansen, 1956 EIpidia glacialis solomonensis Hansen, 1956 EIpidia glacialis sundensis Hansen, 1956

Porcellanaster caeruleus Wy¥. Thomson 1877 Pterastidae gen. (2-3 spp.) Styracaster n. sp. Vitiazaster diakonovi Beljaev, n. sp. O P H I U R O I D E A I (2-3 spp.) I I (1 sp.) ,, I I I (1 sp.) Amphiophiura bullata (Wyv. Thomson, 1877) Ophiacantha opercularis (Koehler, 1909) Ophiura Ioveni (Lyman, 1878)

259

256 257 258

250 251 252 253 254 255

240 241 242 243 244 245 246 247 248 249

Kermadee New Britain Sunda Aleutian Kur.-Kamt. Japan Idzu-Bonin New Britain Kur.-Kamt. Banda New Britain Mariana New Britain Banda Philippine New Britain Tonga Banda Kermadee Pto. Rico Sunda Kermadee Kur.-Kamt. Philippine Kermadec

1

49

1

5 18 c. 1030 20 114 35 3

10

14

1

4 14

7290-7250 6860-9950 9995-10,002 10,630-10,710 6490-7290 6490-6650 6620-8300 8780-8940 6900-7160 6410 8100-9950 6156-7587 9715-9735 8980-9043 8610-8660 6490-7290 8980-9043 10,630-10,710 8780-8940 6490-6650 10,120-10,210 8940 10,687-10,415 6490-7290 6180-8300 7625-7900 7160 6660--7000 8100 9820-10,000 6180-6770

Kern~'adec

Banda Kur.-Kamt. Kermadee Mariana Banda

6660~6770

New Britain Kermadec E. Atlantic

Depth (m) (in trench)

l--contd.

6035 6860-7230 6272-6282 6860 6860-7230 8006 (?) 6620-6730 6035

E. Atlantic Kur.-Kamt. N W Pac. Kur.-Kamt.

Trench(es)

i: l:O

. c. 1787 O0

6

8

2

1

m

1 3 1 177

38

1

No. o f specimens in trenches

APPENDIX

MADS~N, HANSEN, ., ZENK. et HANSEN,

Gal. Vit. Gal. aL, 1955 1956

Vit. Gal.

~t

Gal. Vit.

Vit.

Gal.

Vit.

Gal.

1953 1956

HANSEl, 1956

HANSEN, 1956

ZENK. et al., 1955 HANSEN, 1956 Exhibit., 1958

Exhibit., 1958 ZENK. et al., 1955 Exhibit., 1958

HANSEN, 1956

ZENK. et aL, 1955 Exhibit., 1958

M.~s~N, 1956

Gal. Pr.-A.

Pr.-A. Vit.

KOEI-ILER, 1909 ZENK. et aL, 1755 Exhibit., 1958 Z~NK'{ et aL, 1955 Exhibit., 1958 MADSE~, 1956 KOVJ~LER, 1909

Research ~e$$e[

Recorded by

+ --

--

+ +

+

--

--

~

-+ 4+

--

-+ --

t

545-4809

3292 896-5307

254-2516

2404-4478

70-4750

768-5206 3188-5173

440-1690

2515-4755

2270-5610

1435-5600

--

+

Depth range outside trenches

In one trench only

11

1

2 9 (1)

(I)

3 5

(!)

4

2 5

(1) (I)

7

(1)

c. 20

(1) (1) 29 4 2 2 2

(4)

4

4

1

(2) (1) (1) (2) (1) (i) (3)

23

Total no. o f localities

lb,

-q

Scotoplanes sp.

289 290

288

286 287

285

284

I

I

f

Careproctus n. sp. Liparldae gen. (1 sp.)

I

Careproctus 0Pseudoliparis) amblystomopsis Andr., 1955 ( k

PISCES Bassogigas profundissimus (Roule, 1913) Bassogigas sp.

Culeolus sp.

TUNICATA Cnemidocarpa bythia (Herdman, 1882)

POGONOPHORA 277 Heptabrachla abyssicola Ivanov, 1952 278 Heptabrachia subtilis Ivanov, 1957 279 Lamellinbella Johanssoni Ivanov, 1957 Siboglinum caulleryl Ivanov, 1957 280 281 Spirobrachia I~klemiscl~vi Ivanov, 1957 Zenkevltchlana Iongissima Ivanov, 1957 282 283 ENTEROPNEUSTA (1 sp.)

