ZoologicalJournal ofthe Linnean Society, 68: 99-109. With 4 figures Feb~ uary 1980 BathytyphZops seweZZi (Pisces: Chlorophthalmidae) a senior synonym of B . axorensis, from the eastern North Atlantic with notes on its biology N . R. MERRETT Institute ofoceanographic Sciences, Brook Road, Wormley, Godalming, Surrey GU8 5 U B AlcgPtedfor publication October I978 Evidence provided by six specimens of abyssal 5 a ~ h y ~ y ~ coltected h f o ~ ~ in the eastern North Atlantic indicates the conspecificity of the Atlantic E . azorensu with the Indian Ocean B . sewelli. The data also clarily the distinction between 5. sewelli and its congener, E. marionae. The benthic habit of E . sewelli is contirnied. Its reproductive biology is examined and the hermaphroditic pattern substantiated. The species is evidently a multiple spawner with a fecundity of around 6000 eggs. Observations in support of the probable epipelagic larval development of 5 . sewelli are dicussed. Stomach and intestinal contents analysis suggests that this species Feeds largely on benthopelagic crustaceans. Digenetic trematode parasites were found in the digestive tracts of three specimens. KEY WORDS: - Bathytyphlops sewelli - abyssal fish - taxonomy - reproduction - hermaphrodite - lecundity - feeding - parasites. CONTENTS Introduction . . . . . Materials and methods . . Keaults and discussion . . . Systematic considerations Environmental observations Biological observations . Reproduction . . . Feeding . . . . . Acknowledgements . . . Keferences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 100 101 10 1 107 107 107 108 108 109 INTRODUCTION The chlorophthalmid genus Bathytyphlops (sensu Sulak, 1977 ) is represented in the literature by two specimens of B . sewelli (Norman, 1939) from the tropical western Indian Ocean, 18 specimens of B . marionae Mead, 1958 from the tropical Atlantic and western Indian Oceans and two specimens of B . azorensis Geistdoerfer, Hureau 8c Rannou, 1970 from the tropical and temperate Atlantic Ocean. Sulak (1974) summarized the distribution and depths of capture of all 99 0024-4082/80/0200991 1 1/$02.00/0 0 1980 The Linnean Societv of London N. R.MERRE'IT 100 recorded specimens prior to those ( 7 reported by Golovan ( 197 6) from the slope 1 7 O 36'W, 1740-1920m). While previous waters off West Africa (09O 25", workers have indicated the close relationship of the three nominal species, the chart given by Sulak emphasises the salient difference in depth distribution between B. marionae, an essentially slope-dwelling species (living in soundings shallower than 2000 m or waters warmer than 4°C-Sulak, pers. commn), and the abyssal B . sewelli and B . atorensis. Anatomically the degree of development of the rakers on the first gill arch has been considered the principal character in separating B . marionae from its congeners, B . azorensis (Geistdoerfer, Hureau & Rannou, 1970; Sulak, 1974) and B . sewelli (Nielsen, 1966). The distinction between 5. sewelli and B . azorensis given by Geistodoerfer, Hureau & Rannou (1970) is less trenchant and it is therefore appropriate to reappraise the relationship of these species in the light of a further six specimens of abyssal Bathytyphlops collected from the tropical and subtropical eastern North Atlantic. The results clarify the taxonomy of the genus and add to the knowledge of its biology. MATERIALS AND METHODS The specimens of Bathytyphlops were collected in the course of benthic surveys off northwest Africa carried out by RRS 'Discovery' during 1969-77. The relevant sampling data are given in Table 1, while the localities of capture are shown in Fig. 1 within the context of the overall