PROCEEDINGS OF THE , Biological Laboratory I LtBRARY \ 1 7 1992 CALIFORNIA ACADEMY OF SCIENCES Woods r Vol. 48, No. 2, pp. 27-95, 37 figs., 7 tables. GRENADIERS (PISCES, GADIFORMES) OF THE NAZCA AND SALA Y GOMEZ RIDGES, SOUTHEASTERN PACIFIC By Yuri I. Sazonov Department of Ichthyology, Zoological Museum, Moscow Lomonosov State University, Herzen Street 6, Moscow 103009, Russia and Tomio Iwamoto Department of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. Abstract: Twenty-five species of grenadiers are recorded from the Nazca and Sala y Gomez ridges in the southeastern Pacific. New species include: Caelorinchus immaculatus, C. multifasciatus, C. nazcaensis, C. spilonotus, Hymenocephalus neglectissimus, H. semipellucidus, Kuronezumia pallida, Ventrifossa macrodon, V. teres, and V. obtusirostris. Caelorinchus immaculatus is very similar to C. karrerae from the southeastern Atlantic and Indian Ocean. The H. striatissimus complex is examined using new data. Hymenocephalus semipellucidus and H. neglectissimus appear to be part of this complex. The subspecies H. s. hachijoensis from Japan is elevated to full species status. Kuronezumia, formerly considered a subgenus of Nezumia, is redefined and elevated to generic status to include K. pallida, K. bubonis, K. leonis, K. macronema, K. dara, and two undescribed species. Despite proximity of the ridges to the mainland coast of Peru, relationships of the associated fauna are to the west, particularly the western Pacific and Hawaiian Islands. Of the 25 species from these ridges, eight are definitely known from the vicinity of the Hawaiian Islands: Caelorinchus spilonotus, Cetonurus crassiceps, * Coryphaenoides paradoxus, * H. striatulus, Malacocephalus laevis, * Mataeocephalus acipenserinus, Nezumia propinqua, * and Pseudocetonurus septifer. Three other species whose identifications are undetermined may be part of, or have close counterparts in, the Hawaiian fauna: Gadomus sp. cf. melan- opterus, Hymenocephalus sp. cf. aterrimus, and Trachonurus villosus*"! The four species marked with an asterisk are broadly distributed through the Pacific, Indian, and Atlantic oceans. Malacocephalus laevis is known from the continental slopes of southern California and on seamounts off Baja California, but nowhere else along the Pacific coast of Central and South America. Caelorinchus immaculatus is also recorded from mainland South America; Nezumia convergens is questionably represented by a specimen from the Sala y Gomez Ridge. Received May 1, 1991. Accepted December 1, 1991. Contents Abstract 27 Introduction 2 8 Materials and Methods 28 Classification of Grenadiers 30 Description of Nazca and Sala y Gomez Ridges 32 Key to NSG Species of Grenadiers 32 Bathygadidae 3 6 1. Gadomus sp. cf. melanopterus Gil- bert, 1 905 36 [27] 28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 M ACROURIDAE 3 8 Subfamily Macrouroidinae 38 2. Macrouroides inflaticeps (Smith and Radcliffe, 1912) 3 8 3. Squalogadus modificatus Gilbert and Hubbs, 1 9 1 6 3 9 Subfamily Macrourinae 3 9 4. Cetonurus crassiceps (Gunther, 1 878) 40 5. Caelorinchus immaculatus n. sp 41 6. Caelorinchus multifasciatus n. sp 44 7. Caelorinchus nazcaensis n. sp 46 8. Caelorinchus spilonotus n. sp 49 9. Coryphaenoides paradoxus (Smith and Radcliffe, 1 9 1 2) 5 2 10. Hymenocephalus sp. cf. aterrimus Gilbert, 1 905 54 1 1 . Hymenocephalus gracilis Gilbert and Hubbs, 1 920 55 12. Hymenocephalus neglectissimus n. sp 56 13. Hymenocephalus semipellucidus n. sp 60 14. Hymenocephalus striatulus Gilbert, 1905 63 15. Kuronezumia pallida n. sp 65 16. Malacocephalus laevis (Lowe, 1843) 68 17. Mataeocephalus acipenserinus (Gil- bert and Cramer, 1897) 70 18. Nezumia convergens (Garman, 1 899) 7 1 19. Nezumia propinqua (Gilbert and Cramer, 1897) 72 20. Pseudocetonurus septifer Sazonov and Shcherbachev, 1 982 75 21. Trachonurus villosus (Gunther, 1877) 77 22. Ventrifossa johnboborum Iwamoto, 1982 78 23. Ventrifossa macrodon n. sp 81 24. Ventrifossa obtusirostris n. sp 84 25. Ventrifossa teres n. sp 86 blogeographical considerations 88 Acknowledgments 9 1 Resumen 91 Literature Cited 9 2 cruise between April and November 1979. Pre- liminary studies (Parin et al. 1981) of the ma- terial collected during that cruise suggested the existence of an extremely diverse ichthyofauna on these ridges. The species composition of the macrourid fishes, or grenadiers, differed mark- edly from that known along the nearby conti- nental margins and, in fact, appeared to be more similar to those from the Indo-Pacific region, not the eastern Pacific. Subsequently, between 1979 and 1987, several other Soviet expeditions investigated the ridges and collected many grenadiers from depths of about 300-800 m, with a few trawls below 1 ,000 m. In preparing the section on the grenadiers for the preliminary report (Parin et al. 1981), Sa- zonov recognized 1 1 species, eight of which he considered new or of uncertain taxonomic status. Additionally, Sazonov and Shcherbachev (1982) described a new genus and species (Pseudoceto- nurus septifer) from the Sala y Gomez Ridge. Because of the obvious Indo-West Pacific af- finities of the grenadiers, it was necessary to ex- amine many type specimens deposited in U.S. museums, particularly the U.S. National Mu- seum of Natural History (USNM) and the Cal- ifornia Academy of Sciences (CAS). The idea of studying the fauna on a cooperative basis was initially proposed by N. V. Parin during a Work- shop on Cold-Water Fishes held in Orono, Maine, in 1979. The study was begun in 1981 and a preliminary manuscript was drafted in 1 988 after Iwamoto spent 3.5 months in the Laboratory of Oceanic Ichthyofauna, Russian Academy of Sci- ences, Moscow. This