This Paper Not to Be Cited Without Prior Reference to the Author

This paper not to be cited without prior reference to the author

International Council for C.M. 1973/J:22 the Exploration of the Sea Pelagic Fish (Southern) Committee

Investigations on midwater fish in the Atlantic Ocean by G. Krefft ßundlöl !orscnun san . Institut für .Seefischerei für Fischerei Hamburg der Bi bliothek Bundesforschungsanstalt für Fischerei Hamburg

In recent years a number of oceanic transects were carried out by FRS "Walther Herwig" in order to study the geographical distribution and taxonomy of the midwater fauna in the Atlantic Ocean. Two transects were run from the Bay of Biscay to the mouth of the La Plata River via Madeira, Canary Islands, Cape Verde Islands and Trindade Island(~~y 1966, January/February 1968); each of them covered abt. 5 800 n.m •• A third one crossed the South Atlantic between Mar deI Plata and Cape Town touching Tristan da Cunha and Gough Island (March 1971), its length was abt. 3 600 n.m•• The last of these transects (April 1971) crossed the eastern Atlantic in a lengthwise direction; it was run from Cape Town via St. Helena, Ascension, Cape Verde and Canary Islands to Madeira and covered abt. 4 800 n.m •• Stations along these transects are shown in fig.1.

Benefiting from the facilities of a large stern trawler we used large commercial gear, a 1 600-meshes-herring trawl for sampling the midwater fauna instead of the more conventional IKMT's and • other small nets. Fishing depths were measured by using the "netsonde" down to 1 000 m, deep hauls beyond that depth were measured by using a time-depth recorder (Benthos Co.) attached to the trawl. Very few hauls were made in day-light, most of them after dusk when the fish had finished their vertical upward migrations. Step hauls at various depths choosen according to the deep scattering layers shown on the echo sounder were made in 1966 only. Lateron we decided to tow our gear in a single layer only, starting with the shallowest depth, and going deeper with the next tow(s). Two

-2- -2-

to threo hauls were executed as a rule, except whon shortnge or time did not allow more than one deep haul in abt. 2000 m. The towine time per haul ranged from 15 minutes in the shallower hauls to one hour in deep hauls at a speed or about 3 1/2 knots. ifuen fishing in 2 000 m, the process or hauling the gear had to bo intcrrupted twice, in 1 000 and 500 m depth for about 20 minu tos in order to cool the winch. The net then started fishing again, tho doep hauls thus.converting into step hauls. Because no opening'" closing device exists ror large trawls, the deeper hauls may have bcen contaminatcd to some extent with specimons caught while tho not waS being hauled back. Therefore, it is impossible to calculate actual numbers or rish/hour caught at aselected depth. The depth distribution or hauls, their numbers, nnd those or the specios and specimens obtained isshown on tables 1-4.

-. The total catches were as rollows: Year No.or No.or 1) Haul depths Nos.or hauls hours m (range) Families Species Specimens . 1966 25 34,5 50-650 66 350 ·31 161 1968 72 26 40-2000 09 426 26 373 1971 I 30 17,25 70-2200 60 265 23 017 1971 11 39 19,5 74-2100 85 424 30 017

1) E:x:cept .. timo consumed ror hauline back and stops in deop hauls.

Tho total number or specimens taken along the 4 transects was 110 568; About 500 species are represented, some.40 or which were new to science. or these 17 are published as yet, almost all or them in the "Archiv rür Fischereiwissenschart". Besides, quite a number • or taxonomical descriptions will be round in revisions or ramilies and genera to be published elsewhere. One of the major aims or our studies is the zoo gcoeraphical analysis of thc midwater rishes populating the mesopelagic and bathypelaeic realms of thc Atlantic Ocean. In spite of the encouragine work on the distribution or mesopelagic fishos bogun in rocent years by American scientists or the Woods Hole Oceanographic Institution as weIl as of the Smithsonian Institution, we still are far away from a satisfying knowledge about the patterns

-3- -3-

of dintribution of North Atlantic mosopolagic fishes, and still farther from a thorough knowledge of the South Atlantic fauna. Even less is know about the physical and ?ioloeical factors controlling these patterns, though a number of fruitful hypothesos have been brought forward. As a matter of fact, the analysis of our catches is still at the very beginning and will need years to be finished. Much of the material has been and will be distributed to specialists all over the world, and the help of many colleagues of various countries is highly acknowledged. Here, only a fe,{ examples of typical distribution patterns ob served may be given.