276

$

1

1 I

1

7

12

many

2 2 9

No. of specimens in trenches

Kerrnadec

E. Atlantic Sunda Kur.-Kamt. Japan

Kermadec Kur.-Kamt. Japan NW Pac.

Idzu-Bonin Japan Kur.-Kamt.

Kur.-Kamt.

Trench(es)

6035 7160 7210-7230 6156-7587 7587 6660-6770

6180-7000 7210-7230 6150-6207 6096-6282

8100 9600 (?) 6156-6207 8100 9000-9050 8330-9950 8100

6860

Depth ( m ) (in trench)

A P P E N D I X 1--contd.

ROUTE, 1913 BRUUN, 1953 ANDRL~HEV, 1955 RASS, 1958a, 1958b RAss, 1958b Bauu~, 1953

MILLAR, 1959 ZENK. et aL, 1955 Exhibit., 1958

ZENK. et al., 1955

IVANOV, 1952 IVANOV, 1957

ZENK. et al., 1955

Recorded by

d~J.

Vit.

Gal.

Nit.

Gal.

Vit.

Research vessel

m

+

+

4 + +

In one trench only

?

5610-5860

4850-

23-3400

Depth range outside trenches

(I) (1)

3

3 (1)

(4)

(1)

¢.5

1

c. 20

1 I 1

(1)

Total no. o f localities

o

r-

7,

~e

APPENDIX

2

DECAPODA (CARIDEA) (GALATHEIDEA) PYCNOGONIDA

MYSIDACEA

AMPHIPODA

ISOPODA

CUMACEA TANAIDACEA

PRIAPULOIDEA BRYOZOA COPEPODA (HARPACT.) OSTRACODA CIRRIPEDIA

ECHIUROIDEA SIPUNCULOIDEA

OCTOCORALLIA NEMERTINI NEMATODES POLYCHAETA

HEXACORALLIA

SCYPHOZOA

PORIFERA HYDROZOA

FORAMINIFERA

Group

Nymphon profundum Hilton

Amplyops magna Birstein & Tchindonova

Hirondellea gigas Birstein & Vinogradova

Macrostylis galatheae Wolff

Neotanias serratispinosus hadalis Wolff

Scalpellum japonicum Hoek

Phascolion pacificum Murina

Macellicephala hadalis Kirkegaaxd

Galatheanthemum hadale Carlgren

Stephanoscyphus simplex Kirkpatrick

Hallsiphonia galatheae Kramp

Rheophax nodulosus Brady Thurammina compressa Brady

Deepest identified species

Year 1957 1953 1953 1953 1952 1952 1958 1952 1958 1951 1953 1953 1957 1958 1951 1951 1953 1953 1957 1952 1958 1957 1952 1953 1957 1958 1952 1957 1951 1957 1951 1953 1953 1951 1951 1953 1953

Research vessel Vit. Vit. Vit. Vit. Gal. Gal. Vit. Gal. Vit. Gal. Vit. Vit. Vit. Vit. Gal. Gal. Vit. Vit. Vit. Gal. Vit. Vit. Gal. Vit. Vit. Vit. Gal. Vit. Gal. Vit. Gal. Vit. Vit. Gal. Gal. Vit. Vit.

Locality Tonga Kur.-Kamt. Kur.-Kamt. Kur.-Kamt. Kermadec Kermadec Kermadec Kermadec Mariana Philippine Kur.-Kamt. Kur.-Kamt. Tonga Mariana Philippine Philippine Kur.-Karnt. Kur.-Kamt. Japan Kermadec Kermadec New Britain Kermadec Kur.-Kamt. New Britain Kermadec Kermadec Tonga Philippine Tonga • Philippine Kur.-Kamt. Kur.-Kamt. Celebes Sea Celebes Sea Kur.-Kamt. Kur.-Kamt.