paper presents the results of the cooperative effort. Twenty-five species of grenadiers are herewith recorded from the Nazca and Sala y Gomez ridg- es, ten of which are described as new. Two others may represent undescribed taxa, but the avail- able material does not justify their formal rec- ognition. Considering that for the most part only the shallowest levels of these oceanic ridges have been sampled, the faunal list can be expected to grow, especially in bathybenthic and abyssal forms. Introduction The research vessel Ichthyandr of the Sevas- topol Bureau of Underwater Researches of the former USSR Ministry of Fisheries surveyed the Nazca and Sala y Gomez oceanic ridges (Fig. 1) off the southwestern coast of Peru on its fifth Materials and Methods Almost all specimens reported here were col- lected by Soviet oceanographic and fishery re- search vessels. Most collections were made be- tween 1979 and 1987, although a few specimens found at the Zoological Institute in St. Petersburg SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 29 Figure 1 . Map of southeastern Pacific showing location of the Nazca and Sala y Gomez ridges. Inset shows general location of larger map. Depth contours in fathoms (solid lines represent 2,000 fathoms or about 3,658 m; broken lines represent 1,000 fathoms or about 1,829 m). Based in part on a chart by Mammerickx et al. 1975. 30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 (ZIN) and the Zoological Museum, Moscow State University (ZMMGU) were procured in 1973 and 1975 during cruises of the Hercules and As- tronomer of the Pacific Fisheries and Oceanog- raphy Research Institute, Vladivostok (TINRO). Most of the specimens used for this report are deposited in the Institute of Oceanology, Russian Academy of Sciences, Moscow (IOAN) and ZMMGU, with representatives also deposited in CAS and ZIN. A list of Russian vessels and sta- tion data are provided in Table 1 . Institutional abbreviations follow Leviton et al. (1985) as amended by Leviton and Gibbs (1988). Abbreviations and methods of taking measurements and counts are the same as those described in a previous paper (Iwamoto and Sa- zonov 1988). The abbreviation NSG is used throughout to refer to the combined Nazca and Sala y Gomez ridges. The following abbrevia- tions are also used: cr. — cruise; fm(s) — fathom(s); HL— head length; spec — specimen; sta.— sta- tion; TL— total length; tr.— trawl. The descriptions of species are based mainly on the materials examined from NSG and usu- ally do not include specimens from outside the area, even though such additional materials have been examined when available. These peripheral specimens are usually considered in the Re- marks and Comparisons sections. Several species have been adequately de- scribed and illustrated in recent literature. For these, a brief diagnosis or a short diagnostic de- scription is provided. For other species, more extensive descriptions are given, but synonymies are kept to a minimum; those included are lim- ited to recent references and a few older ones with good descriptions or illustrations. As this is not meant to be a comprehensive revision of the groups treated, generic accounts are for the most part very brief, and synonyms are omitted. Classification of Grenadiers There has been a recent surge of interest in the higher classification of gadiform fishes, which culminated in a workshop on the systematics of the group in 1986 (see Cohen 1989). The work- shop brought together disparate workers in the group to air their views on its classification and phylogeny and search for a modicum of agree- ment. For the so-called macrouroid fishes, or grenadiers, there was none. Howes (1988, 1989), in a radical departure, removed the bathygadine Table 1 . Soviet research vessels and trawl stations on the Nazca and Sala y Gomez ridges at which grenadiers were col- lected. Hercules (1973) tr. 40: Nazca Ridge, 1 9°30.4'S, 80°1 1 .2'W; 950 m; bot. trawl; 20.X.1973. Hercules (1975) sta. 68: Sala y Gomez Ridge, 25°18.8'S, 93°34.2'W; 610 m; bot. trawl; 1. XI. 1975. sta. 70: Sala y Gomez Ridge, 25°01'S, 91°18'W; 610-620 m; bot. trawl; 2.XI. 1975. sta. 74: Sala y Gomez Ridge, 24°57.5'S, 88°27.1'W; 550-630 m;bot. trawl; 5.XI.1975. sta. 1 10: between Nazca and Sala y Gomez ridges, 25°35.0'S, 85°13'W; 1,600-2,000 m; midwater trawl; 3.XII.1975. Astronomer (1975) tr. without no.: Nazca Ridge, 23°30.5'S, 81°45'W; 300-330 m; bot. trawl; 29.VI.1975. sta. 104: Sala y Gomez Ridge, 25°00'S, 88°30'W; 550 m; bot. trawl; 24.VII.1975. tr. without no.: Sala y Gomez Ridge, 25°02'S, 88°35'W; 550 m; bot. trawl; 24.VII.1975. Ichthyandr cr. 5 (1979) tr. 1: Nazca Ridge, 19°40'S, 80°18'W; 980 m; 26.VIII.1979. tr. 13: Nazca Ridge, 21°30 , S, 81°42'W; 330 m; 5.IX.1979. tr. 1 4: Nazca Ridge, 2 1°24'S, 8 1 °4 1 ' W; 330-340 m; 6.IX. 1 979. tr. 15: Nazca Ridge, 21°27'S, 81°41'W; 330 m; 7.IX.1979. tr. 17:NazcaRidge,21°41'S,81°41'W;310-330m;9.IX.1979. tr. 40: Sala y Gomez Ridge, 25°52'S, 86°47'W; 440 m; 24.IX.1979. tr. 44: Nazca Ridge, 21°32'S, 81°38'W; 330 m; 30.IX.1979. tr. 50: Sala y Gomez Ridge, 25°07'S, 99°37'W; 450-480 m; 24.X. 1979. tr. 51: Sala y Gomez Ridge, 25°07'S, 99°39'W; 360-400 m; 24.X.1979. tr. 52: Sala y Gomez Ridge, 25°02'S, 99°26'W; 770 m; 24.X. 1979. tr. 53: Sala y Gomez Ridge, 25°02'S, 88°32'W; 540 m; 29.X.1979. tr. 54: Sala y Gomez Ridge, 25°01'S, 88°27'W; 535-575 m; 30.X.1979. tr. 55: Sala y Gomez Ridge, 25°00'S, 88°31'W; 345-540 m; 30.X.1979. tr. 56: Sala y Gomez Ridge, 25°46'S, 86°33'W; 410 m; 31.X.1979. tr. 57: Sala y Gomez Ridge, 25°42'S, 86°32'W; 420 m; 31.X.1979. Ichthyandr cr. 6(1980) tr. 1: Nazca Ridge, 21°27'S, 81°40'W; 340 m; 5.VIII.1980. tr. 2: Nazca Ridge, 21°08'S, 81°39'W; 320 m; 5.VIII.1980. tr. 3: Nazca Ridge, 21°28'S, 81°39'W; 340 m; 6.VIII.1980. tr. 10: Nazca Ridge, 21°27'S, 81 41'W; 320-340 m; 9.VIII.1980. tr. 33: Nazca Ridge, 21°28'S, 81°4LW; 330 m; 21. IX. 1980. tr. 56: Sala y Gomez Ridge, 25°45'S, 86°35'W; 430 m; 2.X.1980. Odissey cr. 2(1981-82) tr. 11: Nazca Ridge, 22°11'S, 81°21'W; 235 m; bot. trawl; 23.XII.1981. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 31 Table 1. Continued. Table 1. Continued. tr. 14: Nazca Ridge, 22°1 1.6'S, 81°18.8'W; 230 m; bot. trawl; 25.XII.1981. tr. 15: Nazca Ridge, 22°12'S, 81°19'W; 230-250 m; bot. trawl; 25.XII.1981. Akademik Kurchatov cr. 34 (1983) sta. 3594: Nazca Ridge, 21°28.8'S, 81°42.0'W; 340 m; Sigs- bee trawl; 25.XI. 1982. Professor Mesiatzev cr. 13 (1983) tr. 1: Sala y Gomez Ridge, 24°58.6'S, 88°28'W; 565-555 m; 1.IX.1983. tr. 2: Sala y Gomez Ridge, 24°59.8'S, 88°25'W; 550-560 m; 1.IX.1983. tr. 4: Sala y Gomez Ridge, 25°45'S, 86°37'W; 410-420 m; 2.IX.1983. tr. 7: Sala y Gomez Ridge, 25°41'S, 85 31'W; 285-300 m; 3.IX.1983. tr. 10: Sala y Gomez Ridge, 25°34.4'S, 85°20.7'W; 1.070- 1,100 m; 4.IX.1983. tr. 14: Sala y Gomez Ridge, 25°52.7'S, 84°34.2'W; 1,050 m; 5.IX.1983. tr. 17: between Sala y Gomez and Nazca ridges, 23°38.8'S, 85 27.5'W; 1,220-1,230 m; 7.IX.1983. tr. 23: Nazca Ridge, 22°08'S, 81°19'W; 235-250 m; 12.IX.1983. tr. 25: Nazca Ridge, 22°04'S, 81°17'W; 235-225 m; 12.IX.1983. tr. 31: Nazca Ridge, 22°06'S, 81°17'W; 225-240 m; 14.IX.1983. tr. 35: Nazca Ridge, 22°07'S, 81°18'W; 340-325 m; 14.IX.1983. tr. 36: Nazca Ridge, 22°07'S, 81°18'W; 235-230 m; 14.IX.1983. tr. 39: Nazca Ridge, 22°06'S, 81°16'W; 230-240 m: 16.IX.1983. tr. 43: Nazca Ridge, 20°46'S, 80°52'W; 320-325 m; 18.IX.1983. tr. 44: Nazca Ridge, 20°44.9'S, 80°52.3'W; 340-780 m; 18.IX.1983. tr. 94: Nazca Ridge, 19°35.5'S, 80°15.7'W; 940-960 m; 10.XI.1983. tr. 109: Nazca Ridge, 22°06'S, 81°17'W; 225-240 m; 15.XI.1983. tr. 120: Nazca Ridge, 22°05'S, 81°17'W; 230 m; 18.XI.1983. Professor Mesiatzev cr. 15 (1984) tr. 49: Sala y Gomez Ridge, 25°01.7'S, 88°26.9'W; 525-530 m; 10.X.1984. tr. 50: Sala y Gomez Ridge, 25°57'S, 88°31'W; 565 m; 11.X.1984. tr. 52: Sala y Gomez Ridge, 25°45'S, 88°36'W; 400 m; 11.X.1984. tr. 53: Sala y Gomez Ridge, 25°44'S, 86°36'W; 390-385 m; 11.X.1984. tr. 54: Sala y Gomez Ridge, 25°48'S, 86°15'W; 304 m; 12.X.1984. Professor Shtokman cr. 18 (1987) sta. 1826: Nazca Ridge, 20°44.8'S, 80°53.7'W; 300-0 m over bot. depth 330-340, IKMWT; 17.IV. 1987. sta. 1828/35: Nazca Ridge, 20°48.8'S, 80°53.5'W; 730-720 m; traps; 18.IV. 1987. sta. 1845: Nazca Ridge, 21°24.0'S, 81°38.6'W; 330 m; bot. trawl; 19.IV.1987. sta. 1851: Nazca Ridge, 21°23.2'S, 81°38.3'W; 330-350 m; bot. trawl; 19-20.IV. 1987. sta. 1856: Nazca Ridge, 21°43.9'S, 81°04.9'W; 900-920 m over bot. depth 1,270-1,200 m; IKMWT; 20.IV. 1987. sta. 1864/76: Nazca Ridge, 22°04.1'S, 81°16.7'W; 530-500 m; traps; 21. IV. 1987. sta. 1867: Nazca Ridge. 22°06.1'S, 81°17.0'W; 225-247 m; bot. trawl; 21. IV. 1987. sta. 1873: Nazca Ridge, 22°07.1'S, 81°16'W; 235 m; bot. trawl; 22.IV. 1987. sta. 1879/1882: Nazca Ridge, 23°26.4'S, 83°20.1'W; 505 m; traps; 23.IV. 1987. sta. 1887: between Sala y Gomez and Nazca ridges, 24°41'S, 85°25.6'W; 50-300 m over bot. depth 408->2,000 m; IKMWT; 24.IV.1987. sta. 1925: Sala y Gomez Ridge, 25°32.4'S, 85°29.5'W; 50- 300 m over bot. depth l,200-> 2,000 m; IKMWT; 27.IV.1987. sta. 1938: Sala y Gomez Ridge, 25°42.4'S, 86°35.3'W; 380 m; Sigsbee trawl; 28.IV. 1987. sta. 1940: Sala y Gomez Ridge, 25°41.0'S, 86°35.9'W; 380 m; bot. trawl; 28. IV. 1987. sta. 1941: Sala y Gomez Ridge, 25°48.6'S, 86°34.1'W; 410- 385 m; 28.IV.1987. sta. 1949: Sala y Gomez Ridge, 25°37.5'S, 86°29.4'W; 200- m over bot. depth 390 m; IKMWT; 29.IV. 1987. sta. 1955/62: Sala y Gomez Ridge, 24°57.7'S, 88°36.1'W; 560 m; traps; 29-30.IV. 1987. sta. 1956/61: Sala y Gomez Ridge, 24°54.0'S, 88°32.0'W; 580 m: traps; 29-30.IV. 1987. sta. 1964: Sala y Gomez Ridge, 24°56.3'S, 88°32.6'W; 580- 564 m: bot. trawl; 30.IV. 1987. sta. 1965: Sala y Gomez Ridge, 24°58.5'S. 88°29.3'W; 562- 545 m; bot. trawl; 30.IV. 1987. sta. 1970: Sala y Gomez Ridge, 25°34.2'S, 89°09.1'W; 540- 560 m; Sigsbee trawl; 1.V.1987. sta. 1971: Sala y Gomez Ridge, 25°30.4'S, 89°10.3'W; 570- 580 m; bot. trawl; 1.V.1987. sta. 1976: Sala y Gomez Ridge, 25°33.6'S, 89°1 1.9'W; 563- 590 m; 1.V.1987. sta. 1977: Sala y Gomez Ridge, 25°09.9'S, 90° 18. 7' W; 545- 600 m; bot. trawl; 1-2.V.1987. sta. 1996: Sala y Gomez Ridge, 25°08.2'S, 99°25'W; 750- 800 m, 5.V.1987. sta. 2018: Sala y Gomez Ridge, 25°07.9'S, 99°26.8'W; 730- 790 m; 7.V.1987. sta. 2019: Sala y Gomez Ridge, 25°05.7'S, 99°27.7'W; 750 m; Sigsbee trawl; 7.V.1987. sta. 2023: Sala y Gomez Ridge, 25°07'S, 99°40'W; 439-500 m;bot. trawl; 8.V.1987. and trachyrincine grenadiers from the suborder Macrouroidei and included them in the suborder Gadoidei, raising both to family status (see also Howes and Crimmen 1990). Okamura (1989) 32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 maintained the original view of the Macrouroidi- dae as a distinct family in the Macrouroidei and also raised the Trachyrincinae to family status, but he kept them and the Bathygadinae in the Macrouroidei, and also added to this suborder the enigmatic Euclichthyidae. Iwamoto (1989), Markle (1989), and Nolf and Steurbaut (1989a, b) maintained more traditional views of Tra- chyrincinae, Macrouroidinae, Bathygadinae, and Macrourinae as subfamilies of Macrouridae. We recognize that there are apt to be changes in the higher classification of what we now con- sider as macrouroid fishes, and that the tradi- tional view of the family Macrouridae being monophyletic will probably not stand. There will undoubtedly be a sorting-out period before a