Major changes in the species composition along our transects were observod at faunal boundaries as f.i. built by convorgences,

, ' curront systems, etc.' They correspond p1.rtly to those shown by • BACKUS et al., 1970. With these authors we distinguish a number of distribution patterns, the most remarkable ones exemplified here by figs. 2-12. The midwater fishes obtained by the "Walther IIerwig" can be arranged by their, distribution patterns in a number of well defined groups. I. The northern-temperate group. Several distribution patterns seem to be involved, the two most important being as folIows: a) The temperate pattern. Fishes showing this pattern are boreal specios found almost oxclusively north of 35°N. Thcy disappcar ,e from our catches just in front of Madeira. Due to thc small numbcr of hauls falling into this area they are represented in our catches by few specimens only. Examples are Nansenia groenlandica, Stomias boa ferox, MYctophum punctatum, Benthosema glaciale, Lampanyctus crocodilus and Uotoscopelus kro\yeri. The Blue \-Ihiting also belongs to that group. None of these species has a bipolar distribution. b) The temperate-subtropical pattern. Species showing this pattern have a similar distribution as those of pattern a), but range o farther southwardfinding their southern limits at about 20 U. Ichthyococcus ovatus, ArgyroTlelecus olfersi, Rhadinesthes dec'imus,

-4- I \ -4- ,~-----

IIygophum bonoiti, Symbolophorus voranyi, Diaphus holti, D. rafincoguei and Ceratospelus maderenois belong here. Aeain, none of thoso specios was met within subtropical or temperate'

latitudoo of tho southern homispnere by USo

II."The subtropical group. Most of the great number of species forming this group have an antitropical distribution in the subtropical balts of both hemispheros. ·00 . Almost all of them were found between 35 and 20 N, and between about 25 0 and 35 0 S again. In the eastern Atlantic the southern subtropical belt is broader stretching northward to the area of o . St. Helena (or about 16 S) due to the coöling effect of the Benguela ·Current. The bipolar subtropical pattern is exemplified by Mar~rethia obtusirostra, Astronesthes leucopogon, Paralepis atlantica (repre oonted in the South by a doubtfully distinct subspecies), }~croparalepis affinis (fig. 5), M. brevis (entering the tropics by submergence froc • 0 the South to about 10 S), Hygophum hygo~i (fig. 5), Diaphus effulgens, n. metopoclampus (fig. 7), Lampadena chavesi (fig. 5) and Lampanyctus pusillus. Others ohow a simila.r distribution pattern, 'l.lich, however, is restricted to either the northern or the southern subtropico. Such species may or may not be replaced in corresponding latitudes of the other hemisphere by closely related forms. Idiacanthus fasciola (Horth) and 1. atlanticus (South) are cxamples of such an "intrageneric" bipolarity. The fe" specimens of 1.fasciola found in the Cape area obviously are waifs carried into the Atlantic by the Agulhas Current (fig. 6). One other pair here to be mentioned is Cubiceps gracilis and C. coeruleus. Taken only in the northern subtropics were • Astronosthes neopogon, Bathophilus metallicus, Lampadena urophaos atlantica and Poromitra capito as were I~onesthes microcephalus, Grammatostomias dentatus, }Wctophum phengodes and Scopelopsis multipunctata in the southorn ones.

111. The tropical Broup. Soveral patterns of distribution of tropical specis can be observed as exemplified by BACKUS et 0.1., 1970. On hands of our own material we distinguish the following patterns: a) The broadly tropical pattern. We use this term introduced by BACKUS et al., 1970: 191 for the large group of species the distribution