Depth (m) 10,687-10,415 9000-9050 9000--9050 8610-8660 8210-8300 8210-8300 9995-10,002 6960-7000 10,630-10,710 10,150-10,210 8610-8660 7210-7230 10,687-10,415 10,630-10,710 10,150-10,210 10,150-10,210 6860 6860 7587 5850 9995-10,002 7657 (?) 6960-7000 686O 8006 8928-9174 8210-8300 10,687-10,415 9820-10,000 10,687-10,415 10,190 7210-7230 7210-7230 5240-5160 5240-5160 686O 6860

Maximal recorded depth o f major groups o f marine animals and o f deepest identified species (beyond 6000 m ) in each group

ZENK. et al., 1955 ZENK. et aL, 1955

Exhibit., 1958 Exhibit., 1958 BRUUN, 1953 TACt. and ZEV., 1957 Exhibit., 1958 Exhibit., 1958 WOLFF, 1956 Exhibit., 1958 WOLVF, 1956 Exhibit., 1958 D^HL, 1959 BmST. and TCH]ND., 1958 BmST. and TCHIND., 1958

Exhibit., 1958 ST$CIIEDRINA, 1958 STSC~ir~DmNA, 1958 ZV.NK. et al., 1955 KRAMP, 1956 K~MP, 1956 Exhibit., 1958 KRAMP, 1958 Exhibit., 1958 C . ~ t t o t ~ , 1956 ZE~'~K. et al., 1955 7_~NK. et aL, 1955 Exhibit., 1958 Exhibit., 1958 KIRKEGAARD, 1956 BRUUN, 1953 MURINA, 1957 MU~NA, 1957 Exhibit., 1958

Recorded by

¢2L

8

¢3,

PISCES

ENTEROPNEUSTA TUN1CATA

POGONOPHORA

HOLOTHURIOIDEA

ECHINOIDEA

OPHIUROIDEA

ASTEROIDEA

SCAPHOPODA CEPHALOPODA (OCTOP.) CRINOIDEA

LORICATA SOLENOGASTRES GASTROPODA LAMELLIBRANCHIATA

Group

m

Careproctus amblystomopsis Andrlashev

Cnemidocarpa bythis (H©rdman)

Zenkevitchlana longlssima Ivanov

Myriotrochus bruunl Hansen

Pourtalesia sp. 7 aurorae Koehler

Ophiura loveni OLyman)

Eremicaster vlctnus (Ludwig)

Bathycrinus australis (A. H, Clark)

Spinula (Bathysp.) bogorovi Filatova Siphonodentalium sp.

Neomenia sp.

Deepest identified species

2---contd.

7657 6660-677O 10,687-10,415 10,687-10,415 6810 7O00-690O 8100 9715--9735 8210-8300 7585-7615 70OO-6900 8OO6 6660-6770 7290-7250 7290-7250 10,630-1~710 10,120-10,210 9700-9950 9700-9950 8100 7210-7230 6960-7000 7587 7587

Depth (m)

APPENDIX

New Britain Kermadec Tonga Tonga Riu-Kiu Sunda Kur.-Kamt. Idzu-Bonin Kermadec Mariana Sunda New Britain Kermadec Banda Banda Mariana Philippine Kur.-Kamt. Kur.-Kamt. Kur.-Kamt. Kur.-Kamt. Kermadec Japan Japan

Locality

Year 1957 1952 1957 1957 1955 1951 1949 1955 1952 1955 1951 1957 1952 1951 1957 1958 1951 1953 1953 1949 1953 1952 1957 1957

Research vessel Vit. Gal. Vit. Vit. Vit. Gal. Vit. Vit. Gal. Vit. Gal. Vit. Gal. Gal. Gal. Vit. Gal. Vit. Vit. Vit. Vit. Gal. Vit. Vit.

Exhibit., 1958 Exhibit., 1958 Fn.ATOVA, 1958 BRUUN, 1953 7-~NK. et aL, 1955 Exhibit., 1958 Gisl~n, 1956 Exhibit., 1958 MADSEN (in lltt.) Exhibit., 1958 MAX)SEN, 1956 MADSEN, 1956 MADSEN, 1956 Exhibit., 1958 HANSEN, 1956 IVANOV, 1957 IVANOV, 1957 ZHNK. et al., 1955 ZENK. et al., 1955 MnA.AR, 1958 Exhibit., 1958 Exhibit., 1958