consensus is reached based on additional re- search. For the purposes of this paper, we find it convenient to follow a conservative classification and treat the major groups as subfamilies, with the exception of the Bathygadidae, and call them all grenadiers. Description of Nazca and Sala y Gomez Ridges One of the most notable geologic features of the oceanic basin of the southeastern Pacific is a long, deep, and broken submarine range that ex- tends westward from the Peru-Chile Trench in the vicinity of Nazca, Peru, and traverses the basin to connect with the East Pacific Rise around longitudes 1 10°-1 15°W (Fig. 1). It is one of the world's major mountain ranges, but it is exposed at the surface only on San Ambrosio, San Felix, Sala y Gomez, and Easter islands (Norris 1961). The Nazca Ridge forms the easternmost part of this range, beginning at the western rim of the Peru-Chile Trench at a latitude of about 1 5°S and extending in a southwesterly direction for about 600 nautical miles. The northeastern half of the ridge arises from a bottom depth of more than 4,500 m to reach upwards to within 2,200- 2,900 m of the surface. Only a few pinnacles in this part of the ridge are taller. This contrasts with the higher southeastern half, where a con- siderable portion lies above the 2,000-m isobath. Dominating this half is an elongated promi- nence, offset from the main axis of the ridge, that rises to within 320 m of the sea surface. The central part of the range lies to the east of Sala y Gomez Island and forms a well-defined ridge that rises 2,400-3,000 m above the sur- rounding sea floor (Norris 1961). This ridge, called the Sala y Gomez Ridge (Fisher and Norris 1960), extends more than 1,000 nautical miles to the east. It is punctuated by many promi- nences that rise to within 1,500 m of the surface. Several peaks that lie between longitudes 97°W and 101°W come to within the upper 500 m of the surface. Between the Nazca and Sala y Gomez ridges and to the south of the Nazca Ridge are a cluster of high seamounts that rise close to the surface. San Felix and San Ambrosio islands constitute the tips of the easternmost peaks of this cluster. The seamounts in this region were extensively investigated by Russian vessels. Key to NSG Species of Grenadiers This key is presented as a convenient guide to the identification of grenadiers from the Nazca and Sala y Gomez ridges. It should be used only as a guide, however, and preliminary identifi- cations should be confirmed with the descrip- tions. It can be confidently used only for adult specimens, as juveniles undergo ontogenetic changes that render many of the key characters useless. Because its coverage is geographically limited, some character states that key out NSG genera will not agree with the broader range of states given in the diagnoses for genera (e.g., pel- vic rays in Caelorinchus is given as seven in the key but six or seven in the diagnosis because six is found only in one western Pacific species). Fig- ures for the key are mainly from FAO Species Identification Sheets. For a more comprehensive key to the genera and species of grenadiers, the reader is referred to the FAO Species Catalogue on Gadiformes of the world (Cohen et al. 1990). 1 a. One dorsal fin, no portion elevated, spi- nous rays absent; pelvics small or absent; head enormous, globose; eyes 1 or more in HL (Macrouroidinae) (Fig. 2a) 2 lb. Two dorsal fins, the first elevated, first two rays spinous; pelvics small to large; head moderate to large; eyes 5 or less in HL (Fig. 2b) 3 2a. Pelvic fins absent Macrouroides inflaticeps (p. 38) 2b. Small pelvics with 5 or 6 rays Squalogadus modificatus (p. 39) 3a. Mouth large, terminal (Fig. 2c); first (out- ermost) gill slit unrestricted; gill rakers SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 33 Figure 2. (a) Squalogadus, (b) Nezumia, (c) Gadomus. on first arch long and numerous (Fig. 3a); second dorsal begins close behind first dorsal, its rays longer than those of anal; scales without spinules (Bathygadidae) Gadomus sp. cf. melanopterus (p. 36) 3b. Mouth small to large, usually subinferior (Fig. 2b); first gill slit restricted by mem- branes connecting upper and lower por- tions of first arch to gill cover; gill rakers tubercular or tablike (Fig. 3b); second dorsal separated by a distinct gap from first dorsal, its rays usually shorter than those of anal; scales beset with spinules in all but a few species (Macrourinae) 4 Branchiostegal rays 6 5 Branchiostegal rays 7 9 Snout blunt, scarcely protruding; ridge of stout scales, if present, not continuous to preopercle, and ends in a blunt or rounded tip (Fig. 4a); rakers present on both sides of first arch; pelvic rays 9-1 1 (rarely 8) Coryphaenoides paradoxus (p. 52) Snout sharply and distinctly pointed; a continuous ridge of stout scales from tip of snout to angle of preopercle, the ridge ending in a sharp point (Fig. 4b); rakers absent on outer side of first gill arch; pel- vic rays 7 {Caelorinchus) 6 Light organ long, the anterior end just 4a. 4b. 5a. 5b Figure 3. Outer gill arches of (a) Gadomus and (b) Ne- zumia. behind isthmus (Fig. 5a); prominent body markings present 7 6b. Light organ short, the anterior end not extending to a line between pelvic fin origins (Fig. 5b); no body markings 8 7a. A series of about 7 saddlemarks on body; first dorsal not black tipped Caelorinchus multifasciatus (p. 44) 6a. Figure 4. Diagrammatic illustrations of (a) Coryphae- noides paradoxus and (b) Caelorinchus comparing snout shape and suborbital ridge. Figure (a) redrawn from Iwamoto and Sazonov(1988, fig. 28). 