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of which covers subtropical aS tropical waters as weIl. Such species entered our nets in the first two transects as a rule at rirst at }mdeira. They then continucd to be taken along the transect southward to the Subtropical Convergcnce at about 35°S. The most common representants of the broadly tropical pattern are a number of myctophids, e.g. Diaphus mollis, Cerato scopelus warmingi and Notoscopelus resplendens. ~owever, even the two last-mentioned species (figs. 11, 12) differ somewhat in their distribution. Thus, whereas f. warmingi does not show any gaps in its range between }~deira and the South Atlantic Subtropical Convergence, the distribution area of !. resplendens was found to 0 be interrupted between 3 and 23°S in the western Atlantic and 0 0 for a ohorter distance (2 30 1 ' to 11 S) in the eastern part of the ocean, i.e. in the area of the South Equatorial Current. In the • western Atlantic a countcr-clockwise gyral is built up by this current off the coast of.Brazil. The area has great similarity to the Sargasso Sea in the North Atlantic. It is of very low productivity as marked by the sharp decline in numbers,of species and specimens in our catches at stations 189 to 190 (1966, tab. 1) and 20 to 21 (1968, tab. 2). Quite a number of common tropical and broadly tropical species io lacking here. Others, like some species of Eustomias, Chauliodus danae and Lepidophanes gaussi lacking in more productive areas and representing the "Sargasso Sea pattern" of BACKUS et ale reappear again. Thus, the low productivity of these waters most probably is responsible for the observed gap in the distribution of !. resplendens (and others) • b) The tropical patterns. Fishes exemplifying these patterns are • 0 confined to tropical areas, i.e. the region between about 20 N and 25 oS along our transects. However, the southern boundary is much less defined than the northern one. "Amphitropical", that is species occuring in both, eastern and western parts of the Atlantic were found as weIl as others restricted to the eastern ("Guinean" pattern) either or to the western ("Amazonian" pattern) sections of the ocean. Heterophotus ophistoma, Astronesthes indicus, Microstoma microstoma, Stomias affinis,Hygophuo macrochir (s.l.), 11Yctophum asperum, Lampadena luminosa, various Diaphus species and many others are amphitropical. As an example

-6- -6-

the distribution of Astronesthes richardsoni (fig. 2) is given, though it does not cover the whole tropical belt in thc .North - South direction. Eastern tropical speeies are i.a. Paraholtbyrnia cyanocephala and Polyipnus polli (both fig. 4), Stomias lampropeltis, Chauliodus schmidti, ~lereas Diaphus garmani, Brama earibbea and Pseudoseopelus scutntus (fig. 3) are showing the western pattern. In addition severnl other patterns seem to be involved exemplifying distribution areas as f.i. Pollichthys mauli already diseussed by BACKUS et ale (l.e.: 192, fig. 8) or rather loealized habitats, mainly in the eastern Atlantie, as Lampadena pontifex (KREFFT, 1910: 211, fig. 5) and a still undeseribed Symbolophorus speeies in the Cape Verde region show• • One pattern of tropieal distribution eventual~ to be mentioned may be called the Agulhas pattern. This is represented by a number of.speeies of the tropieal Indian Oeean, whieh are drifting into the Atlantie around the Cape with the warm Agulhas Current. Their range inside the Atlantie is elose~ eonfined to the body of Agulhas wnter as shown in figs. 6 (Idiacanthus fasciolc.) and 9 for sot:le other apecies. Obviously these speeies are not able to survive in the surrounding South Atlantic Central Water. This perhaps does not apply to Diaphus diadematus.

IV. The southern groups During the 1911 transeet between }mr deI Plata and Cape Town, running partly through the area of the Subtropieal Convergenee, • partly through the neighbouring waters of the Westwind Drift, FRS "i-laIther Herwig" met with a fish fauna .quite distinet from those alone the other transects. Here, a hiehly mixed fauna of heterogenous origin was found. Of 265 speeies taken in the area 12 were not represented to the North of the Convergenee. These are members of thc follwing distribution patterns: a) The Convergenee pa~rn. GIBBS, 1968 was the first author reporting on a special fauna distributed eireumglobally in the waters of thc southern Subtropical convergenee. Several species have been added to his list sinee. The following speeies taken by us are

-7- -7-

membcrs of that peculiar fauna: Astronesthes boulengeri (fig.8) and two still undescribed Astronesthes species, Eathophilus ater, Eustomias trewavasae, Opostocias micripnus (fig. 8), Diaphus ostenfeldi (~ig. 7) and Scopelarchoides kreffti. b) The Westwind Drift pattern. Fishes of this pattern may be found in a more or less circumglobal belt between the Subtropical and the Antarctic Convergence, i.e. in the Antnrctic Intermedinte Wnter. A. great number of species, espccially myctophids, live here. Diplophos rebninsi, Photichthys argenteus, Luciosudis normani, Scopelosaurus hamiltoni, Protomyctophum subparalleum, P. tenisoni, Electrona carlsbergi, ~. paucirnstra, ~. subaspera, Hygophum hanseni, Lampadena dea, L•. notialis, Hintonia candens, Gymnoscopelus bolini, Lampichthys procerus, ~ nordenskjoeldi, Icichthys austrnlis and Ceratias tentaculatus are typical examples.

c) The broadly Antarct1c pattern. Here belang species having a distri bution area stretching from the consts of Antarctica to the J Subtropical Convergence. Stomias gracilis, Eenthalbella elongata, ~. macropinna, Protomyctophum andersoni, ~. bolini, P. normani, Electrona antarctica, Gymnoscopelus braueri, G. nicholsi, Lampanyctus achirus and the macrourid fish Cynomacrurus piriei may be mentioned. d) The bipolar pattern. Represented by a number of often deep living species occuring as well in subarctic as in subantarctic waters, some of them penetrating even subtropical arens by submergence as e.g. Anotopterus pharao, Borostomias antarcticus and Trigonolampa miriceps exemplify this pattern. v. The bathypelagic group.