Exhibit., 1958

Recorded by

O

Z

O

oo

T h e h a d a l c o m m u n i t y , a n introduction

APPENDIX

119

3

List of trawlings and bottom samplings at depths exceeding 6000 m with information about the following items, according to all records available : Depth, bottom temperature (in °Celcius), position, research vessel, station number, date, gear (see abbreviations below), number of species, approximate number of specimens and details about the fauna (owing to considerations of space the latter is indicated only by means of ciphers which refer to the ones given at the extreme left in Appendix 1 with the total list of the trench animals ; ciphers in italics refer to identified species). After the name of each trench is given its greatest depth recorded*). Abbreviations : H O T : H e r r i n g O t t e r T r a w l ; ST : Sledge (or Agassiz or Sigsbee) Trawl ; D : D r e d g e ; P G : Petersen G r a b ( b o t t o m s a m p l e r 0.2 or 0.25 mS). PACIFIC OCEAN

ALEUTIAN TRENCH (7679m) Vitjaz

Stn. 3340

1955ST

52" 2 5 ' 8 ' N , 170 ° 53'9'E

Vitjaz

Stn. 3357

1955 ST q- P G

Echiuroid. : 114-115-117 l s o p o d a : 156-159

Lamellibr. : 211

6410m

H o l o t h u r . : 259 1.62°C

7246m

Actiniaria : 50 Polychaeta : 82

KURILE-KAMTCHATKA

TRENCH

(10,382 m)

Vitjaz Stn. 3168

c. 6000 m

Vitjaz Stn. 2144

48 ° 2 5 ' 2 ' N , 156 ° 34"2 'E

6860 m

1955 P G

(1-2.6.1953) ST No. o f species : 4 5 + 8 F o r a m i n . : 3-6-8-12-17-

19-21-27 Porifera • 32-34 H y d r o z o a : 38 S c y p h o z o a : 40 A n t h o z o a : 44-56 e. 7 0 0 0 m

7 2 1 0 - 7 2 3 0 m 1.78°C N o . o f species : 4 2 q- 13 F o r a m i n . : 3-5--8-10-1315-16-17-19-

20-21-22-27 H y d r o z o a : 35 A n t h o z o a : 51-56 N e m e r t i n i : 61 Polychaeta : 64-69-84--89-9092-104-110

8100 m No. o f species : 17 + 11 Foramin. : 4-7-8-9-11-21-

23-24-28-29 Porifera : 32 Actiniaria : 44 Polychaeta : 72-81-85-

88-102-111 *After

ZENKEVICH, 1958b.

N o . o f s p e c i m e n s : e. 4 2 5 Polychaeta : 64-110 Echiuroid. : 115 Sipuncul. : 119-120-121 Cirripedia : 126 I s o p o d a : 154 P y c n o g o n : 184

G a s t r o p o d a : 192-193 Lamellibr. : 208-217 Crinoidea : 231 Asteroidea : 235-236-241 Ophiuroid. : 244 H o l o t h u r . : 251-276

Vitjaz

Stn. 3257

1955 P G

49 ° 2 9 ' Y N , 158 ° 41"0'E Vitjaz No. of specimens : e. 690 Echiuroid. : 118 Priapul. : 123 I s o p o d a : 137-148-156-158 A m p h i p o d a : 161 M y s i d a c e a : 182 G a s t r o p o d a : 187

Stn. 2208

22.6.1953 ST

Lamellibr. : 209-212-217 Asteroidea : 241 Ophiuroid. : 244 H o l o t h u r . : 251 T u n i c a t a : 285 Pisces : 288

43 ° 39'N, 152 ° 0 Y E Vitjaz Stn. 162 10.10.1949 ST N o . o f s p e c i m e n s : c. 2 0 0 Echiuroid. : 116 Cephalop. : 228 I s o p o d a : 156 H o l o t h u r . : 259-273 A m p h i p o d a : 161 P o g o n o p h . : 277-280 Lamellibr. : 204 EnteFopn. : 283

120

TORBEN WOLFF

8330-8430 m

N o . o f species : 2 0 Polychaeta : 69-90-110 E c h i u r o i d . : 115 l s o p o d a : 142-145-148-160 8610-8660 m

1.99°C

N o . o f species : 9 Porifera : 32 H e x a c o r r . : 43 O c t o c o r r . : 59 8632-8779 m

2.08°C

N o . o f species : 18 + 5 F o r a m i n . : 14.-18-21-25-26 Actiniaria : 44 Polychaeta : 64--88 9700-9950 m

45 ° 4 1 . 4 ' N , 153 ° 23.8'E Vitjaz Stn. 2216 26.6.1953 ST N o . o f s p e c i m e n s : e. 2 9 0 P o l y c h a e t a : 64 Lamellibr. : 198 E c h i u r o i d . : 118 H o l o t h u r . " 259-260 A m p h i p o d a : 161

