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Ichthyological Bulletin
J.L.B. Smith Institute of Ichthyology
ISSN: 0073-4381
Num. 62, 1995
J.L.B. Smith Institute of Ichthyology
Ichthyological Bulletin No.62, February 1995

A REVIEW OF THE SPECIES OF THE GENUS NEOBYTHITES (PISCES: OPHIDIIDAE) FROM THE WESTERN INDIAN OCEAN, WITH DESCRIPTIONS OF SEVEN NEW SPECIES
by
Jorgen G. Nielsen^1

^1 Zoological Museum, University of Copenhagen, Universitetsparken 15, DK 2100 Copenhagen O, Denmark.


Code Number: FB95001
Size of Files:
    Text: 85.4K
    Graphics: line drawings (gif) - 179.5K

ABSTRACT

Species of the ophidiid genus Neobythites from the western Indian Ocean are revised based on 225 specimens. The majority of specimens came from the 17th cruise (1988-89) of the Soviet R/V VITYAZ. Twelve species are recognized: N. analis Barnard, 1927 (southern Africa; lectotype designed), N. crosnieri sp.n. (Madagascar), N. kenyaensis sp.n. (east coast of Africa), N. malhaensis sp.n. (Saya de Malha Bank), N. meteori sp.n. (Socotra Id.), N. multistriatus Nielsen and Quero, 1991 (Reunion and Rodrigues Islands), N. natalensis sp.n. (South Africa and Madagascar), N. somaliaensis sp.n. (Somalia), N. steatiticus Alcock, 1893 (northern Indian Ocean), N. stefanovi Nielsen and Uiblein, 1993 (Red Sea to Gulf of Oman), N. trifilis Kotthaus, 1979 (Socotra Id.), N. vityazi sp.n. (Madagascar). All 12 species are illustrated, and a key is included.

This paper is the first part of a revision of the genus Neobythites. A second and third part will treat the species from the East Indian/Pacific Oceans and the Atlantic Ocean.

INTRODUCTION

During the 17th cruise of the Soviet R/V VITYAZ, in the southwestern Indian Ocean (1988-89), many specimens of the bathyal, ophidiid genus Neobythites Goode & Bean, 1886 were collected. In treating this material I soon found several problems. Cohen & Nielsen (1978) recognized 14 species stating that "...a much needed revision undoubtedly will show that some of these are conspecific", but they did not foresee any undescribed species.

Until now five described species have been caught in the western Indian Ocean: N. analis Barnard, 1927, N. multistriatus Nielsen & Quero, 1991, N. stefanovi Nielsen & Uiblein, 1993, N. steatiticus Alcock, 1893 and N. trifilis Kotthaus, 1979. The present VITYAZ material included four species of which two appear to be undescribed. A general review of the genus was indicated and an additional ca. 150 Neobythites specimens from the western Indian Ocean were examined. The total material (about 225 specimens) comprises 12 species, of which seven species are new to the western Indian Ocean.

The next step was to ascertain whether any of the seven species were described from outside of the western Indian Ocean. Examination of about 500 non-western Indian Ocean specimens revealed about 30 species including only 13 that were already described. Except for N. steatiticus, none of the western Indian Ocean species were represented among these 30; consequently of the 40-45 species of Neobythites, ca. 25 are yet to be described. The discovery of this large number of new species is the result of the availability of much new material from many localities. Also most of these specimens were collected recently and show distinctive pigmentation characters that are useful in species recognition.

The complete revision of the genus Neobythites will be published in three parts according to the geographical distribution of the species: 1) western Indian Ocean, from the east coast of Africa to the longitude through the tip of southern India; 2) eastern Indian and Pacific Oceans, and 3) Atlantic Ocean. Consequently, most general remarks will be left for the last part.

METHODS

Measurements were taken as specified by Hubbs and Lagler (1958) with the following exceptions: head length is measured from the upper jaw symphysis to the hind margin of the opercle. Eye diameter is measured as the horizontal diameter of the eye window. Gill-rakers are counted from the outer side of the anterior arch, with Roman numerals showing the short or rudimentary rakers and Arabic numerals the long or developed rakers; the long raker in the angle of the gill arch is included in the count for the lower limb. Longest gill filaments are measured on the anterior arch. Vertebral and median fin-ray counts are based on radiographs. The ural elements are included as "1" in the vertebral count. Sex of specimens examined is indicated by F (female) or M (male). Sizes are given as standard length (SL).

Abbreviations for institutional collections follow Leviton et al. (1985) and Leviton and Gibbs (1988). They did not include the Zoological Museum, Moscow State University, which is abbreviated ZM MGU.

Neobythites Goode & Bean, 1886

Neobythites Goode & Bean, 1886:600 (type species N. gilli Goode & Bean, 1886 by monotypy).
? Tetranematopus
Gunther, 1887:100 (nomen nudum).
Watasea
Jordan & Snyder, 1901:765 (type species N. sivicola Jordan & Snyder, 1901 by original designation).

DIAGNOSIS: Mouth terminal or subterminal. Body relatively short. Head length more than 50% of preanal length. Cycloid scales covering head and body. Horizontal diameter of eye window equal to or slightly shorter than snout. Opercular spine strong and straight. Median basibranchial tooth patches two. Sagittal otolith characterized by long, shallow, and straight sulcus with flat and well marked colliculi and normally a pseudoostial opening; ostium about twice as long as cauda, but of same height; ostium and cauda most often well separated, but some species with fused colliculi (Schwarzhans, 1994). Developed rakers on anterior gill arch 6-20. Pseudobranchial filaments 1-8. Dorsal fin rays 88-114, originating above vertebrae 2-6. Anal-fin rays 72-97. Pectoral fin not divided in 2 parts, with 22-34 rays. Ventral-fin rays 2 in each, placed below preopercle. Precaudal vertebrae 11-14.

ADDITIONAL CHARACTERS common to the subfamily Neobythitinae and tribe Neobythitini: Dorsal, caudal and anal fins united. Origin of dorsal fin above pectoral fin. Lateral line uninterrupted, placed high on body. Nostrils midway between upper lip and eye, the anterior with small skin flap, the posterior most often a mere hole. Branchiostegal rays (7)-8. Supraneurals or predorsal bones not developed. Anterior-most neural spine short; neural spines of 3rd to 8th precaudal vertebrae almost as long as that of 2nd vertebra. Parapophyses developed on posterior 6-8 precaudal vertebrae, pleural ribs on vertebrae 3-4 to 12-14 and epipleural ribs on anterior 5-8 vertebrae. Stomach muscular; pyloric caeca 5-15. Genital papilla small. Peritoneum black or dark brown.

REMARKS: The genus Watasea Jordan and Snyder, 1901 was established for those Neobythites species with two spines on the hind margin of preopercle, leaving those with only one spine in Neobythites. Smith and Radcliffe (Radcliffe, 1913: 139) did not find this character important enough to justify a separate genus; however, they seemed to consider it a subgeneric character, as all the species with two spines are cited with Watasea in parentheses after Neobythites. I cannot recognize Watasea as a valid genus, because the number of preopercular spines of present material varies between 0 and 3.

ONTOGENETIC CHANGES: In some species, the hind margin of the vomer tooth patch changes with growth from concave to convex (N. multistriatus [Fig. 7], trifilis [Fig. 12], and vityazi [Fig. 13]). The development of the dorsal fin ocellus was shown for N. stefanovi (Uiblein et al. 1994), the only species for which a developmental series (27 - 198 mm SL) is available. This species also shows a positive allometric growth of body depth and of the length of the gill filaments on the anterior arch. The spines on the hind margin of the preopercle in N. natalensis become stronger with growth.

RELATIONSHIPS: A phylogenetic analysis has not been undertaken for the ophidiid genera. However, Cohen and Nielsen (1978:36) indicated that Neobythites may be related to Bassogigas Goode and Bean, 1896 and to Benthocometes Goode and Bean, 1896.

Relationships (or rather, similarities) have been indicated for the present 12 species, but with about 25 Neobythites species still to be described, the proposed relationships are rather preliminary.

BIOLOGY: Most of the material was trawled from the upper part of the continental slope from 80-760 m; one specimen of N. analis was caught at a depth of 1830 m west of Cape Town, and some of the Red Sea specimens of N. stefanovi came from 800 m. Figure 1 shows the distribution of the western Indian Ocean species.

Stomach and intestinal contents consisted of remains of crustaceans (most of which are unidentifiable), gastropods, bivalves, scales of fishes and dorsal felt from polychetes.

In most species, females were found to be much more common than males. The largest eggs found were 0.5 mm in diameter.

The relatively large eyes and the presence of ocelli in most of the species indicate that sight plays an important role for these fishes (see under N. stefanovi).

    Figure 1. Distribution of western Indian Ocean Neobythites species.