collections. Epibenthic sledges (BN2.4; 2.4 m wide x 0.8 m high and BN1.5; 2.3 m x 0.6 m) took one specimen each, while the remainder were caught in a 14 m headline length Marinovich semiballoon otter trawl (OTSB14).The BN 1.5 incorporates a mouth closing system, in which a blind occludes the entrance of the net at all times other than when the sledge is on the bottom (Aldred, Thurston, Rice & Morley, 1976), but the other two are open nets. Table 1. Collection data for six specimens of Bathytyphlops from the eastern North Atlantic. See Materials and methods for gear notation. ( 0 )denotes non-closing gear std tlon Date Position 7092 8933#3 913l#lO 954 1#6 251x1169 11x1175 191x1176 16lIVl77 2.i007'N, 19"lO'W 24°56.6'N, 1B0O1.2'W 20°15.1'N, 21'35.5'W 20°09.6'N, 21'43.3'W Gear Sounding range (m) BN2.4 OTSB14 BN1.5 OTSB14 331 1-35] 140) 2980-2990-(0) 3950-3952 3929-3929-(0) Number 1 2 1 2 Measurements were made directly from a sliding caliper to the nearest half millimetre. Standard length is used throughout and morphometric characters are expressed in percentages of it. Meristic counts were made from the left hand side of specimens only. Vertebral counts were taken from X-radiographs and exclude the urostyle. In addition to the 'Discovery' specimens, the following material was examined for comparative purposes :B. sewelli, holotype, 300 mm. British Museum (Natural History), BMNH 1939.5.24.458. BATHYTYPHLOPS FROM THE EASTERN NORTH ATLANTIC 101 v 'Canary Isiands 0 9131 c 0 - 15' 0 Cape & * - Q Verde o Islands - 0' .a - D - - 0 10" - . I l l l l l l l l l l l l l r l l Figure I Chai t of the eastern North Atlantic off northwest Africa showing the localities of capture of t h e 'Discovery' 5 ~ ~ ~ ~ among t ~ p other ~ ~ stations o p ~ sampled during 1969-1977 0, 0-500 m, 0, 500-1000 111, A, 1000-1500 m , H, 1500-2000 m, A, 2000-2500m, 7 2500-3000 m , v, 3000-3500 m , 0,3500-4000 m , 0,4000-4500 m , 5500 m *, B . azorensis, holotype, 2 10 mm. Museum National d'Histoire Naturelle, Paris, M N H N 1970-30. B . marionae, Virginia Institute of Marine Science, VIMS 03298, 111 mm, 37O05.4'N, 74O23.6'W, 1520 m sounding; VIMS 05232, 325 mm, 2S049'N, 88'27'W, 1289 m sounding. Supplementary data on B. marionae were kindly provided by K. J. Sulak who made selected measurements on the following VIMS specimens: VIMS 0492 1 , 180 mm, off British Guiana, 914 m sounding; VIMS 05233, 265 mm, 07O41'N, 53O35'W, 882 m sounding; VIMS 05234, 277 mm, 07O20'N, 53OO4'W, 692 m sounding; VIMS 05235,203 and 320 mm, 09'58'N, 76O29'W, 127 1 m sounding; VIMS 04964, 356 mm, 2 8 O 17'N, 7 7 O 10.5'W, 1106 m sounding. RESULTS AND DISCUSSION Systematic considerations The overall size range of the 'Discovery' specimens was 146-282 mm. Their vital statistics are given in Table 2. The ranges of characters for 'Discovery' specimens are compared with those published for B . marionae, B . sewelli and B . azorensis, together with data from hitherto unreported specimens of B. marionae (Table 3). Inspection of Table 3 reveals that the new eastern North Atlantic material overlaps several of the distinguishing proportions and counts previously N. R. MERREIT 102 Table 2. Morphometric measurements, in percentages of standard length, and meristic counts for six specimens of B a ~ ~ ~collected ~ ~ ~ from ~ l ‘Discovery’ o ~ s in the eastern North Atlantic Station number Standard length (mm) 9541#6 146 147 Head length Snout length Premaxillary length Eye diameter Interorbital width Snout to dorsal origin Snout to anal origin Snout to pelvic origin Snout to pectoral base Snout tO dnUS Pelvic to anal origin Anus to anal origin Depth of body at origin of dorsal Length base dorsal Length base