Vitjaz Stn. 3176

44 ° 0 8 . 5 ' N , 150°22'E E c h i u r o i d . " 118

A n t h o z o a : 58 9000-9050 m

46 ° 3 1 ' N , 1540 2 2 ' E Vitjaz Stn. 2120 26.5.1953 ST N o . o f s p e c i m e n s : e. 144 A m p h i p o d a : 161 H o l o t h u r . : 251-259 G a s t r o p o d a : 187 P o g o n o p h . : 282 Lamellibr. : 198-208

2.1°C

N o . o f species : 4 P o l y c h a e t a : 88

6.10.1955 H o l o t h u r . : 259

ST

44 ° 0 7 ' 8 ' N , 150°32.0'E Vitjaz Stn. 2217 29.6.1953 N o . o f s p e c i m e n s : e. 3 2 0 0 E c h i u r o i d . : 118 C r i n o i d e a : 233 G a s t r o p o d a : 187 H o l o t h u r . • 251-259 Lamellibr. : 198-207 P o g o n o p h . : 281-282

ST

43 ° 4 7 ' 6 ' N , 149 ° 54'8'E Vitjaz Stn. 2218 1.7.1953 N o . o f s p e c i m e n s : e. 2 5 0 E c h i u r o i d . : 118 H o l o t h u r . : 251-259

ST

JAPAN 6156-6207 m

Actiniaria : 52 E c h i u r o i d . : 116 Sipuncui. : 120-121 Cirrilmdia : 126 6475-6570 m

Actiniaxia : 52 7190-7280 m

I s o p o d a : 156 7587 m

Echiuroid. : 115 Priapul. : 123

TRENCH

(8597 m )

38 ° 10-9'N, 143 ° 5 6 ' 2 ' E l s o p o d a : 153 Lamellibr. : 201-219 H o l o t h u r . • 259

Vitjaz Stn. 3214 25.10.1955 P o g o n o p h . : 279

41° 2 2 ' 9 ' N , 145 ° 15"4'E E c h i u r o i d . : 116

Vitjaz Stn. 3457

38 ° 0 2 ' N , 143 ° 5 7 ' 4 ' E H o l o t h u r . : 259

Vitjaz

33 n 02"6'N, 144 ° 58-7'E Lamellibr. : 220 H o l o t h u r . : 259

Vitjaz

ST

T u n i c a t a : 285 P i s c e s : 288

21.9.1955 ST I s o p o d a : 143-153-156 Stn. 3227

2.5.1955

ST

Stn. 3571 1957 Pisces : 288-289

ST

N. IV. PACIFIC 6096 m

C i r r i p e d i a : 126 6272-6282 m

Sipuncul. : 120 Lamellibr. : 219

33 ° 18'N, 149°46'E Lamellibr. : 221

Vitjaz Stn. 3232

48 ° 15'N, 169°39'E A s t e r o i d e a : 239-242

Vitjaz Stn. 3363

I D Z U . BO N1N 7305-7315 m

TRENCH

30 ° 3 4 ' N , 142 ° 4 1 ' E

6.5.1955

ST

10.6.1955 T u n i c a t a : 285

ST

(9764 m )

Vitjaz Stn. 3491

7.10.1955

ST

Stn. 3514

1955

ST

Vitjaz Stn. 3494

1955

ST

I s o p o d a : 160 9 6 0 0 m ( 8 5 4 O m ?) H o l o t h u r . : 259

P o g o n o p h . : 278

9715-9735 m E c h i u r o i d . : 118

C r i n o i d e a : 230

Vitjaz

H o l o t h u r . : 259

121

The hadal community, an introduction

R1U - K I U 6810 m (?) 1.84°C Lamellibr. : 218

TRENCH

(7507 m)

Vitjaz

27 ° 30'N, 130 ° 35'E

MARIANA

TRENCH

ST

Vitjaz

Stn. 3689

1955

ST

Vitjaz

Stn. 4004

1958

ST

(10990 m)

7585-7615 m Asteroidea : 234 10630-10710 m No. o f species : 4 (? 6) Actiniaria : 53

Stn. 3528 28.10.1955

No. o f specimens : 12 Polychaeta : 86

Holothur. : 253-263

PHILIPPINE T R E N C H (10497 m) 9820-10,000 m 2.6°C No. o f species : 5 Actiniaria : 48 I s o p o d a : 150