More than 80 species, that is about 16$~ of the total number, have been taken in hauls deepcr than 800 m only. As a rule such species occupy larger ranges than cesopelagic species do. Their distribution areas often stretch from Gubarctic areaa to the southern Subtropical Convereence or even beyond. Gonostona bathyphilum and Malacosteus niger are examples. Howcver, therc are others more or less clearly defined to certain areas as the vicinity of oceanic islands, seamounts etc. Distribution patterns of most of these species still

-8- -8-

remain to be analyzed. Due to this and lack of space a single map is shown here. It represents the distribution of the family Bathylaconidae in the Atlantic Ocean according to NIELSE~,' 1912. Fishes of this family, Bathylaco nigricans and Herwigia kreffti, were found in almost all of our deep stations between the latitudes 35 0 N and 35 0 S, but aro entirely lacking in deep hauls beyond those boundaries (fig. 10). They most probab~ represent what I prelimi narily may call the "North Atlantic Deop vTater pattern". It seems, that ovon true pelagic deep-sea fishes follow individual distribution pattorns rather than to be ovonly distributed over vast areas.

'Concluding remarks. Studios on thc distribution of Atlantic midwater fish reveal a highly complicated three-dimensional cob-wob of distribution patterns. Actually, each specios had to develop its own specific pattern fitting the demands of its abiotic and biotic environment in order to survive. Howover, the threads of the web, represented by these spccific patterns, join to form larger patterns characteristic for more or less well defined faunal communitics populating different geographical areas of the ocean. These areas are separated from each other by boundaries due to physical conditions. Most trenchant along our transects were the o boundaries at abo~t 2o U, that is between thc t1-TO Central 'vater masses, ~nd that formed by the southern Subtropical Convergence. }mjor changes in the faunal composition oceur here. Other boundaries like that äividing subtropical and tropical species in the South Atlantic or the boundary bet\veen the "Guinean" and "Amazonian" regions at about 30 0 Waro much less well-defined. Much work has still to be done in order to understand the eeology of oceanic fishes. Life history • studies wanting for most of the species as weIl as a careful analysis of the various distribution studies one day may enable us such an . understanding.

Summary Investigations on board "vTalther Herwig" dealing with the taxonomy and distribution of Atlantic midwater fish corroborate thc findings of Amorican research groups like that of BACKUS and that of GIBBS. The main patterns of geoeraphical distribution encountered alone 4 transects are discussed and exemplified by 12 maps.

-9- -9-

Fishes are divided into northern-temperate, subtropical, tropical, "southern"and bathypelagic groups, e,ach of these groups showing a number of characteristic distribution patterns. Though still at the beginning of evaluation, patterns observed' so far exemplify the complexity of the environmental conditions responsible for the distribution of species and faunaI communities. References ~~~~~, ~. g., ~.E. 2~~~~~~~, ~. ~. g~~~~!~~, and ~. ~. ~~~~~~: The distribution of mesopelagic fishes in the equatorial and western North Atlantic Ocean. - J. mare Res. 28 (2): 179-201, 1970.

Q!E~~' ~. H.: Photonectes munificus, a new species of melanostomiatid , fish from the South Pacific Subtropical Convergence, with remarks on the convergence fauna. - Los Angeles County Mus., Contributions in Science No. 149: 1-6, 1968.

• !~~!:!:~, G.:, Ergebnisse der Forschungsreisen des FFS "Walther Herwig" nach Südamerika IX. Zur Systematik und Verbreitung der Gattung Lampadena Goode & Bean, 1896 (Osteichthyes, MYctophoidei; }trctophidae) im Atlantischen Ozean, mit Beschreibung einer neuen Art. - Ber.Dt. Komm. Meeresforsch. R1 (1-4): 271-289, 1970. , , !~~!:!:~, Q.: Ergebnisse der Forschungsreisen~•• etc. XVIII. Pseudoscopelus scutatus spec. nov., (Pisces, Perciformes, Trachinoidei, Chiasmodontidae), ein neuer Kreuzzahnbarsch aus dem tropischen Atlantik. - Arch. Fische reiwiss. 22 (3): 165-174, 1971. ~!~~!!~~, ~. Q.: Ergebnisse der Forschung.sreisen.'•• etc.' XX. Additional notes on Atlantic Bathylaconidae (Pisces, Isospondyli) with a new .' genus. - Arch. Fischereiwiss. ~ (1): 29~36, 1972.