anal Length pelvic Length pectoral Length raker in angle of 1st gill arch 22.6 7.5 17.5 0.7 8.6 38.1 65.4 35.3 21.2 43.2 32.5 22.6 12.3 11.6 17.8 16.1 20.9 2.3 22.8 6.8 17.7 0.7 8.5 38.1 67.7 35.0 21.1 42.9 34.4 24.8 11.6 12.2 16.3 17.0 21.8 2.6 12 12 15 8 12 14 6+1+12 64 24 + 40 68 Number dorsal fin rays Number anal fin rays Number pelvic fin rays Number pectoral fin rays Number branchiostegal rays Number gill rakers (1st arch) Number vertebrae - total Number vertebrae - precaudal Number lateral line scales I7 + caudal 8 13 15 6+1+12 65 25 + 42 70 913l#lO 193 21.5 6.7 16.6 0.8 8.8 38.9 68.4 35.0 20.7 43.5 34.2 25.4 11.9 11.4 16.1 14.5 20.7 1.7 11 14 8 12 15 7+1+12 65 24 + 41 70 8933#3 7092 282 25 7 265 21.8 7.4 17.5 0.6 8.6 40.9 69.5 35.8 21.6 44.7 32.9 23.0 13.2 12.8 16.7 16.3 20.2 1.6 22.6 7.2 16.9 0.6 8.7 40.0 66.4 36.0 21.7 44.5 30.2 21.1 12.8 11.3 17.0 16.2 21.1 I .5 21.6 6.7 16.3 0.5 8.5 39.9 67.4 35.6 20.7 44.1 33.2 24.8 13 15 8 13 15 6+1+13 63 23 + 40 68 11 16 8 13 14 6+1+12 63 23+40 69 12 13.1 11.7 18.1 16.8 18.6 1.5 17 8 14 15 6+1+12 64 23 t 4 1 70 used to separate the three species (viz. number of pectoral, dorsal and anal rays, lateral lines scales and gill rakers; relative proportions of body depth and length of dorsal fin base, see Mead, 1958, 1966; Nielsen, 1966; Geistdoerfer, Hureau & Rannou, 1970).As a result it is necessary to re-examine the status of each species in the light of the new data. The only diagnostic feature immediately apparent from Table 3 is the relative length of the anal fin base. This, when judged by the ratio-on-size approach of Pam (19561,is found to be an isometric character (Fig. 2), which is confirmed by a sizeon-size logarithmic regression giving a log linear relationship for each species (slopes; B . sewelli 1.015, Q = & 0.047; B . marionae 1.007, o=f-0.061). In addition, the values of the log of the anal base length when the log of the standard length = 2.000 (= 100 mm SL) are significantly different at the 95% level ( B . sewelli 1.22 1, Q= 0.015; B . marionae 1.123, Q= k 0.025)which isolates B . marionae (anal base length 12.1-14.4% SL) from B . azorensis and B . sewelli, whose proportions (15.5-17.6% SL) are matched by the range found in the ‘Discovery’ specimens (16.1-18.1% SL). Correspondingly, the number of rays in the anal fin of B . marionae is generally lower ( 13-14) than in the sewelli-azorensis-‘Discovery’ specimens ( 14-1 7 ), although the range found among the ‘Discovery’ material makes this no longer invariably diagnostic. The development of‘the rakers on the Table 3. Comparison of the ranges of morphometric measurements, in percentages of standard length, and meristic counts of the ‘Discovery’ Bathytyphlops with similar data for B . sewelli, B . arorensis and B . marionae. (a, Nielsen, 1966; b, Sulak, 1974; c, Geistdoerfer, Hureau 8c Rannou, 1970; d, Mead, 1958; e, unpublished data from VIMS material. ’”, Counts and measurements taken from type material during the course of this study) ‘Discovery’ specimens Standard length (mm) Head length Snout length Prerriaxillary length Eye diameter Interorbital width Snout to dorsal origin Snout to anal origin Snout to pelvic origin Snout to pectoral base Snout to anus Pelvic to anal origin Anus to anal origin Depth ofbody at originofdorsal Length base dorsal Length base anal Length pelvic Length pectoral Length raker in angle of 1st gill arch 146-282 21.5-22.8 6.7-7.5 16.3-17.7 0.5-0.8 8.5-8.8 38.1-40.9 65.4-69.5 35.0-36.0 20.7-21.7 42.9-44.7 30.2-34.4 21.1-25.4 11.6-13.2 11.3-12.8 16.1-18.1 14.5-17.0 18.6-2 1.8 1.5-2.6 Number dorsal fin rays Number anal tin rays Number pelvic fin rays Number pectoral tin rays Number branchiostegal rays Number gill rakers ( G t arch) Number vertebrae- total Numbervertebrae-precaudal Number lateral line scales 11-13 14-17 8 12-14 14-15 6-7+1+12-13 63-65 (23-24) + (40-42) 68-70 + caudal B. azorensts B sewellt 93a 23.0 7.7 18.5 0.9 9.3 40.0 67 36.5 148b 22 21oE B . mananae 300a 21.5 7.0 17.0 0.5 7 - 8.8 8 38 65 - - - 41.5 67 38.0 - 45.0 43.5 - 30.9 21.9 12.3 10.9 17.6 16.6 21.4 1.7’ - 12 17 8 I3 12 I2 17 8 13 13 8 14 16 .. 