Key to Neobythites species of the western Indian Ocean

---------------------------------------------------------------------------
1a.  Preopercular spines 0-1(weak)  ..................................    2
1b.  Preopercular spines 2 (rarely 1 or 3)  ..........................    6

2a.  Dorsal fin with 2 ocelli  .....................    N. kenyaensis sp.n. 
2b.  Dorsal fin with 1 ocellus  ......................................    3

3a.  Vertical bar below ocellus; origin of dorsal fin above operculum  ..   
                                            ..........  N. malhaensis sp.n.
3b.  No vertical bar; origin of dorsal fin above pectoral fin ........    4

4a.  Dorsal fin ocellus above anus; developed gill-rakers 6  .........
                                            .............. N. meteori sp.n.
4b.  Dorsal fin ocellus above anterior part of anal fin; developed          
     gill-rakers 10-14  ................................................  5

5a.  Edge of anal fin transparent; body with 4 ill-defined, brown, 
     vertical bars; longest gill filaments 15-19% head length  ..........   
                                             ............... N. steatiticus
5b.  Edge of anal fin black; no dark vertical bars on body; longest gill    
     filaments 9-16% head length  .........................    N. stefanovi

6a.  Teeth needle-like; black area behind eye.   .............  N. trifilis
6b.  Teeth granular; no black area behind eye  .......................    7 

7a.  Dorsal fin with ocelli or large dark blotches  ..................    8 
7b.  Dorsal fin without ocelli or large dark blotches ................   10

8a.  Dorsal fin with 7 or 8 and anal fin with 3 or 4 dark blotches; 
     4-10 dark vertical bars on body; developed gill-rakers 12-15  ......   
                                             ...........   N. multistriatus
8b.  Dorsal fin with 2 distinct or about 6 indistinct ocelli; 
     no dark bars on body; developed gill-rakers 8-10  ...............    9

9a.  Dorsal fin with 2 distinct ocelli ...............   N. crosnieri sp.n. 
9b.  Dorsal fin with about 6 indistinct ocelli  ......  N. natalensis sp.n.

10a. Distal part of dorsal and anal fins black; longest gill filaments      
     10.5-14% head length  .........................  N. somaliaensis sp.n.
10b. Distal part of dorsal fin not black, anal fin black or not; 
     longest gill filaments 4.4-7.7% head length ....................    11 

11a. Distal part of anal fin black; posterior margin of vomer tooth patch   
     usually concave or straight  ............................    N. analis
11b. Anal fin not black; posterior margin of vomer tooth patch convex  .... 
                                            .............  N. vityazi sp.n.
---------------------------------------------------------------------------

Neobythites analis Barnard, 1927, (Fig. 2)

    Figure 2. Neobythites analis: a) F, 195 mm, ZMUC P77816. Vomer tooth patches; b) lectotype, F, 136 mm, SAM 12487; c) F, 204 mm, USNM 19759. Left sagitta: F, 212 mm, RUSI 12356, d) median view, e) anterior view. Scale bars 1 mm.

Neobythites macrops (non Gunther): Gilchrist & Thompson 1914:89 and 1917: 416; Gilchrist & von Bonde 1924: 19; Smith 1950: 363, fig. 1019; Hulley 1972: 230.
Neobythites analis Barnard, 1927:879 (type locality, 22 miles north of Tugela River mouth, Natal, South Africa); Shcherbachev 1980: 159, fig. 15 (in part); Nielsen & Cohen 1986: 349, fig. 96.18; Shcherbachev et al. 1986: 203.

LECTOTYPE (here selected): SAM 12487, F, 136 mm SL; 22 miles north of Tugela River mouth, Natal; M/V PIETER FAURE st. 11667; 115 m.

PARALECTOTYPES (Note: According to P.A. Hulley, of the South African Museum (per. comm.) the total number of syntypes was 27. Of these 2 were sent to BMNH and 5 to USNM from which 1 syntype was later transferred to ZMUC. The remaining 20 paralectotypes are in SAM.): BMNH 1927.12.6.67, 124 mm and ZMUC P77716, 106 mm (formerly SAM 12487); same data as lectotype. SAM 12084, 114 mm; 11 miles ESE from Bird Island, Algoa Bay; M/V PETER FAURE st. 1129; 99 m. SAM 12087, 134 mm and BMNH 1927.12.6.68, 92 mm; 11.5 miles; NE Nanquas Peak, M/V PIETER FAURE st. 14020; 106 m. SAM 12088, 95 mm; 8.5 miles NE from Bird Island, Algoa Bay; M/V PIETER FAURE st. 14034; 104 m. BMNH 1902.5.28.16, 170 mm; off Natal coast; 366 m.

ADDITIONAL MATERIAL: RUSI 12356 (3; F, 182+?-212 mm); off Durban (29 degrees 45'S, 31 degrees 26'E); M/V PICKLE st. 184, otter trawl; 350 m; 11 Aug 1920. RUSI 12355, F, 205+? mm; off Cape Town (33 degrees 37'S,16 degrees 55'E); M/V PICKLE st. 533, reversible trawl; 1830 m; 29 Nov 1921. USMN 319759, (6; 3 F, 3 M, 133-204 mm); off southern Mozambique (25 degrees 12'S, 34 degrees 04'E); R/V ANTON BRUUN cr. 8, st. 397A; bottom trawl; 230-295 m; 29 Sep 1964. ZM MGU P-15114 (2; F, 176-197 mm); off southern Mozambique (25 degrees 24.2'S, 34 degrees 17.2'E); R/V PROFESSOR MESIATZEV cr.5, haul 77; 330 m; 31 Jan 1976. ZMUC P77816-820 (5; 3 F, 2 M, 160-195 mm); off Natal (29 degrees 46.1'S, 31 degrees 23.1'E); R/V AFRICANA st. A 4709, T-03; bottom trawl; 350 m; 26 Aug 1986.

DIAGNOSIS (based on lectotype; data for 6 paralectotypes and 19 additional specimens in parentheses): Dorsal-fin rays 107 (99-107); anal-fin rays 87 (84-89); 2 (rarely 1 or 3) spines on preopercle; developed gill-rakers 10 (9-11 ); longest gill filaments 7.3 (5.4-7.7%) head length; pseudobranchial filaments 5 (3-6); teeth granular; precaudal vertebrae 14 (13-14); outer part of anal fin black; no ocelli or vertical bars. (Shcherbachev (1980: 160) mentioned that the presence of the dark band on the edge of the anal fin is apparently dependent on sex or age, based on observation of two females and one male. However, Shcherbachev's male specimen is referable to a new species, N. vityazi (p. 14). The best field-character for analis is thus still valid.)

DESCRIPTION (additional meristic and morphometric data are given in Tables 1-8 at end of text): Snout varies from pointed to relatively blunt; horizontal eye diameter slightly longer than snout. Lateral line distinct. Preopercle in majority of specimens with 2 flat spines; 2 specimens with 1 spine on right side 2 on the other; 1 specimen with 1 spine on each side, and 1 specimen with 3 spines on 1 side and 2 on the other. Ventral-fin rays not reaching halfway to anal fin. Rear edge of vomer tooth patch concave in smaller specimens; larger fish with almost straight hind margin (Fig. 2) and in one specimen slightly convex. Pyloric caeca 5-7. Caudal-fin rays 8. Gill-rakers iii-iv, 1-3 + 7-9,v-viii. Sagittal otolith (Fig. 2) twice as long as high with almost smooth dorsal and ventral edges; sulcus closed and deep; cauda half the length of ostium; sometimes without complete separation of colliculi; curvature of inner surface flat.

Colour in fresh specimens mottled brown. Outer edge of anal fin black in its entire length. Lateral line pale, surrounded with dark brown pigment. Dark peritoneum shows through skin. All type specimens examined are uni- formly yellowish without any pigment. RUSI material from 1920 yellowish but anal fin still black.

REMARKS: Only a few specimens with stomach contents, consisting of dorsal felt from polychetes and remains of crustaceans. The material examined comprises 13 females, 5 males and 7 unsexed.

COMPARISONS: Barnard (1927: 880) states that N. analis is very close to N. macrops Gunther, 1887. Other authors (see references above) consider N. analis a junior synonym of N. macrops. Compared to other species from the western Indian Ocean, N. analis seems closest to N. vityazi, both having two preopercular spines, distinct lateral line and no ocelli. They differ from each other by the form of the vomer and by the black edge of the anal fin in N. analis (see key to species).

DISTRIBUTION: Known from off the coast of southern Mozambique southward and round Cape of Good Hope to west of Cape Town (Fig. 1). It is the only species of Neobythites known from the eastern Atlantic. Its northern-most locality is slightly overlapping the area of distribution of N. vityazi. Further south it is the only recorded species of Neobythites. The Indian Ocean specimens were caught in 99-366 m, and the Atlantic specimen at a depth of 1830 m.

Neobythites crosnieri sp.n.(Fig. 3)

    Figure 3. Neobythites crosnieri: a) holotype, M, 192 mm, ZM MGU P-18904. b) Vomer tooth patch of holotype. Left sagitta of holotype: c) median view, d) anterior view. Scale bars 1 mm.

HOLOTYPE: ZM MGU P-18904, M, 192 mm; off SW Madagascar (22 degrees 19.1'S, 43 degrees 6.1'E); R/V VITYAZ cr. 17, st. 2644, bottom trawl; 330-335 m; 2 Dec 1988.

PARATYPES: MNHN 1992-534, F, 113 mm; off SW Madagascar (22 degrees 29'S, 43 degrees 39'E); A. Mauge, coll., R/V STELLA MARINA, 125-250 m; 10 Mar 1968. MNHN 1992-532, F, 160 mm; off SW Madagascar (15 degrees 24.5'S, 46 degrees 2'E); A. Crosnier, coll., R/V VAUBAN, P.4, trawl 43; 250-265 m; 7 Nov 1972. ZMUC P77823, M, 125 mm; off NW Madagascar (15 degrees 20.5'S, 46 degrees 9.5'E); A. Crosnier, coll., R/V VAUBAN, P.4, trawl 45; 310-350 m; 7 Nov 1972. MNHN 1992-533, M, 165 mm; off W Madagascar (18 degrees 54'S, 43 degrees 55'E); A. Crosnier, coll., R/V VAUBAN, trawl 88; 280-310 m; 24 Nov 1973.

DIAGNOSIS (based on holotype; data for paratypes in parentheses): Dorsal-fin rays 98 (94-98); anal-fin rays 82 (79-82); 2 spines on preopercle; developed gill-rakers 9 (8-10); longest gill filaments 5.0-6.4% head length; pseudobranchial filaments 4 (4-5); vomer tooth patch triangular (Fig. 3); teeth granular; precaudal vertebrae 14 (13-14); total vertebrae 58 (57-58); posterior half of dorsal fin with 2 ocelli; outer part of posterior half of anal fin black; no vertical bars on body.