" . Tab1e 1 : I \/ alt her Her 11 i 9 • Transect 1966 Distribution of fish species (upper figures) and spedimens (10wer figures) according to station and depth

0 0 0 Latitude 35 N 20 N 0 25°N Tot. 0 of No.of Station 174 175 176 177 178 179 180a 180b 181 182 183 184 185 186. 187 188 189 190 191 192 193 19 195 196 197 specilens hauls depth(.) -.2i T 20 .3838 25 ~ j- 100 -- - .7;74- - . --- -. ------.. ----.------I- •------'398- -- .. 50- • 33J - 401------9891 6 200 ------·----·-f······ ···1-'..·:r···;;;···;ii····jC· ..;l ;~ ;?i ;~ .. ;:;··;;:;···;!:····J·····T···!~!~··!~~1. 3!!3···;~~····5;··;;1:····6~·· . .~. )00 ••••••• 10' '41; ;1lr , ••••• 1011· Joro' i . '1' ,1 .. 1 . '1' t '. .'SY" . 459)" 6018 1 400 ••••••••••,.~ J: Jr j- !f ,...... ~ ""'1["" j[ .

:::: :: ::::1:: ::::::.:::::.:.:::::::::::::: ::::::::::::1::::::::::::::.::.:::: 15141 11 700

minutes*) 60 75 75 75 80 90 65 90 90 75 43 80 80 75 80 80 60 120 70 80 70 110 110 105 105 31161 25 fi shed 350 speCles

*) tiae for shooting aod hauling excluded

I step hauls, depth range fished Table 2 : • Wal t her Her \/ i 9' T r ans e c t 1 9 6 8 Distribution of flsh specles (upper figures) and specimens (lol/er figures) according to stations and depth

0 0 Latitude 35 ll 20 N 0° 25°5 ToLNo.of No.of Station 1 2 3 4 5 6 7 8 10 11 12 13 14 15 16 17 18 19 20 21 22 23 2; 25 26 27 28 29 30 specilens haula depth(.) ...... i .ll ...fg ...E ~ 11 1 2 941 25~ 706 510 65 3 4 2523 8 100 -•• - -~~- •••• --.--.--. ------_.------._--...---._------._------.- -._----.------._._------_.--- 11-!.J2.lQ 64 ~ 44 48..§l 51 15 29 16 18 23 6.1 22.J2. 11 667 185 381 232 617 462 56 182 tiö 112 190 9 11 m 183 87 4064 20 200 .-.- -~~.-~~~-.~~~--~~~ -.-._._--_.-._--.-_.~ ..------._---- -_._------.-._--~_ .._-.-.---.---.- ---.--.- --_._-._-..----.--..-- -..------_.------.li 34 60 .Jl 26.11 .1Q

160 _a._~_~222 "3M._'~_ ._._~ _~ 516 ._~_.108 163 207 1819 7 300 .••• -._••-••-•••••-••• .._._._.. .. •• __ ._._. __ . __ •• • • .._. ._.• _ .J.§. ...ll 16 J2 36 .l2. l 36 .l1 ..11 169 149 37 51 262 235 476 132 391 1902 9 400 -'•• -.--•••• --••••------._------.---_..•-Q---.-._-. ------.------._.._-.-----_._---.- .------_.------.-

500 --.- ---.-.•-.-----•••- --.-.--.•••--.-.---••.-- -.------•••--•.-••-••_- .-..--.--••-...•--.-•••-.-••••----••-.- .---••••------.------.-.------.---.--••-----.- ~~~ ~1~~ ~~~ ~~~ ~~ ~~~ ~ 150 131 1~9 625 996 778 751 646 617 548 408 212 265 333 268 3~5 589 655 ~66 18 600 -.------.---.--.---. ------.---.--••-----••-•••--._---.------.-.--- ••••••••------.-•••-----••----.-- -.-.-••------.------••--••••••-- ---••-.-••-••••-----.