0.6 - - 32 - - 9.1 9.7 16.0 17.5 10.0 15.5 13 - - 3.8 11 16 8 12 14 7+1+11 62 24 + 38 c.70 1.8* 11 16 8 12 14 6+1+11 63 25 + 38 c.65 10 I7 1.9 22.1 7.1 - 0.7 8.0 40.4 67.1 7+1+12 7+1+12 65 64* 24 + 40* 67 263d 22.4 6.6 16.6 0.11 8.5 40.3 69.0 37.9 21.7 I 12.2 12.4 15.4 17.1 4.2 12 279d 23.1 7.5 17.0 0.7 8.6 39.1 69.6 37.5 21.3 - 22.0 16.0 12.7 14.0 16.7 20.1 - 13 14 8 13 17 15 b + lc(12-14) 62 63 L5+37 65 64 llOe 22.7 7.3 17.2 09 8.2 39.0 67.2 36.4 21.8 45.5 33.6 22.7 11.4 11.8 13.6 16.4 17.2 3.2 315e 22.8 7.3 16.2 0.5 8.2 37.4 69.5 38.0 21.2 46.4 35.2 24.4 13.6 13.3 13.3 13 13 13 14 12 8 8 SO@ - - - - - - 14.5 - - 3.3 15 15 15 7+1+13 7+1+13 65 62 25+40 24+38 64 66 13 16.8 20.6 3.5 I3 8 14 15 7+1+13 62 24+38 64 104 N. R.M E R R E I T Standard length (mm) Figure 2 Length of the anal fin base, as a percentage of the standard length, plotted against standard length in B. manonut (A,published data; A, VIMS data), B sewelli (0).E azorensu (m) and ‘Discovery’ Bathytyphlops (0).Data for closed symbols taken from Table 3, with the addition off, Norman, 1939. first gill arch, hitherto considered sufficient to distinguish marionae from both sewelli and azorensis, must now be qualified as a diagnostic feature. The degree of development of rakers on the epi-, cerato- and hypo-branchial bones varies and, as the photographs in Fig. 3 show, this is particularly marked in small specimens. The rudimentary nature of the gill rakers in larger specimens of B. murionue, however, serves as a useful confirmatory character (see Mead, 1958 for description). The relative length of the enlarged raker in the angle of the epibranchial and ceratobranchial bones can be used to distinguish marionae from both sewelli and azorensis, at least among specimens larger than 130mm. In contrast to the isometric growth of the anal fin base, however, this gill raker is strongly negatively allometric among the sewellz-atorensis-‘Discovery’ material group (Fig. 4),so it is of value only in relation to the size of specimen concerned. This allometry is confirmed by logarithmic regression as above, while the data for B. mmianae indicate again a log linear relationship (slope; 1.133, Q= 0.110), which is thus isometric. Individual data are not given by Golovan (1976)for his seven specimens (302-353 mm) of B . marionae, but the ranges shown for the length of the base of the anal fin (11.9-15.1% SL), the number of rays in this fin (12-14) and the length of the enlarged raker in the angle of the first gill arch (3.6-4.2% SL) are consistent with, and so serve to confirm, the features found here to be diagnostic for this species. The status of B. marionae is further strengthened by its predominantly slope-dwelling habit (p. 1001, which contrasts with the abyssal occurrence of the sewelli-azorensis-‘Discovery’ group of specimens. The comparison of the features above serves, not only to confirm the distinction of B. marionae, but also to unite the nominal species sewelli and atorensis. Two of these features (uiz. the number of anal fin rays and the relative Figure 3. Comparison o f t h e first gill arches o f t h e 'Discovery' Bathytyphlops (A, 146 mm: B, 147 mm; C, 193 inin; E, 257 m n ; F, 265 inin; G, 282 mrn) with the holotypes of E . azorensis (D, 