DESCRIPTION (data for paratypes, if different from holotype are given in parentheses; additional meristic and morphometric data are given in Tables 1-7 and 9): Snout slightly longer than eye window; upper jaw reaches just posterior to eye. Lateral line indistinct. Ventral-fin rays reaching 2/3 (1/2 - 2/3) distance to anal fin. Caudal fin rays 8. Pyloric caeca 11. Gill-rakers iii, 1 + 8,vi (ii, 1-2 + 7-8,vii). Sagittal otolith (Fig. 3) twice as long as high, with smooth dorsal and ventral edges; sulcus closed; cauda half length of ostium; colliculi partly separated; curvature of inner surface rather strong.

Colour of body mottled brown. Dorsal fin with 2 distinct black ocelli placed well behind anus. Outer edge of posterior dorsal fro, caudal fin and posterior anal fin dark. Edge of mouth, oral cavity and peritoneum dark. Dark brown pigment along lateral line and at base of dorsal fin.

REMARKS: Remains of crustaceans were found in the stomach and intestine of 3 specimens.

COMPARISONS: Among the species of Neobythites with 2 spines on preopercle and ocelli on the dorsal fin, N. crosnieri seems most similar to N. natalensis, both having 2 or more ocelli on dorsal fin, none on anal fin, 8-10 developed gill-rakers and a subtriangular tooth patch on vomer. N. natalensis differs from N. crosnieri in having about 6 indistinct ocelli on the dorsal fin.

DISTRIBUTION: Known from five localities off the west coast of Madagascar, where it was caught on the upper slope between 125 and 350 m of depth (Fig. 1).

ETYMOLOGY: N. crosnieri is named in honour of Mr. A. Crosnier of the Museum National d'Histoffe Naturelie, Paris who made an excellent collection of fishes from off Madagascar and collected all the paratypes of this new species.

Neobythites kenyaensis sp.n. (Fig. 4)

    Figure 4. Neobythites kenyaensis: a) holotype, 124+ mm, RUSI 13952. b) Vomer tooth patch of holotype. Left sagitta of holotype: c) median view, d) anterior view. Scale bars 1 mm.

Neobythites steatiticus (non Alcock): Norman 1939:76 (in part)

HOLOTYPE: RUSI 13952, 124+ mm (caudal tip missing); off Kenya (2 degrees 50'S, 40 degrees 31'E); P.C. Heemstra, coll., R/V FRIDTJOF NANSEN; bottom trawl; 275 m; 17 Dec 1980.

PARATYPE: BMNH 1939.5.24.1439, F, 103+ mm (caudal tip missing); off Zanzibar (5 degrees 34'24"S, 39 degrees 14'05"E); R/V JOHN MURRAY st. 105, Agassiz trawl; 238-293 m; 11 Jan 1934.

DIAGNOSIS: No spines on preopercle; developed gillrakers 9-10; longest gill filaments 7.4-8.3% head length; pseudobranchial filaments 2-3; vomer tooth patch diamond-shaped (Fig. 4); precaudal vertebrae 13; dorsal fin with 2 ocelli, the first one located anterior to anus.

DESCRIPTION (data for paratype, if different from holotype are given in parentheses; additional meristic and morphometric data are given in Tables 1-7): Pectoral-fin rays 29; first dorsal-fin ray above vertebra 4 (5); first anal-fin ray below vertebra 16 (15) and below dorsal-fin ray 22 (20); 1st ocellus placed on dorsal-fin rays 14-20 (12-17); 2nd ocellus placed on dorsal-fin rays 36-43 (35.43). Snout blunt and equal in length to eye window; maxilla reaches vertical at rear margin of eye. Lateral line indistinct. Ventral-fin rays reaching 1/3 distance to anal fin. Teeth in upper jaw needlelike and in lower jaw granular (teeth granular in both jaws of paratype). Pyloric caeca 12. Gill-rakers ii,2 + 7-8,vii. Sagittal otolith (Fig. 4) 1.5 times as long as high, with somewhat undulating anterior and posterior part; ostium a little less than twice of length of cauda, from which it is almost separated; curvature of inner surface moderate.

Colour of body mottled brown, with 7 broad, indistinct, vertical bars. Dorsal fin with 2 distinct ocelli; posterior third of dorsal fin, caudal fin and posterior 5/6 of anal fin black. Brown pigment on lower and upper jaws. Peritoneum brown and eye lens blue.

REMARKS: Unidentifiable remains of crustaceans were found in the intestine of the holotype.

COMPARISONS: N. kenyaensis differs from all other species of Neobythites by lacking spines on the preopercle and by having 2 ocelli on the dorsal fin. It is difficult to determine which of the four other western Indian Ocean species of Neobythites that lack spines on the preopercle is most similar to N. kenyaensis.

DISTRIBUTION: Known from 3 degrees S to 6 degrees S off the east coast of Africa at 240-300 m of depth (Fig. 1).

ETYMOLOGY: The specific name refers to the type locality.

Neobythites malhaensis sp.n. (Fig. 5)

    Figure 5. Neobythites malhaensis: a) holotype, M, 123 mm SL, ZM MGU P-189 15. b) Vomer tooth patch of holotype. Left sagitta: paratype, F, 132 mm; ZM MGU P-18917; c) median view; d) anterior view. Scale bars 1 mm.

Neobythites sp.: Shcherbachev et al. 1986: 203..

HOLOTYPE: ZM MGU P-18915, M, 123 mm; Saya de Malha Bank (11 degrees 02'S, 62 degrees 15'E); 250 m. The data for the holotype are incomplete, but it was most probably caught during the same PROFESSOR MESIATSEV cruise as the two paratypes (Yu. Sheherbachev pers. comm.).

PARATYPES (all from Saya de Malha Bank): ZM MGU P-18916, F, 117 mm; 11 degrees 08'S, 62 degrees 16'E; R/V PROFESSOR MESIATZEV trawl 476; 235-239 m; 7 Oct 1977. ZM MGU P- 18917, F, 132 mm; 11 degrees 06'S, 62 degrees 19'E; R/V PROFESSOR MESIATZEV trawl 478; 240 m; 7 Oct 1977. ZMUC P77840, F, 135 mm; R/V FIOLENT; (no other data).

DIAGNOSIS (based on holotype, data for paratypes given in parentheses): Origin of dorsal fin above operculum [vertebrae 2 or 3]; a diffuse, black, vertical bar on body below ocellus on dorsal fin; dorsal-fin rays 99 (99-103); anal-fin rays 78 (78-82); preopercle with or without a weak spine; developed gill-rakers 12-13; longest gill filaments 7.7 (7.58.9%) head length; pseudobranchial filaments 3 (3-4); vomer tooth patch triangular (Fig. 5); needle-like teeth on jaws; precaudal vertebrae 13; total vertebrae 57 (57-59).

DESCRIPTION (Data for paratypes, if different from the holotype, are given in parentheses; additional meristic and morphometric data are given in Tables 1-7 and 10): Snout subequal in length to eye window; maxilla ends well behind rear margin of eye. Lateral line indistinct. Preopercle with one weak spine (one paratype [ZM MGU P- 18917] with no spine). Ventral-fin rays reaching 2/3 distance to anal fin. Caudal fin rays 8. Pyloric caeca 15. Gill-rakers iii,3 + 9-10,vi (ii-iii,3-4 + 9-10,v-vii). Sagittal otolith (Fig. 5) 1.5 times as long as high, with undulating dorsal edge and rest of edge smooth; ostium twice as long as cauda with an incomplete separation between colliculi; curvature of inner surface moderate.

Colour of body brownish with an indistinct dark, vertical bar below dorsal fin ocellus, which is placed posterior to a line through anus. Peritoneum brown, and eye lens blue.

REMARKS: ZMUC P77840 with remains of crustaceans in the intestine.

COMPARISONS: N. malhaensis is not closely related to any of the western Indian Ocean species of Neobythites, but it may be closer to one of the species from the eastern Indian Ocean.

DISTRIBUTION: All specimens were caught off the Saya de Malha Bank north of Mauritius at depths between 235 and 250 m (Fig. 1).

ETYMOLOGY: The specific name refers to the type locality.

Neobythites meteori sp.n.(Fig. 6)

    Figure 6. Neobythites meteori: a) holotype, F, 102 mm, ZMH 5621. b) Vomer tooth patch of holotype; scale bar 1 mm.

Neobythites unimaculatus (non Smith & Radcliffe): Kotthaus 1979: 13, fig. 462.

HOLOTYPE: ZMH 5621, F, 102 mm; off Socotra Id., northern Indian Ocean (11 degrees 33.9'N, 52 degrees 54'E); R/V METEOR st. 102, Agassiz trawl; 175-337 m; 20 Dec 1964.

DIAGNOSIS: Dorsal-fin rays 91; anal-fin rays 75; caudal-fin rays 7; no spines on preopercle; developed gill-rakers 6; longest gill filaments 13.5% head length; pseudobranchial filaments 3; vomer tooth patch triangular (Fig. 6); precaudal vertebrae 13 (total vertebrae 53); dorsal fin with a large ocellus above anus.

DESCRIPTION (see also Tables 1-7): Pectoral-fin rays 27; first dorsal-fin ray above vertebra 5; first anal-fin ray below vertebra 15 and below dorsal-fin ray 20; ocellus placed on dorsal-fin rays 17-25. Head length 22% SL; depth at anus 19% SL; horizontal eye window 4.9% SL; preanal length 45% SL; predorsal length 24.5% SL; snout-ocellus 39% SL. Snout equal in length to eye window; maxilla ending below posterior part of eye. Lateral line indistinct. Ventral-fin rays reaching almost halfway to anal fin. Pyloric caeca 5. Teeth needle-like on anterior half of jaws and granular posteriorly. Gill-rakers iii-iv + 6,ii. Sagittal otolith in poor condition.