700 ..•- •.--•••••..-••---. ~~--••-•••--••----.-.-.-- •.•--••------•••-••••• --.--.---•• --.--.----.------.-.-.-.---. --.-----•.•-••-•••-.-.-.------.--. --.--••••--•• -.-.-••- 52 308 1

::: -.-- :::::::::::::::::: ~::::::::::::::::::::::l:::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: ::::::::::::::::::::::::::::::::::: ::::::::::::: :::::::: 52 416 416 1 1000 --- .--••-•••--••------••---.---.------•.-••• -.------.-•••------.. ---•.••-.--.--.-.--.-.--.--••----.------.------.--.------•••-.-•••--- --.-••••------.---. 1100 1800 1900 .- --._ ..-._------.------....------.._.- .-.------.------.------.------_._------88 61 .JQ.. ill ...22 .J2**) ..!!l .:N 6875 8 2000 2947 940 ~23 ~02 170 363 1039 591 ToLNo.of 20 25 42 At6 52 75 102 102 126 04 125 88 97 90 95 85 83 57 47 43 53 65 62 58 67 76 83 73 70 426 specie! 72 spadas ot.~o.of T 272 449 270 334 308 1357 3369 1082 1917 406 1321 983 1969 1589 423 402 1120 655 325 170 485 999 372 4At2 At81 1111 1222 949 591 26373 :;PCClll:C:1S lBi nutes t) 60 60 60 60 30 45 90 60 60 60 30 45 60 60 30 30 60 60 60 30 60 60 45 60 60 60 45 60 60 fli shed * *) tll1e for shoot1ng and hauling excluded. **) in day light. "Table 3 :• Wal t her Her wi g' T r ans e e t 1 9 7 1 I I (Mar del Plata - Cape Town) Distribution of fish speeies (upper figures) and speeilens (lover figures) aeeording to stations and depth

Tot.No.of No.of Station Jlt8 350 354 358 363 367 371 376 380 381t 395 399 1t02 1t06 1t09 1t12 1t17 1t19 1t23 speeileAs hauls depth(l) .J1 11 2 159 -- .2~0_ -32 "7 -- 7:2 It_ -- 100 - - - • - jf -3j - 13- -24 - --~J~ --- "11 -- .J1 --- -.Jf -rli 31~ 492 113 1292 290 551 184 660 791 1t687 9 200 - - . -J§.- -- 26------.:l2.------.... _----- .. ------2108 3 817 538 753 300 - - foo ------}i ------10- .:.f2. " ------~- -42 668 54 803 668 1t16 2609 5 1t00 ------. --. ------

500 ------~ ------443 443 1 600 -- - -~------.:li------foo------708 1t25 64 2297 3 700 ------61 ------"'1"164-- .------.-- }1 ------.- foo ------• 713 633 131t6 2 800 -----~_._------_ .. _------560 560 1 -. -----. ------.. ------. ------.. - -1000 - - =il ------J!- -- . -..-.------. - --. -. ~ --. ---. .. 1663 2 998 665 1100 ------..--.. -. .. .. --. . -. -. -. --.. -. - 1200 ------. -- - --•------•------1300 ------•------•---•--•-----•---•---.-.-•-----.------

1400 - - -. ------. - ----.------• f------1500 -.--- - .---.------72 ------525 525 1 1600 .-

l700 ------•-•-- - -. ----•------.------

,,------66 ------... ------.. - ---. - -- - - 475 475 1 1900 ------69 ---- - :-l1- -.- 94- - - - 48 ------6S ------. - - - . ------1273 572 1193 149 839 5026 5 2000 ------• ------f------••-• - ---- 2100 ------7------m 526 1 2200 Tot.No. f 43 71 79 75 72 71 85 94 71 48 66 15 63 68 79 80 72 77 86 265 species 18 speeleS Tot.No.q! 998 1839 1765 1316 2673 1572 874 1193 2468 149 475 284 1537 839 1293 533 525 1459 1215 1lQ1l hauls speell1en~ .~nutes 15 60 45 45 45 60 45 90 45 30 120 30 45 90 30 105 60 30 45 flshee Table -\: • Wa1:...... :..t....:.;h...:e:...:r:..--..::H.....:e:...;r:....::..l1..:.i...:l91-·_...:..T...:..r..:a:...:n::...:..s:..• ...:::c.....:t:..----..:..1..!.9....l71-1~1~II (Cape TOlln - Madeira) Distribution of fish speeies (upper figur.s) and speei.ens (lover figures) aecording to stations and d.pth