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 25 mm Figure 5. Diagrammatic ventral views of Caelorinchus showing (a) long light organ extending from anus forward to large fossa on chest and (b) short light organ, scarcely apparent, anterior to anus. 7b. Not more than 2 or 3 saddlelike marks on body; first dorsal fin black tipped Caelorinchus spilonotus (p. 49) 8a. Snout rather short, much less than orbit diameter; anterolateral edge of snout not supported by bone Caelorinchus nazcaensis (p. 46) 8b. Snout long, much longer than orbit; an- terolateral edge of snout completely sup- ported by bone Caelorinchus immaculatus (p. 41) 9a. Ventral striae, consisting of parallel lines of black over a silvery ground, over much of isthmus, shoulder girdle, and ventral aspects of abdomen; ventral light organ with two lenslike bodies, one immedi- ately before anus and connected to an- other on the chest by a black midventral line (Fig. 6). (Hymenocephalus) 10 9b. No ventral striae; light organ not as above 14 10a. Body terete, head about as deep as wide; small serrations along leading edge of second spinous ray of first dorsal fin; gill rakers on lower limb of outer arch fewer than 15 Hymenocephalus gracilis (p. 55) Figure 6. Diagrammatic lateral (a) and ventral (b) views of Hymenocephalus showing extent of ventral striations and location of vent and light organs. 10b. Body laterally compressed, head deeper than wide; second spinous ray of first dorsal smooth; gill rakers on lower limb of outer arch more than 1 5 1 1 1 la. Pelvic rays 8; orbits 42-55% HL 12 1 lb. Pelvic rays 13-15; orbits 21^43% HL 13 1 2a. Many pigment cells scattered ventral and posterior to black blotch on dorsum of trunk and part of caudal region; pectoral rays i 1 4 — i 1 9 (usually i 1 6— i 1 7); interor- bital width 16.5-22.1, suborbital width 8.5-1 1.8, snout length 20.9-27.0% of HL Hymenocephalus semipellucidus (p. 60) 1 2b. Black blotch on dorsum with sharp out- lines, surrounded by singular pigment cells with few (if present) above and be- hind blotch; pectoral rays i 1 2-i 1 4 (rarely il5), usually i 1 2— i 1 3; interorbital width 20.8-27.4, suborbital width 5.2-9.5, snout length 16.4-23.8% of HL Hymenocephalus neglectissimus (p. 56) 13a. Orbits 21-23% HL; pelvic rays 13; bar- bel absent Hymenocephalus sp. cf. aterrimus (p. 54) SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 35 25 mm Figure 7. Diagrammatic ventral views of (a) Cetonurus and (b) Nezumia showing positions of pelvic and anal fins relative to periproct region. Arrow in (b) points to small fossa between pelvic fin bases. 13b. 14a. 14b. 15a. 15b. 16a. Orbits 35-42% HL; pelvic rays 14-15; barbel rudimentary or obsolescent Hymenocephalus striatulus (p. 63) Anus and urogenital opening surrounded by a broad naked margin, the entire re- gion (periproct) closely abutting or nar- rowly separated from anal fin origin, the anus closer to anal origin than to pelvic insertions; no accessory fossa of light or- gan anterior to anus (Fig. 7a) 1 5 Periproct moderate to small, broadly separated from anal fin origin by several scale rows, anus usually closer to pelvic insertions than to anal origin; usually a small fossa anterior to periproct (Fig. 7b) 17 Spinous ray of first dorsal smooth; pelvic origin below midbase of first dorsal fin, far behind vertical through pectoral or- igin Trachonurus villosus (p. 77) Spinous ray of first dorsal serrated; pel- vic origin below or anterior to origin of first dorsal fin, about on same vertical as pectoral origin 16 Head stout, somewhat depressed; snout long and pointed; no enlarged scales along dorsal and anal fins Mataeocephalus acipenserinus (p. 70) Figure 8. Diagrammatic lateral (a) and dorsal (b) views of Cetonurus. Arrow in (b) points to enlarged scales along second dorsal fin. 1 6b. Head soft, inflated; snout broadly round- ed (Fig. 8a); a series of enlarged spiny scales along dorsal and anal fins (Fig. 8b) Cetonurus crassiceps (p. 39) 17a. Lower jaw teeth large, widely spaced in 1 row; second spinous dorsal ray smooth; scales on branchiostegal membrane; an- terior fossa of light organ bean shaped (Fig. 9) Malacocephalus laevis (p. 68) 1 7b. Lower jaw teeth small, closely spaced, in 1 or more rows; spinous dorsal ray ser- rated or smooth; no scales on branchios- tegal membrane, or if scaled, dorsal ray serrated; anterior fossa usually tear-drop shaped 18 1 8a. Head notably large and broad, preoper- cle and suborbital bones deep and large; interorbital width 33-44%, suborbital width 19-26%, orbit-preop. 53-64% HL Pseudocetonurus septifer (p. 75) 1 8b. Head more normally proportioned for a macrourid; interorbital <31%, subor- bital <21%, orbit-preop. <47% HL 19 1 9a. Snout distinctly pointed and tipped with a spiny bifid tubercle; a suborbital shelf 36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 9. Diagrammatic lateral (a) and ventral (b) views of Malacocephalus. Arrows point to scales on branchiostegal rays. formed of two rows of coarse, modified scales; snout partially or extensively na- ked ventrally; mouth small, upper jaw length 26-38% HL {Nezumia) 20 1 9b. Snout bluntly rounded to pointed, tipped with a single, rather small tubercle or terminal tubercle absent; suborbital shelf of more than two rows or no modified scales; snout almost entirely scaled ven- trally; mouth moderate to large, upper jaw 35-55% HL 21 20a. Pelvic rays 14—18; inner gill rakers of first arch 9-12 Nezumia propinqua (p. 72) 20b. Pelvic rays 1 1 ; inner gill rakers of first arch 6-10 Nezumia convergens (p. 71) 21a. Pelvic rays 11; inner gill rakers of first arch 8-9; suborbital about 20% HL Kuronezwnia pallida (p. 65) 2 1 b. Pelvic rays 8—10; inner gill rakers of first arch 12-19; suborbital 8-14% HL (Ven- trifossa) 22 22a. Spinous second ray of first dorsal fin smooth