10 ° 20'N, 126 ° 41'E No. o f specimens : 8 A m p h i p o d a : 174 Lamellibr. : 203

Galathea

1 0 , 0 2 0 - 1 0 , 1 2 0 m 2.6°C No. of species : 1 A m p h i p o d a : 169

9 ° 49'N, 126 ° 44'E No. o f specimens : 3

Galathea

Stn. 429

10120 m 2-6°C No. of species : 1 Holothur. : 266

10°28'N, 126°39'E No. of specimens : 1

Galathea

Stn. 424 27.7.1951

10190 m 2.6~'C No. o f species : 5 Actiniaria : 48 Polychaeta : 83

10" 13'N, 126 '~ 43'E No. o f specimens : 133 A m p h i p o d a : 169 Lamellibr. : 203

Galathea Stn. 418

10 ° 19'N, 126 ° 39'E No. o f specimens : 24 Echiuroid. : 112

Galathea Stn. 419

10150-10210 m No. of species : 4 Actiniaria : 48 Polychaeta : 83

2.6°C

BANDA

TRENCH

Stn. 435

7.8.1951

ST

Holothur. : 274

2.8.1951

21.7.1951

D

PG

ST

Holothur. : 266

22.7.1951 ST

H o l o t h u r . " 266

(7298 m)

6490-6650 m 3.5°C No. o f species : 10 Scyphozoa : 41 Polychaeta : 99-109

5 ~ 18'S, 131 ° 18'E Galathea Stn. 497 23.9.1951 H O T No. o f specimens : e. 70 A m p h i p o d a : 172 Holothur. : 254-255-261Pychnogon. : 183 265-268

6580m 3.5°C No. o f species : 8 Polyehaeta : 73-99-108-109 I s o p o d a : 149

5 ° 21'S, 131 ° 17'E No. of specimens : 12 A m p h i p o d a : 167 Lamellibr. : 202

Galathea

7270m 3.6°C No. o f species : 3 I s o p o d a : 151

5° 36'S, 131° 06'E No. o f specimens : 5 A m p h i p o d a : 167

Galathea

7240-7290 m 3.6°C No. o f species : 3 Polychaeta : 78-100

5 ° 36'S, 131 ° 01'E No. of specimens : 4 Holothur. : 268

Galathea

7290-7250 m 3.6°C No. o f species : 14 Actiniaria : 47 Polychaeta : 67-82-83-96-

5 ° 26'S, 130 ° 58'E Galathea Stn. 495 22.9.1951 H O T No. o f specimens : c. 100 A m p h i p o d a : 164 Echinoidea : 250 Lamellibr. : 222-225 Holothur. : 254-261-268

100-109

Stn. 499

24.9.1951

PG

Holothur. : 255

Stn. 496

23.9.1951

PG

Lamellibr. : 202 Stn. 494

22.9.1951

ST

122

TORBEN WOLFF

NEW

BRITAIN

TRENCH

6920 m 7657 m O s t r a c o d a : 125 C u m a c e a : 129 (8006 m ?)

(9140m)

Vitjaz

Stn. 3655

Vitjaz

Stn. 3663 1957 S T O p h i u r o i d : 245 ( 8 0 0 6 m ?)

L o r i c a t a : 185

1957

ST

8830-8780 m 2.7°C N o . o f species • 2 H o l o t h u r . : 257-264

5 ° 59'S, 153 ° 2 8 ' E No. of specimens : 76

Galathea

Stn. 521

16.10.1951

ST

8940 m 2.7°C N o . o f species : 4 P o l y c h a e t a : 66 (tube)

6 ° 31'S, 153 ° 5 8 ' E Galathea N o . o f s p e c i m e n s : 19 H o l o t h u r . : 257-264-266

Stn. 517

13.10.1951

ST

Stn. 3 6 6 3 - A 1957 H o l o t h u r . : 259-262

ST

Vitjaz

8980-9043 m Polychaeta : 66-68-76-95 l s o p o d a : 136

G a s t r o p o d a : 189

NEW

HEBRIDES

TRENCH

6680-6830 m

(7570m)

Vitjaz TONGA

TRENCH

1958

ST

(10840 m )