Latitude 25°S 00 200 N Tot.No. No.of Station 421 431 435 439 443 441 451 455 459 463 467 471 478 482 486 490 494 498 502 506 510 512 speci •• hauls depth(lI.J1-1~ _8~ "1Ö~~ :7~0. 3 100------:li --- - • - m· .. -2l------63 --- )1- . - -59 f- • __ -- - 696 715 1436 904 880 1183 5814 6 200------6~ ------.- 300- --- }2 -- rli ----4b --- ~ --- ]i --- -60 --- ~4~ --- - • --- ali :O~O_ ~ -- 476 274 456 906 2112 4 400- -

500- -- ::.&i --::-- - -.------~ ------.- 600- - "" ~! "" "" - - -""- -""- "- "~ """eil - "- """- "11;! "- -1l~1 -----""""";:~;- · ~'- - 1177 700- - . --.-- -68 --- -6~ - . -- - .--.-----. ~ .114 ------522 476 1096 2094 3 --.-_. -.------_ .. ------.--_._---_.----.------

900- - 821 1000 ------~ ------. ------1100 ------. ------1200 ------. . .--. -.--. -. -.--. --. ---. -. ------. 121 1329 1 1300 ------.---- l329------1400 ------. . ------. ------. - - 1500 ------•------. ------. - . - • 1600 .

------1i1· - - -~- -- -112- ---. ------•------. ------______898. 4~4 983 • __ . _~_ ___ .~ ~9~4_ 5 1900. - -- - -~ 5 2000 _ ~ 538 116 1Q2 86 3409 - - -1Ö4------•---- ~------_. 754- 667 - . - 825 - ~9~0- 2 624 1286 2100 Tot.~o. 90 88 99 86 1~ 77 111 111 87 106 115 116 127 135 121 107 127 129 116 102 74 86 424 39 speC1es specie Tot.~o.spec,.. ()'25 1640 538 1565 624 833 898 1879 484 3069 983 2812 h286 2515 1329 2036 968 3056 754 667 631 825 30017 .jnut~s fl shed 60 .,L5 60 45 60 45 60 45 60 45 60 50 60 45 60 30 60 45 60 60 60 60 • 20 w 20

. 40 ." -.

30 30 e Atlantic Transects 20 o 1000 - 2100 m • 0-1000 m " 1966 - 1971 10 -_._- 10

I ___ \_ _ __ _~..l>!.---+=:"':':':::'------4_---A'--I --~ 186. o ~ ~16 167 t,~ • 1966 Stat. 174 -197/66 1O~• 1l&.-.1'__+-- 4--?&-r.3 ~ 1968 Stat. 1 - 30/68 459 w Q455 1971 Still. 348-512/71 0451 .447 •20 = J---nn-~r----+-----04:-+-3----\' 30~ 30 042'7

0395 0 • .~ ~{:> 40~• --+---J40 ~, •~ •,I •~ 50~• ;------1f------+------t------4------+--4lr50 • •~

-.-..:_---...,--=-~-=-=~ ·l...... Me------4:--• ..-.:wpa:.--=:."*,,.--:..---. • • ... - --=-=--- ..... _. 60 40 20 W 0 E 20 Fig.1 60 20 w 0 E 20

40~---+---"'"---+------+----/ ,. '. 40

30----+------+------+---i• 30 .~ 20Ac----+------+----~.-+-~ Astronesthes richardsoni

10 10

TROPICAL DISTRIBUTION (northern') 20 • •30 • 40~• ~ 50~ ~--+------+------+-----~------+-~50 • ~ ·l...... ~ __ ••...-.:-;-=-=-~~~ ~ .__...... - ...-.:~~ 60 40 20 W 0 E 20 Fig. 2 2cf~----+----__.}+-~ Pseudoscop-elu s scutatus

o negative Stations e.

o o

e . TROPICAL DISTRIBUTION (western) Fig. 3 From G. Kre tft, 1971 60 20 W 0 20

60 :'

40~---+------+------t----I "

30. 30 .~

20~::-----+------+----"""'" • Paraholtbyrnia ~y'anocel2bala ~lyjQOus asteroides 10 10

0 ~ 0 ~ 10 TROPICAL • ~10 DISTRIBUTION w 11 (eastern) • •20 i20 30~ •30

~ .. 40~• 40 I ~ • I 50 :•I. -=-=1If--=-~. =-=- •_.....---- -_ ...~---- - ....-.----- 60 40 20 W 0 E 20 Fig. 4 w 20 .