Ventrifossa macrodon (p. 81) 22b. Spinous second ray of first dorsal fin ser- rated 23 23a. Barbel long, much greater than orbit di- ameter, about one-half HL Ventrifossa obtusirostris (p. 84) 23b. Barbel moderate to short, 20) and long. Branchiostegal rays 7, upper 2 on epihyal. Scales small, bearing few to many fine erect spinules. One dorsal fin, no portion elevated, spinous rays absent; pelvics ab- sent. Remarks. — Only the single species known. Macrouroides inflaticeps Smith and Radcliffe, 1912 Macrouroides inflaticeps Smith and Radcliffe in Radcliffe, 1912: 138, pi. 31, fig. 2 (Philippines, near Batan I., Lagonoy Gulf, Luzon, 13°23'S, 124°00'30"W, 408 fms [746 m]. Albatross sta. 5450). Norman 1939:48 (1 spec, near Maldives, Indian Ocean); Marshall 1964:92; 1973:516; Marshall and Taning 1966:1-5, pi. 1 (1 spec, South Atlantic off St. Helena); Parin et al. 1981:1 1 (1 spec, se. Pacific on Nazca Ridge); Shcher- bachev and Piotrovskiy 1982:45^47 (10 spec, Indian Ocean); Arai in Uyeno et al. 1983:209 (12 spec, Suriname and French Guiana). Parin 1 990: 1 7 (listed from area between Nazca and Sala y Gomez ridges). SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 39 Diagnosis.— As for genus. Distribution. — Philippines; Indian Ocean (including offSW tip of Australia); E and W trop- ical Atlantic; and E Pacific on Nazca Ridge. Mid- water to bottom in bathyal and abyssal depths (747-4,000 m) (Shcherbachev and Piotrovskiy 1982). Size.— To more than 48 cm. Remarks.— Only one specimen of this dis- tinctive species was taken off NSG. That 475 + mm individual, the largest known, was reported by Parin et al. ( 1 98 1 ). Radcliffe ( 1 9 1 2) and Mar- shall and Taning (1966) provide good descrip- tions and illustrations; Shcherbachev and Pio- trovskiy (1982) provide a distribution map plotting all known records of the species, includ- ing theirs from the Saya de Malha Bank and Naturaliste Plateau in the Indian Ocean. Material Examined. -ZMMGU 17683 (158 mm HL, 475 + mm TL); Nazca Ridge, 1,600-2,000 m; Hercules; midwater trawl 110. Squalogadus Gilbert and Hubbs, 1916 Type species: Squalogadus modificatus Gilbert and Hubbs, 1916, by original designation. Diagnosis.— As for Macrouroides, but having a small pelvic fin of five or six rays. Remarks.— Only the single species known. Squalogadus modificatus Gilbert and Hubbs, 1916 (Figure 2a) Squalogadus modificatus Gilbert and Hubbs, 1916:156, pi. 8, fig. 2 (offKyushu, Japan; 32°32'N, 132°23'W; 720 fms [1317 m]; Albatross sta. 4956). Golovan' 1978:222 (2 spec. West Africa, 1,450-1,550). Pakhorukov 1981:26 (3 spec. Whale Ridge; 850-1,550 m). Shcherbachev and Piotrovskiy 1982: 47 (17 spec, Indian Ocean; Pacific over Lord Howe Rise, 800-1,740 m). Shiobara 1982:143-146(2 spec, SurugaBay, Japan). Sawada in Amaoka et al. 1983:192 (1 spec, Iwate Pref., Japan; about 39°N). Parin 1990: 17 (listed from area between Nazca and Sala y Gomez ridges). Squalogadus intermedius Grey , 1959:330, fig. 53 (Gulf of Mex- ico, 29°07'N, 87°54'W; 600 fms [1 ,097 m]; Oregon sta. 1426) (See Marshall [1973] for additional earlier references.) Diagnosis.— As for genus. Size.— To more than 41 cm. Distribution. — Worldwide in tropical seas, and in temperate western North Pacific and North Atlantic, but not yet recorded from central and eastern North Pacific and western South Atlan- tic. Shcherbachev and Piotrovskiy (1982) pro- vide a distribution map, to which may be added our Nazca specimen, the first record from the eastern South Pacific. They state that the species is restricted to bathyal depths of 600-1,740 m, "where it may be found together with M. infla- ticeps." Remarks.— Our two specimens had six pelvic fin rays on each fin compared with the five gen- erally reported in this species. Squalogadus mod- ificatus is occasionally taken in great numbers in the north-central Gulf of Mexico. In an unpub- lished report of cruise 1 4 of the R/ V Oregon II (issued by the National Marine Fisheries Service, Pascagoula Fisheries Laboratory, Pascagoula, Mississippi), during a three-week period in Jan- uary 1970, "Squalogadus intermedius ^Squalo- gadus modificatus] . . . was taken frequently, catches ranging up to 2 1 9 pounds [about 1 00 kg] per 4-hour tow with a 70-foot [21.3 m] trawl." Deepwater trawling during that cruise was con- ducted at 500 to 1,000 fathoms [914-1,829 m]. Material Examined. -ZMMGU 1 7684 ( 1 52 mm HL, 459 + mm TL) and CAS 67408 (154 HL, 410 TL); Nazca Ridge, 1,050 m; Prof. Mesiatzevcv. 13, tr. 14. Subfamily Macrourinae The largest subfamily in the Macrouridae with more than 250 species, generally allocated to 30 or more genera. A diverse assemblage that may not be monophyletic. Size at maturity ranges from 12 cm in some Hymenocephalus to more than 1 60 cm in Albatrossia pectoralis (Gilbert, 1 892). Species predominantly benthopelagic, but a few strictly bathypelagic species known. Distribution worldwide in depths from about 200 m to more than 6,000 m. Cetonurus Gunther, 1887 Type species: Coryphaenoides crassiceps Gunther, 1878, by monotypy. Diagnosis. — Macrourine grenadiers with 7 branchiostegal rays. Head large, broad; interor- bital width much greater than orbit diameter; suborbital width more than orbit diameter; snout high, broad. Mouth rather small, subterminal; armed with small teeth in 1-3 irregular rows. Barbel very small to rudimentary. Head and body essentially completely covered with small scales beset with fine erect spinules; a series of enlarged scales along each side of dorsal fins and anteriorly along anal fin. Lateral line without grooved scales; lateral line course marked by a series of widely 40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 spaced black papillae (free neuromasts). First dorsal with a serrated spinous ray; V. 8-10. Anus within a broad naked area (periproct) that abuts anal fin origin. Swim bladder with 2 retia mirabi- lia. Distribution.