Body yellowish. Eye lens blue; peritoneum brown.

REMARKS: Intestine with unidentifiable crustacean remains. The holotype and only known specimen is a female with well developed ovaries containing eggs of about 0.5 mm in diameter.

COMPARISONS: Among the species with one distinct ocellus on the dorsal fin and no spines on the preopercle, N. meteori seems much closer to an undescribed species caught off Lombok, Indonesia, than to any of the species in the western Indian Ocean. It differs from the Lombok species mainly by the position and size of the dorsal-fin ocellus. Kotthaus (1979: 13) identified the holotype as N. unimaculatus Smith & Radcliffe, 1913, but this species differs in many characters (e.g., it has 2 spines on the preopercle and 8-11 developed gill-rakers).

DISTRIBUTION: Known only from off Socotra Id. in 175-337 m (Fig. 1).

ETYMOLOGY: The specific name refers to the R/V METEOR which caught the holotype.

Neobythites multistriatus Nielsen & Quero, 1991 (Fig. 7)

    Figure 7. Neobythites multistriatus: a) holotype, F, 178 mm, MNHN 1988-1945; b) F, 76 mm, ZM MGU P-18914. Vomer tooth patches: c) F, 76 mm, ZM MGU P-18914; d) paratype, F, 165 mm, MNHN 1988-1946; e) holotype. Left sagitta paratype: F, 15 cm, ZMUC P77809, f) median view, g) anterior view. Scale bars 1 mm.

Neobythites multistriatus Nielsen & Quero, 1991: 194, fig. 1, type-locality, off Reunion, 20 degrees 57.9'S, 55 degrees 14.5'E).
Neobythites fasciatus (non Smith & Radcliffe): Shcherbachev et al. 1986: 203.

MATERIAL: Type material listed in Nielsen and Quero (1991). New material: ZM MGU P-18914, F, 76 mm; off Rodrigues Id. (19 degrees 23'S, 61 degrees 31'E); R/V AKADEMIK KURCHATOV cr. 36, st. 3765, bottom trawl; 410 m; 24 Apr 1983.

DIAGNOSIS: (based on type material only): Dorsal-fin rays 106-109; anal-fin rays 91-93; 2 spines on preopercle; developed gill-rakers 13-15; longest gill filaments 5.8-6.3% head length; pseudobranchial filaments 5; teeth granular; dorsal fin with 7-8 and anal fin with 34 more or less distinct dark blotches; body with 9 indistinct vertical bars.

DESCRIPTION (see also Tables 1-7 and 11; N. multistriatus is described in detail by Nielsen & Quero, 1991:194): Snout slightly longer than eye window. Maxilla ends well behind posterior margin of eye. Lateral line indistinct. Preopercle with 2 strong spines. Ventral-fin rays reaching halfway to anal fin. Caudal fin rays 8. Teeth granular. Vomer tooth patch varies in form apparently due to size (Fig. 7). Pyloric caeca 14-15. Gill-rakers ii-iii,3-4 + 10-12,iv-vi. Sagittal otolith (Fig. 7) 1.5 as long as high with rounded anterior and posterior edges and a flat dorsal rim; sulcus long and narrow; ostium twice as long as cauda, from which it is only partly separated; ostial canal not developed; sagitta thick with strong curvature of inner surface.

Colour of body generally light brown. Dorsal fin with 7-9 more or less well defined dark blotches or ocelli, 3-4 on anal fin and 7-9 diffuse dark, vertical bars on body.

REMARKS: Stomach and intestinal contents consisted of crustaceans and a few fish bones. All 5 specimens examined are females. In addition to the 4 type-specimens, Tables 1-7 & 11 show data from a fifth specimen (Fig. 7B) caught off Rodrigues which differs in several characters from the types of N. multistriatus. Some of the differences such as shape of vomer tooth patch, number of ocelli in dorsal fin, bars on body and the presence of a tube around the opening of the posterior nostril might be due to the small size (76 mm) of the Rodrigues specimen (the types are 149+-178 mm SL). Additional material may show whether differences in fin ray counts and number of pseudobranchial filaments are caused by geographical variation.

COMPARISONS: N. multistriatus seems most similar to N. fasciatus Smith & Radcliffe in Radcliffe, 1913, which also has several dark vertical bars on the body, dark blotches or ocelli on the dorsal and the anal fins, and 2 spines on the preopercle. N. fasciatus has fewer developed gill-rakers (8-10) and anal-fin rays (84-88).

DISTRIBUTION: The 4 type-specimens were all caught off Reunion between 300 and 490 m. The specimen from off Rodrigues was trawled at a depth of 410 m (Fig. 1).

Neobythites natalensis sp.n. (Fig. 8)

    Figure 8. Neobythites natalensis: a ) holotype, 53 mm, ZMUC P77824; b) paratype, 53 mm, MNHN 1992-535. c) Vomer tooth patch of holotype. Scale bar 1 mm.

HOLOTYPE: ZMUC P77824, 53 mm; off Natal, South Africa (25 degrees 20'S, 35 degrees 17'E); R/V GALATHEA st.202, herring otter trawl; sounding 590 m, fishing depth near bottom; 21 Feb 1951.

PARATYPES: MNHN 1992-535, (3, 53-69 mm); off W. Madagascar (15 degrees 20.5'S, 46 degrees 9.5'E); A. Crosnier, coll., R/V VAUBAN, P.4, trawl 45; 310-350 m; 7 Nov 1972.

DIAGNOSIS (based on holotype; data for paratypes in parentheses): Dorsal-fin rays 101 (100-102); anal-fin rays 84 (84-86); 2 spines on preopercle; developed gill-rakers 8 (8-9); longest gill filaments 3.5 (3.3-4.5)% head length; pseudobranchial filaments 2 (2-4); vomer tooth patch triangular (Fig. 8); teeth granular; precaudal vertebrae 13; 6 more or less distinct ocelli on dorsal fin; a brown line through snout and eye.

DESCRIPTION (data for paratypes, if different from the holotype, are given in parentheses; additional meristic and morphometric data are given in Tables 1-7 and 11): Snout blunt (snout pointed) and equal in length to eye window; maxilla ends below rear margin of eye (in the smaller paratype [Fig. 8B] the maxilla ends well behind the eye). Lateral line indistinct. Preopercle with 2 rather weak, flat spines (largest paratype with upper preopercle spine strong and lower weak). Ventral-fin rays reaching about halfway to anal fin. Caudal fin rays 8. Pyloric caeca 14. Gill-rakers iii, 1 + 7,fii (iii,1+7-8,ii-v). Sagittal otolith in poor condition.

Body yellowish; 6 brown blotches or ocelli on dorsal fin (many indistinct blotches on dorsal fin). Diffuse brown line from front of snout through eye (faint, horizontal rows of brown pigment on anterior part of body). Eye lens blue and peritoneum brown.

COMPARISONS: Among the species with 2 spines on preopercle and 2 or more dark spots on the dorsal fin, N. natalensis seems most similar to N. crosnieri (cf. p. 5).

DISTRIBUTION: Off Natal and off west coast of Madagascar at depths between 310 and 590 m (Fig. 1).

ETYMOLOGY: The specific name refers to the type locality.

Neobythites somaliaensis sp.n. (Fig. 9)

    Figure 9. Neobythites somaliaensis: a) holotype, F, 205 mm, ZM MGU P- 18905; b) F, 74 mm, ZM MGU P-18913. Vomer tooth patches: c) M, 74 mm, ZM MGU P-18913; d) paratype, F, 123 mm, ZM MGU P-18911; e) holotype, F, 205mm, ZM MGU P-18905. Left sagitta: paratype, F, 172mm, ZM MGU P-18911: f)median view, g) anterior view. Scale bars 1 mm.

HOLOTYPE: ZM MGU P-18905, F, 205 mm; the Brothers Islands, SW of Socotra (12 degrees 16.6'N, 53 degrees 8.2'E); P/V VlTYAZ cr. 17, st. 2560, bottom trawl; 380 m; 27 Oct 1988.

PARATYPES (13, 123-198 mm): ZM MGU P-18906, F, 152 mm and ZMUC P77836, F, 173 mm; same data as holotype. ZM MGU P-18907, F, 170mm; SW of Socotra (11 degrees 33.2'N, 52 degrees 54.6'E); R/V DMITRY STEFANOV cr.3, trawl 34; 300 m; 25 Dec 1988. ZM MGU P-18908 (2; F, 160-165 mm) and ZMUC P77837-838 (2; F + M, 139-168 mm); SW of Socotra (11 degrees 31.4'N, 53 degrees 1.2'E); R/V DMITRY STEFANOV cr.3, trawl 35; 405-442 m; 25 Dec 1988. MGU P-18909, M, 177 mm; SW of Socotra (11 degrees 35.3'N, 39.6'E); R/V DMITRY STEFANOV cr. 3, trawl 42; 400 m; 28 Dec 1988. ZM MGU P-18910, 198 mm; SW of Socotra (12 degrees 21.6'N, 53 degrees 16.5'E); R/V DMITRY STEFANOV cr.3, trawl 50; 370 m; 2 Jan 1989. ZM MGU P-18911 (2; F, 123-172 mm) and ZMUC P77839, M, 140 mm; SW of Socotra (12 degrees 14.8'N, 53 degrees 6.2'E); R/V VITYAZ cr. 17, st. 2830, bottom trawl; 420-395 m; 16 Jan 1989. ZM MGU P-18912, M, 132 mm; Gulf of Aden off Democratic Yemen (14 degrees 45.2'N, 50 degrees 18.2'E); R/V DMITRY STEFANOV cr.3, trawl 75; 490 m; 2 Feb 1989.