7345 m

Vitjaz

Stn. 3818

1957

ST

c. 9 7 0 0 m P o l y c h a e t a : 65

Vitjaz

Stn. 3823

25.12.1957

ST

Vitjaz

Stn. 3 8 2 3 - A 28.12.1957

ST

10687-10415 m N o . o f species : 17 q- 4 Foramin. : 2 Porifera : 33 N e m a t o d z s : 62 Polychaeta : 6 5 - 7 5 - 1 0 1 - 1 0 3

N o . o f s p e c i m e n s : c. 1 0 0 I s o p o d a : 134 A m p h i p o d a : 162-175-176 G a s t r o p o d a : 189

KERMADEC

TRENCH

Lamellibr. : 199 H o l o t h u r . : 267

(10030 m )

6180 m 1.2°C N o . o f species : 10 S c y p h o z o a : 41 Actiniaria : 48 O c t o c o r r . : 59

32 ° 09'S, 176 ° 3 5 ' W No. of specimens : 63 P o l y c h a e t a : 93-94 A m p h i p o d a : 174 A s t e r o i d e a : 237

6620-6730 m 1.3°C N o . o f species : 24 Porifera : 30 S c y p h o z o a : 41 Actiniaria : 49 O c t o c o r r . : 59 N e m a t o d e s : 63

32 ° 20'S, 176 ° 5 4 ' W Galathea Stn. 650 15.2.1952 ST N o . o f s p e c i m e n s : c. 4 2 5 P o l y c h a e t a : 79-82-91-93G a s t r o p o d a : 196 94-97-105 Lamellibr. : 205-216 Cirripedia : 127 A s t e r o i d e a : 236 l s o p o d a : 152-157 O p h i u r o i d . : 246 A m p h i p o d a : 177 H o l o t h u r . : 256-269-275

6 6 6 0 - 6 7 7 0 m 1.3°C N o . o f species : 3 2 Porifera : 30 H y d r o z o a : 37 Actiniaria : 45-49-54 P o l y c h a e t a : 71-82-83-91-

35 ° 51'S, 178 ° 3 1 ' W Galathea N o . o f s p e c i m e n s : e. 1190 Cirripedia : 127 T a n a i d a c e a : 131 I s o p o d a : 144-147-152-155 A m p h i p o d a • 170-181 G a s t r o p o d a : 191-194-197

94-97-106 Echi~roid. : 113

Galathea

Stn. 653

17.2.1952 H O T

H o l o t h u r . : 269-275 T u n i c a t a : 281

Stn. 658

20.2.1952

ST

Lamellibr. : 214-224 A s t e r o i d e a : 238 O p h i u r o i d . : 249 H o l o t h u r . : 269-272-275 Pisces : 290

The hadal community,

an introduction

123

6960-7000 m 1.3°C 32 ° 10'S, 177 ° 1 4 ' W Galathea N o . o f species : 31 N o . o f s p e c i m e n s : c. 2 3 5 Porifera : 30 C i r r i p e d i a : 127 S c y p h o z o a : 41 T a n a i d . : 132-133 Actiniaria : 49 I s o p o d a : 139-140-141-144P o l y c h a e t a : 80-82-83-94-97 146-147-152 E c h i u r o i d . : 113

Stn. 651 16.2.1952

HOT

A m p h i p o d a : 163-165-177-

179-180 Lamellibr. : 2 1 0 - 2 1 5 - 2 1 6 H o l o t h u r . : 269-272 T u n i c a t a : 284

Galathea

7640 m 1.4°C N o . o f species : 1 A m p h i p o d a : 168

35 ° 20'S, 178 ° 5 5 ' W No. of specimens : 1

20.2.1952

ST

8210-8300 m 1.5~C N o . o f species : 2 0 H y d r o z o a : 39 Actiniaria : 46-49 Polychaeta : 7 9 - 8 0 - 8 3 - 9 3 - 9 8 Echiuroid. : 113

35 ~ 16"S, 178 ° 4 0 ' W Galathea Stn. 649 14.2.1952 N o . o f s p e c i m e n s : c. 2 1 0 0 T a n a i d a c e a : 133 B r y o z o a : 229 A m p h i p o d a : 173-177 C r i n o i d e a : 232 G a s t r o p o d a • 195 H o l o t h u r . : 256-269 Lamellibr. : 2 0 5 - 2 1 0 - 2 2 6