— Warm waters of the Atlantic, Pacific, and Indian oceans. Usual capture depths range from about 1,000-1,800 m, but C. globi- ceps recorded from more than 4,200 m. Remarks.— Only two broadly distributed and closely related benthopelagic species. Cetonurus crassiceps (Gunther, 1878) Coryphaenoides crassiceps Gunther, 1878:25 (n. of Kermadec Is., Challenger sta. 170 and 171; 520 and 650 fm). Coryphaenoides {Cetonurus) crassiceps: Gunther 1887:143, pi. 37. Cetonurus crassiceps: Marshall 1973:613 (in key); Pakhorukov 1 976:327 (4 spec., Rio Grande Rise); 1981:25 (Walvis Ridge); Sazonov and Shcherbachev 1985:180, fig. lb, 2, 3 (diagnosis, distribution). Parin 1990:15 (listed from Nazca and area between Nazca and Sala y Gomez ridges). Diagnosis.— Orbits, 4.3-5.7 in HL (4.3-4.6 in NSG specimens); interorbital width about one- half HL; barbel very short to rudimentary, less than 3.5% HL; upper jaw extends to below pos- terior half of orbit. Counts and Measurements. — 1 D. 11,8-10; IP. il6-il9; V. 10-1 1; GR-I (outer/inner) (4)7- 9/1 + 1 + (10-11), GR-II (0-1) + 1 + (10-12) /l + 1 + (10-11); scales ID. 12-14, mid.-lD. 6-11, 2D. 10-13; caeca 10-11 (3 spec). Measurements in percent HL (8 spec, smallest excluded): postrostral 68.2-72.5; snout 35.4-42.7; preoral 32.3-38.0; orbit 21.7-23.0; interorbital 46.6-53.6; suborbital 23.7-30; postorbital 50.7- 58.9; orbit-preop. 50.3-58.2; upper jaw 31-37.1; gill slit 9.4-1 2.9; pre- ID. 99-1 06; pre-V. 95-1 27; pre-A. 118-139; body depth 73-95; V.-A. 23- 31; height ID. 39^16 (3 spec); base ID. 17.3- 21.7; 1D.-2D. 25.6-32.8; IP. 41-47 (2 spec); V. 26^42 (4 spec). Size.— To more than 44 cm. Distribution. — Cetonurus crassiceps has been reported from the subtropical South Atlantic, central tropical Atlantic, off Kermadec Islands, and off Hawaii (see Sazonov and Shcherbachev 1985: fig. 3). Records from the Nazca and Sala y Gomez ridges are the first in the southeastern Pacific, but are not unexpected. Surprisingly, however, the species is yet to be found in the Indian Ocean. Comparisons.— Sazonov and Shcherbachev (1985) readily differentiated C. crassiceps from its congener C. globiceps (Vaillant, 1888) and showed the circumglobal distribution of the lat- ter species. Their records show a common oc- currence of the two species only on the Sierra Leone Rise in the eastern Atlantic. The NSG specimens of C. crassiceps show no apparent dif- ference from representatives from other locali- ties. Material Examined.— (9 spec.) Sala y Gomez Ridge: ZMMGU 17688(5:29-68 mm HL, 112+-314+ mm TL) and CAS 75971 (3:41-73.5 HL, 183-315+ TL); 1,070-1,100 m; Prof. Mesiatzev cr. 13, tr. 10. Nazca Ridge: ZMMGU 17689 (1, 62 HL, 320+ TL); 940-960 m; Prof. Mesiatzev cr. 13, tr. 94. Caelorinchus Giorna, 1809 Type species: Lepidoleprus caelorhincus Risso, 1810, by sub- sequent designation. Diagnosis. — Macrourine grenadiers with branchiostegal rays 6. Snout blunt and barely protruding to greatly elongated and sharply pointed. Orbit large, oval to elliptical, the long axis horizontal (or nearly so). Mouth small and inferior to moderately large (to about 40% HL). Outer gill slit greatly restricted; no gill rakers on outer side of first arch. Ridges of head often stout and sharply spined. A stout suborbital ridge formed of modified sales running from tip of snout posteriorly onto preopercle, ending in a sharp point. Scales covered with spinules, which vary widely in size, shape, arrangement, and den- sity. Spinous first dorsal ray smooth (a few ter- minal denticles in a few species). Pelvic fin rays almost always 7 (6 in one species). Anus at or near anal fin origin. Ventral light organ variously developed; elongated with two large dermal win- dows at each end in some, to almost rudimentary in others; lens absent. Abdominal vertebrae 1 1- 1 2. Swim bladder oval to bilobed anteriorly; retia mirabilia usually 4, but number variable in a few species (as many as 9-1 1 in C. canus). Distribution. — Worldwide in tropical to temperate seas, but absent in high polar latitudes. Two species found south of Antarctic Conver- gence. Species most numerous in tropical waters. Remarks.— We follow Eschmeyer (1990:70) in the use of the spelling of the generic name. The dipthong ae as originally used by Giorna ( 1 809) was changed to oe by later workers on the SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 41 ^38* Figure 10. Caelorinchus immaculatus new species; holotype, ZMMGU 17692 (69 mm HL, 268+ mm TL) captured on Sala y Gomez Ridge in a trap set at 560 m, Prof. Shtokman cr. 18, sta. 1955/62, 29-30 April 1987. Photograph by Sergei Dudarev. assumption that the name Coelorinchus was de- rived from 'hollow snout.' The name Caelorin- chus, meaning burin snout, alludes to the loz- enge-shaped snout tip of the type species, C. caelorhincus (note: a burin is an engraving tool, sometimes having a lozenge- or diamond-shaped tip). The genus is a distinctive and diverse group with more than 100 species, although only about three-quarters of that number are named. Its re- lationship is closest to Macrourus (Okamura 1970a, b; Iwamoto and Sazonov 1988). Three of the four species recorded from NSG are most closely similar to species of the Indo-West Pa- cific. The fourth (C. nazcaensis) appears similar to C. aconcagua Iwamoto, 1978 from the con- tinental slopes of the eastern Pacific. Caelorinchus immaculatus new species (Figures 10, 11a, 12) Coelorinchus innotabilis: Iwamoto 1978:329-332 (in part; 1 spec, Chile).