210 mm) and B . rewvlli iH, 300 nun) and two specimens o f E . rnarzonac (I, 110 mm, J , 315 mm). j All removed from LHSexceptforD.)Scalelines= 10 mm. N. R.MERRETT 106 A .a* 4- - *a d h f m .a 3 ‘ Q 3- 2 - z A A 0 c 0 f In 2- .b tl x z *f .C 0 0 - 0 0 ’c f m a nI I- 0- 1 100 1 1 1 1 I50 1 1 1 l 1 1 200 1 1 1 1 1 250 1 1 1 1 1 300 1 1 1 1 351 Figure 4. Length of the raker in the angle of the first gill arch, as a percentage of standard length, plotted against standard length in B. manonae (A, published data; A. VIMS data), B . sewelli (01,B. azorewts (U)and ‘Discovery’Bdhytyphlops (0). Data for closed symbols is the same as given for Table 3 and Fig. 2, with the addition ofa”, Nielsen, 1966. length of the raker in the angle of the first gill arch) were among the four adopted by Geistdoerfer, Hureau & Rannou (1970) to distinguish B. azoremis from B. sewelli. The number of rakers on the first gill arch, their third diagnostic character, has already been invalidated by the range evident from the ‘Discovery’ data (Table 3). Finally they stressed the differences in the obscurity of the eye between azorensis and sewelli. Examination of the ‘Discovery’specimens suggests a regression in the conspicuousness of the eye with size. The degree of obscurity of the eyes of the holotypes of B . azorensis and B . sewelli matches this trend in relation to their length. At all sizes the skin over the eyes bears scale pockets, but in small specimens little opaque tissue separates this from the minute eye. With body growth the eye appears to become deeper set, but barely any larger. Overlying tissue renders it less conspicuous to the point where the holotype of B . sewelli led Norman ( 1939) to comment on “the complete absence of eyes” in describing the species. Hence the new data overrule the distinctions apparent to Geistdoerfer, Hureau & Rannou ( 19 701,while further comparison of the ‘Discovery’specimens with the type material of both species reveals no other differences worthy of specific rank. The discrepancy in number of caudal vertebrae between the two s ecimens of sewelli (38-Table 3) and the uzormis-‘Discovery’ fishes (40-42) is t e largest variation, but it is comparable with that observed in marionae (37-40). I t is considered to be within the range of individual variation for a species of Bathytyphlops, especially as the range in the total number of vertebrae from the ‘Discovery’ material conforms with the ranges in both B. azorensis and B. sewelli. Hence, all the available evidence indicates that B . azorensis is a synonym of B. sewellz. K BATHYTYPHLOPS FROM THE EASTERN NORTH ATLANTIC 107 Environmental observations With this new information, the known range of B . sewelli is extended into the Atlantic Ocean to give a geographic distribution broadly similar to that of B . murionae, except for the absence of B . sewelli in the western Atlantic (cf. Sulak, 1974: fig. 2). The lack of a swimbladder in B . sewelli and its robust skeleton are indicative of a bottom-living habit, which is confirmed by the capture of one ‘Discovery’B . sewelli in the mouth-closing epibenthic sledge (BN1.&see Materials and methods). A conductivity-temperature-depth profile taken at Stn. 9541#4, in the same area as one of the captures (Stn. 9541#6) and a few hours prior to it, gave a temperature of 2.3OC at 3944 m depth in a sounding of 3950 m. This confirms my initial conclusion (based on the maps in Fuglister, 1960) that all the ‘Discovery’ specimens were collected from temperatures lower than 3OC. The substrates at two of the collection sites, sampled by box corer, were calcareous clay (Stn. 8933#3) and pink calcareous ooze (Stn. 9 131#10)(S.E.Calvert,pers. commn). Biological