Additional (non-type) referred specimen: ZM MGU P-18913, M, 74 mm; SW of Socotra (11 degrees 35.3'N, 52 degrees 39.6'E); R/V DMITRY STEFANOV cr.3, trawl 42; 400 m; 28 Dec 1988.

DIAGNOSIS (data for paratypes in parentheses): Dorsal-fin rays 104 (103-109); anal-fin rays 87 (87-91); 2 flat spines on preopercle; developed gill-rakers 9/10 (9-12); longest gill filaments 11.5 (11-14)% head length; pseudobranchial filaments 5 (5-8); vomer tooth patch subtriangular (Figs. 9D and E); teeth granular; precaudal vertebrae 13 (13-14); no ocelli on fins; no dark vertical bars on body; distal part of vertical fins black; 2 diffuse, brown, horizontal lines from tip of snout through eye to upper part of operculum.

DESCRIPTION (data for paratypes in parentheses; additional meristic and morphometric data are given in Tables 1-7 and 12): Snout blunt and equal (shorter than or equal) in length to eye window; maxilla ends just behind eye. Lateral line distinct. Ventral-fin rays reaching halfway to anal fin. Caudal fin rays 8. Pyloric caeca 12 (11-15 in 6 paratypes). Gill-rakers ii-iii,2 + 8-9,v-vi. Sagittal otolith (Fig. 9) about twice as long as high, with rather smooth edges; ostium nearly twice length of cauda; colliculi separated; small ostial canal developed; curvature of inner surface moderate.

Body yellowish with scattered brown pigmentation; lateral line surrounded by brown pigment. Peritoneum and branchial cavity dark; eye lens green.

The 74 mm specimen (Fig. 9B) is not considered a paratype as it differs slightly in several meristic and morphometric data from those of the type-specimens of N. somaliaensis. It is consequently excluded from the diagnosis. Tables 1-7 & 12 show differences in number of dorsal-fin rays, vertebrae, pseudobranchial filaments, and developed gill-rakers and morphometric characters. Differences in morphometric characters may be due to the much smaller size of this specimen. It also has the preopercle with 2 strong and pointed spines; posterior nostril with low rim; and hind margin of vomer tooth patch straight (Fig. 9C). Especially the latter character indicates that this specimen may belong to a species different from N. somaliaensis, as it is normal in Neobythites that if there is an ontogenetic change in the form of the vomer tooth patch it goes from juveniles with a concave hind margin to adults with a straight or convex margin (cf. Fig. 2). Sagittal otolith in bad condition. I have refrained from describing a new species on the basis of this single, small specimen. More material may show its status.

REMARKS: Gastropods and crustacean remains were found in the intestine of 5 specimens.

COMPARISONS: Among the species with 2 preopercle spines, no ocelli on the dorsal fin and no dark bars on the body, N. somaliaensis seems closest to N. analis which also has granular teeth and a subtriangular vomer tooth patch with concave hind margin. However, N. analis differs in having shorter gill filaments, shorter head and fewer anal and pectoral-fin rays.

DISTRIBUTION: All specimens were caught between Somalia and Socotra Island at depths between 300 and 490 m (Fig. 1).

ETYMOLOGY: The specific name refers to the area off which the type material was caught.

Neobythites steatiticus Alcock, 1893 (Fig. 10)

    Figure 10. Neobythites steatiticus: a) F, 113 mm, USNM 46756; b) vomer tooth patch of same specimen. Left sagitta F, 127 mm, USNM 256238; c) median view, d) anterior view. Scale bars 1 mm.

Neobythites steatiticus Alcock, 1893: 181, pl. IX, fig. 3 (type locality, east coast of India, 15 degrees 4.7'N, 80 degrees 25.7'E); Alcock 1898: pl. 21, fig. 2; Alcock 1899: 82; Beaufort & Chapman 1951: 417, fig. 67 (description of syntypes of N. malayanus Weber, 1913); Menon & Yazdani 1968: 149; Shcherbachev 1980: 162.

Holotype: ZSI 13435, F, 129 mm; Bay of Bengal (15 degrees 4.7'N, 80 degrees 25.7'E); R/V INVESTIGATOR st. 137, Agassiz trawl; 234 m; 3 Mar 1893. USNM 46756 (formerly ZSI 13475), F, 113 mm; Bay of Bengal; R/V INVESTIGATOR. USNM 256238 (formerly ZSI 13474), F, 127 mm; Bay of Bengal (13 degrees 51.2'N, 80 degrees 28.2'E); R/V INVESTIGATOR st.162, Agassiz trawl; 265-458 m; 30 Jan 1894. BMNH 1895.1.3.3 (formerly ZSI 13477), F, 143 mm; Bay of Bengal (13 degrees 1.1'N, 80 degrees 36'56"E); R/V INVESTIGATOR st. 170, Agassiz trawl; 196 m; 16 Feb 1884. BMNH 1895.1.3.4, F, 114 mm; Bay of Bengal (13 degrees 51.2'N, 80 degrees 28.2'E); R/V INVESTIGATOR st. 162, Agassiz trawl; 265-458 m; 30 Jan 1894. BMNH 1904.5.25.1, M, 150 mm; Gulf of Oman (25 degrees 32'N, 57 degrees 47'E); 320 m; F.W.Townsend, coll. BMHN 1910.1.31.7-10 (4; 3 F + 1 M, 132-145 mm); Gulf of Oman (25 degrees 35'N, 57 degrees 47'E); 311 m; F.W.Townsend, coll. BMHN 1910.1.31.23, F, 130 mm; 10' west of Dabai, Oman, Arabian Gulf; 267 m; F.W.Townsend, coll. (the Register and the labels give different localities but both in the Arabian Gulf off Oman. I consider the depth indication "14 fins" a mistake.).

DIAGNOSIS (based on holotype; data for paratypes in parentheses): Dorsal-fin rays 88 (88-93); anal-fin rays 75 (72-76); no spines on preopercle; developed gill-rakers 12 (11-14); longest gill filaments (15-19)% head length; pseudobranchial filaments 2-3 (2-4) (long); vomer tooth patch boomerang-shaped (Fig. 10); precaudal vertebrae (12); teeth granular to slightly needle-like; dorsal fin ocellus posterior to anus; middle part of anal fin black, with a pale band both distally and proximally; body with 3-4 indistinct, broad, dark vertical bars.

DESCRIPTION (based on 11 specimens, 113-150 mm SL, including holotype; additional meristic and morphometric data are given in Tables 1-7 and 13): Snout slightly pointed and equal in length to eye window; maxilla ends well behind eye. Lateral line indistinct. Pectoral-fin reaches vertical at anal fin origin. Ventral-fin reaching 1/2-3/4 distance to anal fin. Caudal fin rays 8. Teeth usually granular, but a few specimens with needle-like teeth in premaxilla. Pyloric caeca 8-9. Gill-rakers i-iii,2-5 + 8-10,vi-ix. Sagittal otolith (Fig. 10) 1.5 times as long as high with smooth edge; ostium twice as long as cauda; colliculi separated and rather broad; curvature of inner surface flat. Alcock (1893, pl. IX, fig. 3) showed a distinct ocellus on dorsal fin and 4 broad, irregular, vertical bars on body, all of which have now disappeared.

REMARKS: The material examined comprises 9 females and 2 males. The 127 mm specimen from the Bay of Bengal has relatively large eggs, 0.5 mm in diameter. Five of the specimens examined are from the Bay of Bengal, not far from the type-locality, and the other 6 specimens are from off Oman. Only one character seems to show geographical variation: Head length is 28-31% SL in specimens from the Bay of Bengal and 25.5-27 % SL in specimens from off Oman (Table 5). It should also be noted that the latest caught specimen is from 1910 so the colour pattern is much faded.

COMPARISONS: Among the species with one distinct ocellus on dorsal fin and 0-1 spines on preopercle, N. steatiticus is most similar to N. stefanovi (see next species) and to N. malayanus Weber, 1913. Beaufort & Chapman (1951: 417) considered N. malayanus and N. unimaculatus Smith & Radcliffe, 1913 as junior synonyms of N. steatiticus. However, N. malayanus has 8-11 more slender developed gill-rakers, shorter gill-filaments (5-15 % head length) and a weak, hidden spine on the preopercle. N. unimaculatus belongs to the group of species with 2 distinct spines on the preopercle, and N. malayanus has been discussed above.

DISTRIBUTION: Known from the Bay of Bengal, Gulf of Oman and Arabian Gulf at depths of 196-458 m (Fig. 1).

Neobythites stefanovi Nielsen & Uiblein, 1993 (Fig. 11)

    Figure 11. Neobythites stefanovi: a) M, holotype, 166 mm, ZM MGU P-18923. Vomer tooth patches: b) paratype, 61 mm, SMF 26429; c) paratype, F, 94 mm, SMF 26427; d) paratype, M, 158 mm, USNM 309008; e) paratype, F, 163 mm, ZMUC P77843; f) holotype, M, 166 mm, ZM MGU P-18923. Left sagitta: paratype, F, 180 mm, ZM MGU P-18928: g) median view, h) anterior view. Scale bars 1 mm.

Neobythites stefanovi Nielsen & Uiblein, 1993:110 (type locality: Gulf of Aden); Uiblein et al., 1994.
Neobythites steatiticus (non Alcock): Norman, 1939:76 (in part).