40~---+------+------+---=# .'. 40 "

30 +.-----1~---___i----____+_l• 30 • Hygophum hygomi •20~,-----+------+------' • Lampadena chavesi ~ 20 • Macroparalepis affinis

AN T/TROP/CAL 10 DISTRIBUTION ( intraspecific)

~ I-+-----.-----...... ---I-"III--I-.+---~--+------+---+ 40 I ~ •~

~ I ~=-=-:=-.-. :•••-~-- -~...... -=-=---.~.----- ... _:.-'-:~-=-:J~-=-:tI 60 40 20 W 0 E 20 Fig.5 • 60 w 0 E 20

50l 50

40~---+------,-.+------l----I " 40 .,.

30 Idiacanthus

)( fasciola •20 • atlanticus 20 )( both species 1 0 negative Stations 10 10 0

0 0 0 ·0 0 0 0 0 CD ANTITROPICAL 0 0 10 0 10 DISTRIBUTION 8 0 J> 0 (intrageneric) 0 20 0 0 20 19 • •30 30 • • • • X ••• 0 .-.!~ • • 40 ---- • 40

----+----·---1t--~50

60 40 20 w 0 E Fig. 6 , 20

50~ 50

40*------t------+------+----J

30 DiaQhus • ostenfeldi • Je metopoclampus • both species o negative Stations 10. 10

~ o ~ (: ~ 0 ~ Ci • lLl0 11 8 0 Je o· • 0 • 0 • 0 O 0 r o o Je. o ~ .. . --_. - _. --f40 • o I•I I • ----+---t-+o .l...... -:a--*.-----~_ ...... ,~...."~.. ~_J.~ ••-.--J-.•. ~lJ 60 40 20 W 0 E 2u Fig.7 w 0 E 20

,. 60 ·"l-+--~60

40~---+--·----+------+-----I .•. '. 40

. 30+-----+- 30 11 .~ o Astronesthes boulengtlri .. 20~---+------:-+------+--4 ~20 ...... ~ . • Opostomias micripnus I 10 - ...... +------+------+--~ 10

o ~O

SUBTROPICAL 10 ~10 CONVERGENCE • DISTRIBUTION • •20 20 30 30

o 40 •~ ~ 50 ~--+------+-----+-----+-----+----+50 •~ ~

Fig.8 w 0 20

60 "

40~----;.------+----- o 40 , ,

30~ 30 0' • Persparsia kopua • Bathophilus cwyanor • Eustomias bulbornatus 20 0• Diaphus diadematus 20

• 4••• l:::A Diaphus parri 0 Diaphus richardsoni 10 + Myctophum spinosum 10

0 0

Distribution 10 of INDIAN OCEAN ~10 Species • •~ 0 20 •20 0 30 30 IR 4 40 I 50 .50 l ..-.t-:-...... :.: -:Ir. W:~..~ •••=--=-:--- 60 40 20 w 0 E 20 Fig. 9 •• 60 40 20 ~.~-=-=iIfX:_WO~-=-::J.~':' ...------.•.• •

I •~ 5a~ ~

40-PI-----t------t------t------t ,.

'6 .:30+----+-- 30 10 , Distribution • I• 10 of a Family In ,~'1 •I NORTH ATLANTIC • DEEP WATER o • 11 .ai .' iÜ ~ 30~ ~30 •~ I 0 o • o I 0 40~ o O~-+-J- ~ ~O r·-

I ,J---+- ----+------+------+------+---~50 •~ ~ ~__ * ~::Jr.-.:llt:---: .-::..-=----.•J - _. -••:.:.-~~ 60 40 20 W 0 E 20 Fig. 10 (ac cording to J. G. N·ielsC?n, 1972 ) 60 40 20 W 0 E 20 ...... -~. ~ ... ----~----=-:.-~~~:.~.,.,. .... •"

, 40 > •

Ceratose o~elus •20~"'----1------+-----.IIIH-~ ~20 warmingi ~ 10 ~10

• °i BROADlY TROPICAl •• 1O~ DISTRIBUTION • •• • • • O~ • ~ • 30. 30 • • • • -+-----.---J I I I • rL----+---4---+---+--.--+iso ~.--:~. ~ -~.~ .--~-lJ 40 20 w 0.- E 20 Fig.11 • 60 20 •

r1l---""'60

50 50

40*----+--·--...:----+------+-----1 > • •

30-- 30

NotoscORelus 20~,..----+------~---.-...~ !:§plendens 20

10 10

~ mOADLY TROPICAL 1O~ • ~10 DISTRIBUTION • • • 20~ • • e , • ~ • r 30~ ~30 • • • • 40~ I (0 50~~ ~~--~~--~--~--~~I .~~~~-~~-=-=-t--"'--~--.~-----. ---~ _.~-~-D 60 40 20 W 0 E 20 Fig.12