observations Reproduction Nielsen ( 1966) studied the reproductive biology of several of the ipnopine species in detail, but B . sewelli was not among them. Examination of the ‘Discovery’ material largely confirms the reproductive pattern he revealed. Thus B. sewelli is hermaphrodite. The testicular portion of the ovotestes in adults is a relatively inconspicuous line of light-coloured tissue running mid-ventrally along the length of the gonad. First maturity had not been attained in the three smaller specimens ( 146-196 mm). Oocyte development was in the early stages prior to chorion formation (oocyte diameter < 0.1 mm) and the testicular string was barely distinguishable in the unstained gonad. Maturation of the ovotestes was more advanced among the three larger ‘Discovery’specimens (257-282 mm) and the holotype. While none of the testicular portions were swollen with milt, the ovarian sections were in the growth phase of maturation with yolk formation in the developing ova well advanced (c. 0.6-0.8 mm diameter). The left hand side of the ovotestis of the 282mm fish was removed and the most advanced generation of eggs counted. There were 2940 present and, with the right hand side of similar size, a total fecundity in this generation of about 6000 eggs would be expected. This contrasts with the considerably lower count of 300 or less in Zpnops and Bathymtcrops (Nielsen, 1966). Apparently two other generations, represented by resting oocytes of < 0.1 mm diameter and ova in the chorion formation stage of oogenesis of 0.1-0.2 mm diameter, were present in this and the other maturing ovaries examined. Also, free in the lumen of the left hand ovary of the 282 mm specimen were ten residual ovulated eggs, presumably of a previous spawning. Although collapsed, they were noticeably larger than any developing ova, probably > 1 mm diameter. A certain amount of degeneration had occurred internally, but a conspicuous oily globule was present in each. Whether or not such a globule would provide buoyancy in healthy eggs is uncertain, yet the observation lends added credence to the identification of B u t ~ ~ p h i o plarvae s from near surface waters (Okiyama, 1972). In the related genus Zpnops, Sulak (1977 ) found that ovarian eggs were not buoyant and he used this observation, together with the low fecundity, to conclude that the eggs developed close to the bottom. In contrast, the 20-fold 108 N.R. MERRETT’ greater fecundity of B . sewelli argues for epipelagic development. Oval tailless spermatozoa, about twice the size of blood cells, were found in the testis of this 282mm fish, which accords with their morphology in B. marionae and other Ipnopinae (Nielsen, 1966). It would appear from these specimens that B. sewelli is a multiple spawner, which attains first maturity around 200-250 mm. The interpretations given by Nielsen ( 1966) on synchronous hermaphroditism in this group remain substantially unaltered in the light of the present data. While the collection is too small for observations on seasonality, for future reference it is noted that the specimens with ripening gonads were caught in November (cf. Sulak (1977) regarding Zpnops). Feeding Stomach and intestinal contents were removed and examined separately. No stomachs were more than half full or contained undigested organisms. The only identifiable remains of prey were crustacean. Fragments were found amongst the amorphous digested contents of the stomachs and/or intestines of all but one fish. The intestine of the 193 mm specimen contained mysid remains, while decapod material was identified from three fishes ( 2 5 7 mm-stomach, 265 mm-stomach, 282 mm-intestine; the