DIAGNOSIS: (based on the 29 type-specimens, 42-198 mm): Dorsal-fin rays 89-94; anal-fin rays 73-78; no spines on preopercle; developed gill-rakers 10-13; longest gill filaments 9.7-16% head length; pseudobranchial filaments 3-6; teeth needle-like (rarely granular); precaudal vertebrae 12; one ocellus on dorsal fin posterior to anus (poorly developed in specimens less than about 50 mm SL); outer part of vertical fins black in larger specimens.

DESCRIPTION (additional meristic and morphometric data are given in Tables 1-7): Snout rather blunt, shorter than horizontal diameter of eye window; maxilla ends behind eye. Anterior nostril with small flap; posterior nostril with or without low rim. Branchiostegal rays 8. Lateral line distinct and pale. Ventral-fro rays reaching 3/4 distance to anal fin. Caudal fin rays 8. Teeth needle-like. Rear edge of vomer tooth patch concave, straight or convex (Fig. 11); the 163 mm paratype (Fig. 11 E) shows an aberrant vomer tooth patch with a concave hind margin, a state that is normally found in much smaller specimens. Gill-rakers i-iv,24 + 8-10,v-viii. Sagittal otolith (Fig. 11 g & h) 1.5 times as long as high, with smooth edges; ostium twice length of cauda; colliculi broad and separated from each other; ostial canal present; curvature of inner surface flat.

Body mottled brown. Gill cover, peritoneum and pectoral fins dark.

REMARKS: A comparison of Red Sea specimens with the type material from the northern Indian Ocean revealed some slight differences (Nielsen and Uiblein, 1993). Uiblein et al. (1994: 18, Fig. 4) showed that the development of the dorsal-fin ocellus in Red Sea specimens can be divided into eight stages, which are positively correlated with size, and that the smaller (younger) stages occur at greater depths than the larger (older) stages. They concluded that "A well developed ocellus reduces the mortality risk from visually hunting predators ... and thus may allow N. stefanovi to adapt to increasing predation risk in shallower habitats."

The stomach and intestinal contents comprised crustaceans and gastropods. More than half of the specimens examined were too small for sexual identification.

COMPARISONS: Among the species with one large ocellus on the dorsal fin and lacking spines on the preopercle, N. stefanovi seems closest to N. steatiticus. They share many meristic and morphometric characters (Table 1), but differ from each other in adult coloration. In N. stefanovi the margin of the dorsal and anal fins is black and there are no dark bars on the body; in N. steatiticus there is hardly any black margin on the dorsal fin, the edge of the anal fin is transparent, and there are broad, irregular, dark, vertical bars on the body. Furthermore, the gill filaments are shorter in N. stefanovi (5.4-16 vs 15-19% of head length).

DISTRIBUTION: Known from 11 localities in the Gulf of Aden and from one in the Gulf of Oman; depth distribution 80-549 m. The Red Sea specimens come from ten localities with depths of 434-804 m (Fig. 1).

Neobythites trifilis Kotthaus, 1979 (Fig. 12)

    Figure 12. Neobythites trifilis: a) holotype, F, 157 mm, ZMH 5620. Vomer tooth patches: b) F, 117 mm, ZMUC P77826; c) holotype. Left sagitta: F, 178 mm, ZM MGU P-18900: d) median view, e) anterior view. Scale bars 1 mm.

Neobythites trifilis Kotthaus, 1979:11, fig. 461 (type-locality 11 degrees 33.9'N, 52 degrees 54'E); Shcherbachev et al. 1986: 203.

DIAGNOSIS (based on 70 specimens, 117-190 mm SL including the holotype): Dorsal-fin rays 99-105; anal-fin rays 83-89; 2 strong spines on preopercle; developed gillrakers 7-10; longest gill filaments 9.5-14.5% head length; pseudobranchial filaments 3-7; teeth needle-like; precaudal vertebrae 12-13; black area behind eye and on gill cover; no ocelli or vertical bars on body.

DESCRIPTION (additional meristic and morphometric data are shown in Tables 1-7 and 14): Snout blunt and slightly shorter than eye window; maxilla ends posterior to eye. Lateral line distinct. Ventral-fin rays reach halfway or less to anus. Caudal fin rays 8. Shape of vomer tooth patch changes with growth (Fig. 12 b & c). Gill-rakers i-v,1-2 + 6-8,v-ix. Sagittal otolith (Fig. 12 d & e) elongate, twice as long as high with smooth edges; cauda half as long as ostium, from which it is separated; ostial canal present; curvature of inner surface moderate. One specimen with 12 precaudal vertebrae and the rest (including the holotype) with 13.

Body of holotype mottled brown; indistinct brown band from snout through eye to gill cover; no distinct ocelli. Recently caught specimens tend to have brown pigmentation concentrated on dorsal part of body, especially along lateral line and at base of anterior part of dorsal fin; outer part of unpaired fins greyish; often longitudinal lines of brown pigmentation from tip of snout between eyes, below eyes and on ventral part of head.

REMARKS: In 17 specimens the ventral-fin rays are thicker and rather flat distally; this is neither a sexual dimorphism nor a size-dependent difference. Kotthaus ( 1979 ) named his new species trifilis, as he found that the left ventral-fin of his single specimen ends in 3 thread-like rays (Fig. 12). However, this specific name is misleading, as examination of an additional 69 specimens shows that each ventral-fin comprises only 2 unbranched rays. Examination of gut contents showed a great variety of food items, such as foraminiferans, dorsal felt from polychaets, gastropods (Cyclostrematidae, Naticidae, Trochidae, Buccinidae), bivalves (Mytilidae), pteropods, isopods, copepods, anomuran crabs, and fish-remains (scales, otoliths and bones). The sex distribution of 60 specimens was 39 females, 17 males and 4 indeterminable. The most ripe female (about 150 mm SL) had eggs 0.5 mm in diameter.

COMPARISONS: The closest related species seem to be N. analis and N. vityazi, with which trifilis shares a number of meristic characters, the presence of 2 spines on the preopercle and the absence of ocelli on the fins and vertical bars on the body. N. trifilis differs by having needle-like teeth (the 2 other species have granular teeth) and by having much longer gill filaments. It also differs from N. analis by having 13 (rarely 12) precaudal vertebrae vs 14 (rarely 13) and by not having the black band on the anal fin.

DISTRIBUTION: N. trifilis is known only from south and southeast of Socotra in 175-420 m (Fig. 1).

MATERIAL: Holotype: ZMH 5620, F, 157 mm; off Socotra (11 degrees 33.9'N, 52 degrees 54'E); R/V METEOR st. 102, Agassiz trawl; 175-337 m; 20 Dec 1964. ZM MGU P-18900 (26:17 F, 5 M and 47, 125-190 mm) and ZMUC 1377825-828 (4:2 M, 27, 117-181 mm); off Socotra (10 degrees 18.7'N, 56 degrees 7.3'E); R/V VITYAZ cr. 17, st.2573, bottom trawl; 408 m; 30 Oct 1988. ZM MGU P-18901 (2: F and ?, SL 145 mm); off Socotra (11 degrees 35.3'N, 52 degrees 39.6'E); R/V DMITRY STEFANOV cr. 3, trawl 42, bottom trawl; 400 m; 28 Dec 1988. ZM MGU P-18902, F, 175 mm; off Socotra (12 degrees 21.6'N, 53 degrees 16.5'E); R/V DMITRY STEFANOV cr. 3, trawl 50, bottom trawl; 370 m; 2 Jan 1989. ZM MGU P-18903 (22:12 F and 10 M, 134-188 mm), USNM 308998 (10:143-176 mm) and ZMUC P77829-832 (4:3 F and 1 M, 143-173 mm); off Socotra (10 degrees 19.5 'N,59 degrees 8.8'E); R/V VITYAZ cr. 17, st.2825, bottom trawl; 395-420 m; 14 Jan 1989.

Neobythites vityazi n. sp. (Fig. 13)

    Figure 13. Neobythites vityazi: a) holotype, F, 170 mm, ZM MGU P-18918. Vomer tooth patches: b) paratype, 107 mm MNHN 1992-526; c) holotype; d) paratype, M, 201 mm, ZM MGU P-18919. Left sagittal paratype, M, 182 mm, MNHM 1992-530: e) median view, f) anterior view. Scale bars 1 mm.

Neobythites analis (non Barnard): Shcherbachev 1980: 159, fig. 15c.

HOLOTYPE: ZM MGU P-18918, F, 170 mm; off SE Madagascar (22 degrees 19.1'S, 43 degrees 6.1'E); R/V VITYAZ cr 17, st. 2644, bottom trawl; 330-335 m; 2 Dec 1988.