ST

8928-9174 m T a n a i d a c e a : 130 9995-10002 m N o . o f species, : 11 + 5 Foramin. : 1 S c y p h o z o a : 42 N e m a t o d e s : 63

Stn. 656

Vitjaz

Stn. 3827

1958

ST

Vitjaz

Stn. 3831

1958

ST

N o . o f s p e c i m e n s : 177 P o l y c h a e t a : 74 C o p e p o d a : 124 l s o p o d a : 135

G a s t r o p o d a : 190 Lamellibr. : 200-223 H o l o t h u r . " 252

INDIAN OCEAN

SUNDA

TRENCH

(7317 m )

6730 m 1-4"C N o . o f species : 3 Porifera : 31

10 ~ 02'S, 107 ° 52'E No. of specimens : 3 Polychaeta : 77-80

Galathea

Stn. 462

7000-6900 m 1.4°C N o . o f species : 7 H y d r o z o a : 36 A n t h o z o a : 57 P o l y c h a e t a : 70

10 ~ 20'S, 109 ° 5 5 ' E No. of specimens : 28 Lamellibr. : 206 S c a p h o p o d a : 227

Galathea

Stn. 465

7160 m 1.4"C N o . o f species : 7 A c t i n i a r i a : 55 P o l y c h a e t a : 82

lff ~ 21'S, 110 '~ 12'E Galathea Stn. 466 6.9.1951 H O T N o . o f s p e c i m e n s : e. 3 1 7 0 C u m a c e a : 129 H o l o t h u r . : 258-271 A m p h i p o d a : 178 Pisces : 287 ATLANTIC

PUERTO 7625-7900 m

No. o f species : 7 P o l y c h a e t a : 87-107 l s o p o d a : 138

5.9.1951

PG

D

A s t e r o i d e a : 236 H o l o t h u r . : 258

OCEAN

TRENCH

(9218 m)

19 ° 4 9 ' N , 65 ° 01 ' W Albatross Stn. 370 17.8.1948 19 ° 4 2 ' N , 64 ° 4 4 ' W N o . o f s p e c i m e n s : c. 35 A m p h i p o d a : 163-166-171 H o l o t h u r . : 270

CAPE 6035 m N o . o f species : 4 Asteroidea" : 240

RICO

3.9.1951

SOT

VERDE B A S I N

12 ° 0 7 ' 3 0 " N , 33 ° 3 2 ' 4 5 " W No. of specimens : 6 O p h i u r o i d . : 247-248

Princesse-Alice 6.8.1901 Pisces : 286

ST

124

TORBEN WOLFF

APPENDIX 4 The hadopelagic fauna recorded from a haul with closing net from 6000-8500 m depth in the Kurile. Kamtchatka Trench. No. of specimens CRUSTACEA COPEPODA Aetidius sp. ? Calanus tonsus Brady, 1883 Heterorhabdus compactus (Sars, 1900) Lucicutia ushakovi Brodsky, 1955 Metridia okhotensis Brodsky, 1950 Metridia similis Brodsky, var. abyssalis Brodsky, 1955 Mimocalanus distinctocephalus Brodsky, 1950 Parascaphocalanus zenkevitchi Brodsky, 1955 Pareuchaeta sp. Pseudeuchaeta sp. Pseudochirella gurjanovae Brodsky, 1955 Scaphocalanus bogorovi Brodsky, 1955 Scaphocalanus sp. Scolecithrix birshteini f. major Brodsky, 1955 Scolecithrix birshteini f. minor Brodsky, 1955 Scolecithricidae gen. 1 Scolecithricidae gen. 1[ Spinocalanus similis Brodsky, var. profundalis Brodsky, 1955 Xanthocalanus pavlovskii Brodsky, 1955 Zenkevitchiella abyssalis Brodsky, 1955 OSTRACODA (4 spp.) AMPHIPODA Andaniexis subabyssi Bst. and Vinogr., 1955 Halite quarta Bst. and Vinogr., 1955 Hirondellea gigas Bst. and Vinogr., 1955 Hyperiopsis laticarpa Bst. and Vinogr., 1955 Vitjaziana gurjanovae Bst. and Vinogr., 1955

Recorded by

BRODSKY, 1955

37 1

97 2 1

13 1 1 1 1 1 1

64 2 1

BIRSTEIN et al., 1954 2 4 1

2

BIRSTEIN and VINOGRADOV, 1955 ,,

(DAHL,1959)