contents of the two largest specimens have already been reported by Marshall & Merrett (197 71, under the preliminary diagnosis of B . uzorensis). Decapod remains could be recognised as natant forms in the 257 and 282 mm specimens. Together with these fragments in the largest fish were a few foraminiferan shells of the Globigerina type. This was the only bottom substrate present in any of the specimens and it might have originated from the decapod prey. Such general absence of sediment and the active nature of the identifiable prey suggests that, while B sewelli without buoyant means must be a benthic species, it has a marked preference for pelagic food. It presumably feeds on benthopelagic organisms swimming close to the sea-land interface and must compete with the more active buoyant fishes which can “hover and explore” for food (see Marshall & Merrett, 1977). Trematode parasites were present in the digestive tracts of the three specimens in stomach; 265 mm-6 in stomach, 2 in mentioned above (257 mm-1 intestine; 282 mm- 10 in intestine). They were identified as undescribed opecoelid digeneans of the genus Neolebouria Gibson, 1976 (D. I. Gibson, pers. commn). It is noteworthy that this genus, which was described from antarctic material (Gibson, 19761, should have an abyssal semi-tropical representative. ACKN 0WLED GEMENTS The loan of specimens was kindly arranged from the British Museum (Natural Histo ) by Mr A. C. Wheeler, from the Museum National d’Histoire Naturelle, Paris,?y D r J.-C. Hureau and from the Virginia Institute of Marine Science by Dr K. J. Sulak. Dr Sulak also generously provided data from additional material. I am grateful to Dr D. I . Gibson (British Museum (Natural History)) for identifying the gut parasites and to Dr A. L. Rice (1.0.S.) for identifying the food organisms. Finally, I wish to thank Mr J. R. Badcock and Mr P. M. David (I.O.S.),Dr J . G. Nielsen (Zoologisk Museum, Copenhagen), Dr J.-C. Hureau and Dr K. J. Sulak for their helpful criticism of the manuscript. BATHYTYPNLOPS FROM T H E EASTERN N O R T H ATLANTIC 109 REFERENCES ALDRED, R. G., THURSTON, M. H., RICE, A. L. & MORLEY, D. R., 1976. An acoustically monitored opening and closing epibenthic sledge. Deep-sea Research, 2 7 : 167-1 74. FUGLISTER, F. C . , 1960. Atlantic Ocean atlas of temperature and salinity profiles and data from the International Geophysical Year of 1957-1958. The Woods Hole Oceanographic Institution, Wood.i Hole Atlas Series, 1 : 209 pp. G I B S O N , D. I . , 1976. Monogenea and Digenea from fishes. Discovery Reports,36: 179-266. GEISTDOERFER, P., HUREAU, J.-C. c! RANNOII, M., 1970. Deux poissons abyssaux nouveaux raptures darts I’Atlantique Nard et Est: Eathjtyphlops azorensi,! n. sp. (Ipnopidae) et Lycenchelys labradorensis n. sp. (Zoarcidae). Bulletin du Musium National d’tltstoire Naturelle, ( 2 )42: 452-459. GOLOVAN, G. A,, 1976. Redkie i vpenye obnaruzhennye na materikovom sklone Zapadnoi Afriki khryashchenvye i kosmye ryby. (Rare and firstly recorded chondrostean and teleostean fishes of the continental slope ofWest Africa.) Trudy Instituta Okeanologii tm P. P . Shirshou, 104. 277-3 1 7 . (In Russian). MARSHALL, N. B. & MERRETT, N. R., 1977. The existence o f a benthopelagic fauna in the deep sea. In: M. V. Angel, (Ed.), A Voyage ofDiicovety. George Deacon 70th Anniuersav Volume: 483-497, Supplement to DeepSea Research, 24. MEAD, G . W., 1958. Ichthyology-Three new species of-archibenthic iniomous fishes from the western North Atlantic. Journal ofthe Washington Academy ofSciences, 48: 362-372. MEAD, G. W., 1966. Family Ipnopidae. In G. W . Mead (Ed.). 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