PARATYPES: MNHN 1992-526 (2; 107-124 mm); off N. Madagascar (12 degrees 52.3'S, 48 degrees 10.4'E); A. Crosnier, coll., R/V VAUBAN, P.1, trawl 3; 403-415 m; 4 Mar 1971. MNHN 1992-527, 150 mm; off W. Madagascar (18 degrees 54'S, 43 degrees 55'E); A. Crosnier, coll., R/V VAUBAN, trawl 88; 280-310 m; 24 Nov 1973. MNHN 1992-528 (2; 149-152 mm); off SW Madagascar (23 degrees 36'S, 43 degrees 31.6'E); A. Crosnier, coll., R/V VAUBAN, trawl 56; 395-410 m; 26 Feb 1973. MNHN 1992-934, F, 153 mm; off SW Madagascar (23 degrees 35.9'S, 43 degrees 32.7'E); A. Crosnier, coll., R/V VAUBAN, trawl 57; 285-305 m; 27 Feb 1973. MNHN 1992-529, M, 170 mm; off SW Madagascar (23 degrees 26.2'S, 43 degrees 30.5'E); A. Crosnier, coll., R/V VAUBAN, trawl 58; 510 m; 27 Feb 1973. MNHN 1992-530, M, 182 mm; off SW Madagascar (23 degrees 35'S, 43 degrees 28.6'E); A. Crosnier, coll., R/V VAUBAN, trawl 65; 740-760 m; 28 Feb 1973. MNHN 1992-531, M, 154 mm; off SW Madagascar (23 degrees 36.4'S, 43 degrees 31'E); A. Crosnier, coll., R/V VAUBAN, trawl 66; 450-460 m; 28 Feb 1973. ZM MGU P-18919, M, 201 mm; off southern Mozambique (25 degrees 24.2'S, 34 degrees 17.2'E); R/V PROFESSOR MESIATZEV cr. 5, haul 77; 330 m; 31 Jan 1976. ZM MGU P-18920, M, 177 mm; off NE Madagascar (12 degrees 31'S, 48 degrees 8.5'E); R/V VITYAZ cr.17, st. 2608, bottom trawl; 690-695 m; 13 Nov 1988. ZM MGU P-18921 (4:3 F and 1 M?, 110-183 mm) and ZMUC P77833-835 (3:2 F and 1 M, 133-173 mm); same data as holotype. ZM MGU P18922, F, 113+?mm; off SW Madagascar (22 degrees 14.8'S, 43 degrees 07'E); R/V VITYAZ cr.17, st.2662, bottom trawl; 310-315 m; 4 Dec 1988.

DIAGNOSIS (based on 20 specimens, 107-201 mm SL; data for paratypes in parentheses): Dorsal-fin rays 103 (99-105); anal-fin rays 83 (81-87); 2 strong spines on preopercle; developed gill-rakers 9 (9-10); longest gill filaments 5.5 (4.4-6.9)% head length; pseudobranchial filaments 3-6; rear margin of vomer tooth patch convex (Fig. 13); teeth granular; precaudal vertebrae 13; diffuse, horizontal, brown lines on snout; no ocelli or vertical bars on body or fins.

DESCRIPTION (data for paratypes, if different from the holotype, are given in parentheses; additional meristic and morphometric data are given in Tables 1-7 and 14): Snout rather blunt, slightly shorter than or equal to diameter of eye; maxilla ends behind eye. Ventral-fin rays reach about halfway to anal fin. Rear edge of vomer tooth patch convex (Fig. 13). Pyloric caeca 12 (10-14). Gill-rakers iii, 1 + 8,v (i-iv,2-3 + 7-9,v-viii). Sagittal otolith (Fig. 13 e & f) more than twice as long as high, with smooth margin; ostium 2.5 times as long as cauda; colliculi partly separated; ostial canal indicated; curvature of inner surface strong.

Body mottled brown; lateral line distinctly pale. Peritoneum, oral and branchial cavities brown. Eye lens greenish. Indistinct brown lines on snout.

REMARKS: Gut contents from many specimens comprised large foraminiferans, unidentifiable crustacean remains, the cumacean Diastylis and gastropods Turridae, 4 mm, and Pyranudellidae, 7 mm). The size of the liver varies considerably, ending anterior to the stomach in some specimens, and in other specimens, ending near the anus. Fig. 13 shows that variation in shape of the vomer tooth patch is much less than in most other species; even a 107 mm SL specimen has the posterior margin convex.

COMPARISONS: Among the species with 2 spines on preopercle and lacking ocelli and dark bars on the body, N. vityazi seems most similar to N. analis, which also has a distinct lateral line, granular teeth and relative short gill filaments. In addition to the characters used in the key to species (p. 3), these two species differ in that N. vityazi usually has a deeper body (depth at anus 16-20.5% SL vs 14.5-17.5% SL in N. analis).

DISTRIBUTION: Known from the Mozambique Channel in depths of 280-760 m (Fig. 1).

ETYMOLOGY: Named after the R/V VITYAZ which has made a significant contribution to marine biology with her many valuable collections.

Table 1. Frequency distributions of dorsal and anal-fin ray counts for Western Indian Ocean species of Neobythites.

---------------------------------------------------------------------------
                                Number of dorsal-fin rays
                    88  89  90  91  92  93  94  95  96  97  98  99  100 
--------------------------------------------------------------------------
analis                                                           1   1
crosnieri                                    1           1   3    
malhaensis                                                       2
meteori                          1     
multistriatus        
natalensis                                                           1
somaliensis         
steatiticus          1       4   3   1   2              
stefanovi                2   5  12  16  11   4   3
trifilis                                                         1
vityazi                                                          1
---------------------------------------------------------------------------
                          Number of dorsal-fin rays
                 101  102  103  104  105  106  107  108  109  110  111
---------------------------------------------------------------------------
analis             2    3    5         3    3    2
crosnieri 
malhaensis                   2
meteori 
multistriatus                               1         1    1         1*
natalensis         2    1
somaliaensis       1#        1    1    3    4    4         1
steatiticus 
stefanovi 
trifilis          12   16   11    4    3
vityazi            4    3    2    1    1
---------------------------------------------------------------------------
                               Number of anal-fin rays 
                 72  73  74  75  76  77  78  79  80  81  82  83  84 
---------------------------------------------------------------------------
analis                                                            4    
crosnieri                                     2   1   1   1 
malhaensis                                1   1   1       1
meteori                     
multistriatus                   
natalensis                                                        1      
somaliaensis    
steatiticus       1   1   2   4   3 
stefanovi             3  13  13   5   3   2 
trifilis                                                      2   1   
vityazi                                                1   1  4   6    
---------------------------------------------------------------------------
                                 Number of anal-fin rays 
                   85   86   87   88   89   90   91   92   93   94   95
---------------------------------------------------------------------------
analis              5    3    5    1    1
crosnieri         
malhaensis    
meteori   
multistriatus                                     2         1         1* 
natalensis          2    1
somaliaensis                  2    4     6#  2    1
steatiticus    
stefanovi        
trifilis            8   12   16    7     6
vityazi             3    1    2
---------------------------------------------------------------------------
* specimen from Rodrigues
# 74mm SL specimen

Table 2. Frequency distributions of counts of pseudobranch filaments and developed gill-rakers for Western Indian Ocean species of Neobythites.

---------------------------------------------------------------------------
               Pseudobranch filaments         Developed gill-rakers
               ----------------------   -----------------------------------
                2  3  4  5  6  7  8      6  7  8  9  10  11  12  13  14  15

---------------------------------------------------------------------------
analis             2  8  8  5                     3  13   8 
crosnieri             4  1                     2  2   1 
kenyaensis      1  1                              1   1 
malhaensis         3  1                                       2   2 
meteori            1                     1 
multistriatus      1 *   3                                    1*  1   2   1
natalensis      1  1  2                        2  2 
somaliaensis          1# 6  6  1  1         1 *   3   9   1   1   1 
steatiticus     2  7  2                                   1   6   3   1 
stefanovi       1  6 14  8  2                         2  13  23   8 
trifilis           9 40  6                  9 43  7   2 
vityazi            1  6  6  6                    12   8
---------------------------------------------------------------------------

Table 3. Frequency distributions of counts of precaudal and total vertebrae for Western Indian Ocean species of Neobythites.

---------------------------------------------------------------------------
               Precaudal               Total number of vertebrae
              -----------     ---------------------------------------------
               12 13 14       52 53 54 55 56 57  58  59  60  61  62  63  64 
---------------------------------------------------------------------------
analis             5 19                           4   8   5   2
crosnieri          1  4                       2   3
kenyaensis         2                    1  1 (caudal tip missing)
malhaensis         4                           2  1   1 
meteori            1              1 
multistriatus*     5                                                  4 
natalensis         4                              2       2 
somaliaensis       3 12                                   1*  2   8   3   1 
steatiticus   10                  2  6  1  1 
stefanovi     50               1 13 19  7  2 
trifilis       2 59                               1  26  20   6 
vityazi          20                            1 12   2   4
---------------------------------------------------------------------------

Table 4. Frequency distributions of positions for dorsal-fin origin (with respect to vertebrae 2-6) and anal-fin origin (with respect to dorsal-fin rays 17-25) for Western Indian Ocean species of Neobythites.

---------------------------------------------------------------------------
                   Vertebrae            Dorsal-fin rays / Anal-fin origin
                 ---------------      -------------------------------------
                  2  3  4  5  6        17  18  19  20  21  22  23  24  25 
---------------------------------------------------------------------------
analis                    20  4                 1   7  11   3   2 
crosnieri                  4  1                 1   2   2 
kenyaensis              1  1                        1       1 
malhaensis       3   1                                          1   1   2
meteori                    1                        1 
multistriatus              4* 1             1*  1   1   2
natalensis              1  3                1       2   1
somaliaensis              10# 5                 1#  3   8   2 
steatiticus             5  5                1   3   4   2
stefanovi              37 12            1  10  14  16   5 
trifilis                3 58            2  13  23  16   7 
vityazi                   17  3                10   6   4
---------------------------------------------------------------------------
* includes specimen from Rodrigues
# 74 mm SL specimen

Table 5. Frequency distributions of head length (as % SL) for Western Indian Ocean species of Neobythites Size ranges (in mm of SL) are given in parentheses.

---------------------------------------------------------------------------
                                     Head length in % of SL
                       ----------------------------------------------------
                        19  20  21  22  23  24  25  26  27  28  29  30  31
--------------------------------------------------------------------------- 
analis(92-212)           1 3 4 3 6 2 2
crosnieri (113-192)              1 1 2 1 
malhaensis ( 117-135)            2 2
meteori (102)                            1 
multistriatus (76-178)         1*1   1   1 
natalensis (53-69)             1   2   1
somaliaensis (74-205)                1#  5 6 3
steatiticus(113-150)                           [1 2 1 1]^1)[3   1     2]^2)
stefanovi(29-198)                            3 11 5 5 8 9 3 1 1 1 
trifilis(117-190)                1 4 11 13 15 8        2 
vityazi (107-201)        1   3   2 4  1  4  4
---------------------------------------------------------------------------
^1)From Guff of Oman
^2) From Bay of Bengal

Table 6. Frequency distributions of body depth at anus (as % SL) for Western Indian Ocean species of Neobythites. Size ranges (in mm of SL) are given in parentheses.

---------------------------------------------------------------------------
                                  Depth at anus in % of SL
                       ----------------------------------------------------
                        14    15    16    17    18    19    20    21    22
---------------------------------------------------------------------------
analis (92-212)             1  5  4  3  1  3  1 
crosnieri (113-192)                     1        3  1
malhaensis (117-135)                       2     1     1
meteori (102)                                                1
multistriatus (76-178)   1*    1  1     1
natalensis (53-69)             2  1  1
somaliaensis (74-205)       1  1  5  2# 3  2     1
steatiticus (113-150)                         3  2  2  2  1
stefanovi (29-198)                3  2  3  2  4  5  5  3  3  4  3  5  2  3 
trifilis (117-190)                2  2  8 11 11  6  4  4 
vityazi (107-201)                    1     5     5  2  2  2  1  1
---------------------------------------------------------------------------

Table 7. Frequency distributions of gill filament lengths (as % HL) for Western Indian Ocean species of Neobythites. Size ranges (in mm of SL) are given in parentheses.

---------------------------------------------------------------------------
                       Length of longest gill filament in % of head length
                        3  4  5  6  7  8  9  10 11 12 13 14 15 16 17 18 19
---------------------------------------------------------------------------
analis(92-212)                 4 4 3 6 5
crosnieri (113-192)           2  2 1
kenyaensis (103+-124+)                 1  1
malhaensis (117-135)                   2  1 1
meteori (102)                                         1
multistriatus (76-178)           4*1
natalensis (53-69)      2 1  1
somaliaensis (74-205)                         1#1 2 6 3 1   1
steatiticus (113-150)                                          1  2 3 2   2 
stefanovi(29-198)                1 4 1      1 1 4 4 4 7 6 1 1 2 2 3 1   3 1 
    trifilis(117-190)                           1 4 9 51316 3 6 1 2 1 
vityazi (107-201)          4 3 4 5 3 1
---------------------------------------------------------------------------
* Rodrigues specimen
# 74 mm SL specimen

Table 8. Data for Neobythites analis: L = lectotype, PL = 6 paralectotypes, AS = additional specimens; n = number of additional specimens examined.

--------------------------------------------------------
                            L       PL       AS      n 
Standard length            136    92-134   92-212    25 

                                Meristic data 
                           -----------------------------
Pectoral fin rays           27     25-27    25-27    19 
Anal fin origin/vertebra    16     14-16    14-16    24

                             Morphometric data in % SL
                           -----------------------------
Head length                19.5    19-20    19-22    21 
Depth at anus              16.5    15-17    14-17    18 
Eye diameter                4.4   4.4-5.1  4.2-5.2   19 
Preanal distance           41.5    38-40    37-41    19 
Predorsal distance         21.5    21-22    21-25    19
---------------------------------------------------------

Table 9. Data for Neobythites crosnieri: H + P = holotype and 4 paratypes.

-----------------------------------------------------
                                   H           H+P 
Standard length                   192        113-192

                                    Meristic data
                                ---------------------
Pectoral fin rays                  26         25-27 
Anal fin origin/vertebra           15          15 
1st ocellus on dorsal rays       41-48        38-48 
2nd ocellus on dorsal rays       65-70        65-75

                             Morphometric data in % SL
                             --------------------------
Head length                      21.0         21-22
Depth at anus                    18.0         16-18
Eye diameter                      4.2          4-5 
Preanal distance                 41.5         39-43
Predorsal distance               25.0         25-26
Snout-1st ocellus                65           59-65
Snout-2nd ocellus                82           80-84
-------------------------------------------------------

Table 10. Data for Neobythites malhaensis: H = holotype, H + P = holotype and 3 paratypes.

-------------------------------------------------------
                                  H           H + P
Standard length                  123         117-135 

                                    Meristic data
                                -----------------------
Pectoral fin rays                 30           30 
Anal fin origin/vertebra          15           15 
Ocellus on dorsal rays           25-36        25-37

                            Morphometric data in % SL
                            -------------------------
Head length                       21          21-22 
Depth at anus                     17          17-19 
Eye diameter                       4.9         4-4 
Preanal distance                  36          36-40 
Predorsal distance                20          20-21 
Snout-ocellus                     42          42-45
------------------------------------------------------

Table 11. Data for Neobythites multistriatus and natalensis: H = holotype, H + P = holotype and 3 paratypes.

-------------------------------------------------------------------
                                multistriatus        natalensis
                           -----------------------  ---------------
                            H      H+P    ZM MGU      H     H+P 
                                          P-18914 
                          -----------------------------------------
Standard length            178   149+-178    76       53    53-69 

                                      Meristic data
                          -----------------------------------------
Pectoral fin rays         29/30   28-30      30     28/29   27-29 
Anal fin origin/vertebra    16    14-16      15       14    14-15

                                 Morphometric data in % SL
                          -----------------------------------------
Head length                22.0    21-23    20.5     21.5    20-22 
Depth at anus              16.5    15-16    14.0     15.0    15-16 
Eye diameter                4.3   4.3-4.4    4.3      4.7   4.6-4.7 
Preanal distance           41.5    40-43    35.5     37.0    37-40 
Predorsal distance         24.0    24-25    23.0     23.5    23-24
--------------------------------------------------------------------

Table 12. Data for N. somaliaensis: H = holotype, H + P = H and 13 paratypes.

-----------------------------------------------------
                            H         H+P     ZM MGU 
                                             P-18913
                           --------------------------
Standard length            205      123-205     74

                           Meristic data
                           --------------------------
Pectoral fin rays           28       28-29      28 
Anal fin origin/vertebra    16       15-16      15

                           Morphometric data in % SL
                           --------------------------
Head length                 23.5     23-24     22.0 
Depth at anus               18.0     14-18     16.0 
Eye diameter                 5.1    5.0-6.1     5.1 
Preanal distance            43.5     39-44     39.0 
Predorsal distance          28.0     24-28     25.0
-----------------------------------------------------

Table 13. Data for Neobythites steatiticus: H = holotype, AS = holotype and n additional specimens.

-----------------------------------------------------
                               H        AS        n    
Standard length               129     113-150    11                 

                              Meristic characters
                              ------------------------
Pectoral fin                   24      24-27      9 
Anal fin origin/vertebra.       -      14-15     10 
Ocellus on dorsal rays          -      24-37     10

                             Morphometric data in % SL  
                             -------------------------
Head length                   28.0     25-31     11    
Depth at anus                  -       17-19     10 
Eye diameter                   -      4.8-5.9     9 
Preanal distance              40.0     40-46     10 
Predorsal distance            24.5     24-28     10 
Snout-ocellus                  -       46-51     10
-------------------------------------------------------

Table 14. Data for N. trifilis (holotype + 31 - 60 specimens) and N. vityazi: H = holotype; H + P = ranges for holotype and 18 - 19 paratypes; number of specimens in brackets.

-------------------------------------------------------------
                        trifilis           vityazi 
                     --------------   -----------------------
                                       H       H+P
                                      ---------------
Standard length      117-190 [61]     170    107-201    [20]

                              Meristic data
                     --------------------------------
Pectoral fin rays     27-30  [32]      26     25-28     [20] 
Anal fin origin/ 
vertebra              14-16  [61]      15     14-16     [20]

                       Morphometric data in % SL
                     --------------------------------
Head length           21-24  [54]     23.0    20-23     [19] 
Depth at anus         15-19  [48]     19.5    16-20     [19] 
Eye diameter         5.1-6.4 [48]      5.3   4.3-5.3    [19] 
Preanal distance       3543  [40]     41.0    38-44     [19] 
Predorsal distance    23-28  [35]     25.5    23-26     [19]
-------------------------------------------------------------
ACKNOWLEDGEMENTS

I wish to thank the following colleagues for making material available and/or for providing information: M.E. Anderson (RUSI), M.L. Bauchot (MNHN), G. Duhamel (MNHN), P.C. Heemstra (RUSI), P.A. Hulley (SAM), S. Jewett (USNM), W. Klausewitz (SMF), N.R. Merrett (BMNH), N.V. Parin (IOAN), J.-C. Quero (lFREMER, La Rochelle), Yu. Sazonov (ZM MGU), W. Schwarzhans (Mulheim, Germany), B. Seret (MNHN), Yu. Shcherbachev (IOAN), P.K. Talwar (ZSI), F. Uiblein (SMF), H. Wilkens (ZMH). J. Knudsen (ZMUC) kindly identified stomach contents.

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LEVITON, A.E.R.H. GIBBS, Jr., E. HEAL & C.E. DAWSON. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: 802-832.

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-- & J.-C.QUERO. 1991. Quelques Ophidiiformes de l'ile de la Reunion: description d'une espece nouvelle. Cybium 15: 193-198.

-- & F. UIBLEIN. 1993. Tiefenwasser- und Tiefsecfische aus dem Roten Meer. XVI. A new species of Neobythites from the NW Indian Ocean and the Red Sea. Senckenbergiana maritima 23: 109-113.

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SCHWARZHANS, W. 1994. Sexual and ontogenetic dimorphism in otoliths of the family Ophidiidae. Cybium, 18(1): 71-98.

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Copyright 1995 J.L.B. Smith Institute of Ichthyology


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