-Parin et al. 1981:1 1. "Coelorinchus sp. nova 2 Sazonov et Iwamoto": Parin 1990: 1 5 (listed from Nazca and Sala y Gomez ridges). Parin et al. 1990:42 (stomach contents). Kotlyar and Parin 1990:104, fig. 3v (otolith). Diagnosis. — Snout sharp, slender, longer than orbits, anterolateral margins completely sup- ported by bone. Subopercle forms a narrow pointed triangle. Upper jaw teeth in broad short bands that fall well short of posterior angle of jaws; upper jaw about one-fourth HL. Underside of head scaleless except for a small patch postero- ventrally on preopercle. Scales with short, greatly reclined spinules in parallel rows; scales below 2D. (7-8.5) + 1 + (1 1-13). Anterior rays of sec- ond dorsal almost as well developed as opposite members of anal fin; 1 D.-2D interspace less than base of first dorsal. No bold markings on body. Anus separated by 2-3 scale rows from anal fin; ventral light organ small with short pedicel. Counts and Measurements (data for holo- type in square brackets).— D. 11,10 [11,10] + 102- 1 13, A. 99-1 16, IP. il8-i21 [il9]; GR-I (outer/ inner) 0/(1-2) + (0-1) + (6-7) 8-9 total [1 + 1 + 7], GR-II + 1 + (5-7), 6-8 total/(l-2) + (6-8) 8-10 total [1+0 + 8]; scales below ID. 7-9 [9.5], below 2D. (7-10)+ 1 + (10.5-15) [9.5 + 1 + 10] (18.5-24 total scales in diagonal row to A.), below mid- ID. 5.5-8 [5.5], lat.l. 39-48 [41]. Total length of specimens examined 137-315 mm, HL 37.9-70 mm; the following in percent HL: postrostral 58-65 [6 1.7]; snout 37-43 [39. 1]; preoral 35^5 [40.7]; orbit 28-32 [31.0]; inter- orbital 19-23 [19.3]; suborbital 12-15 [13.6]; postorbital 27-33 [30.9]; orbit-preop. 30-35 [31.7]; upper jaw 22-26 [23.9]; gill slit 8-1 1 [10.9]; barbel 5.6-13.5 [12.3]; body depth 40-56 [55]; pre-A. 133-151 [151]; V.-A. 31^*5 [36]; height ID. 36-57 [47]; 1D.-2D. 5-15 [10.4]; IP. 37-45 [45]; V. 29^19 [37]. Description. — Head 3.8-4.2 in TL, about as wide as deep. Trunk and tail moderately com- pressed laterally. Anus slightly removed from anal origin a distance equal to about 0.5 diameter of pupil. Head ridges rather strong, sharp, and gen- erally narrow; head divided into dorsal and ven- tral parts by suborbital ridge. Snout sharply pointed; median and lateral processes of nasal bones connected and forming a complete bridge across anterolateral edges of snout. Interopercle completely hidden behind preopercle; ventral tip of subopercle produced into a flexible, pointed tab, its tip slightly exposed beyond posteroven- tral margin of preopercle. Gill membranes broadly connected to isthmus without a free pos- terior fold. Gill slits restricted, the outermost about as long as pupil diameter. Uppermost (epi- 42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 ^*&: ■ y" Figure 1 1 . Scales from dorsum below interspace between dorsal fins of: (a) Caelorinchus immaculatus; (b) C. innotabilis\ (c) C. nazcaensis; and (d) C. spilonotus. Drawn by Tomio Iwamoto. Scale bars equal 1.0 mm. branchial) gill rakers padlike; lower rakers tu- bercular and armed with very small spines. Chin barbel thin, short, less than diameter of pupil. Scales on body (Fig. 1 la) of moderate size and covered with short, reclined spinules, each aligned in close longitudinal rows (9-12 rows in 48 mm HL specimen, 12-13 in a 58 mm HL specimen); outer rows on field slightly divergent from mid- dle rows. Posteriormost spinules extend beyond scale margin. Middle spinule row not enlarged. Scales on head generally with more divergent spinule rows; those dorsally between head ridges mostly with small, reduced spinules or spinules absent. A small scutelike scale at posterior end of occipital sensory canal and before lateral line; no supraoccipital ridge scale. Terminal snout scute somewhat arrowhead-shaped in dorsal view, dorsoventrally flattened, and armed with several longitudinal rows of small spinules, the rows diverging posteriorly from the anterior apex. Broad areas around nostrils and behind leading anterolateral margin of snout naked. Underside of head, including mandible, completely and smoothly naked except for a narrow file of thin scales on posterior end of preopercle. Head pores not prominent. Light organ small, externally manifested in some specimens by a blackish median streak be- fore anus; streak lacking in others. Luminescent gland housed within body wall and in a flattened, black, oval structure connected to rectum. Entire length of gland about 0.5-1.2 pupil diameter. Intestine with multiple loops and somewhat similar to that of C. smithi Gilbert and Hubbs, 1 920, as illustrated by Okamura (1970b: fig. 65B). Pyloric caeca of three specimens 9, 9, and 10, short and unbranched. Gas bladder with 4 large retia and gas glands. Fins generally small; first dorsal shorter than postrostral length of head, its spinous second ray scarcely longer than adjacent segmented rays. Second dorsal close behind first dorsal, their in- terspace less than length of base of latter; ante- riormost rays of second dorsal relatively long for a macrourine grenadier, slightly shorter but dis- tinctly slimmer than anterior rays of anal fin. Outer pelvic fin ray somewhat prolonged beyond inner rays, reaching to anteriormost 1 to 3 anal rays; inner rays extend, at most, only to anus. Color in alcohol generally light brown to some- what swarthy on head, but abdomen and oper- culum dark bluish to blackish; scale pockets prominently outlined. Ventral surfaces of head grayish; in smaller specimens peppered with small melanophores. Anterior rim of orbits and sep- tum between anterior and posterior nostrils black. Fins all blackish or dusky. First dorsal in smaller specimens blackish basally, but pale distally; the distinction less obvious in large Prof. Mesiatzev specimens. Oral, branchial, and peritoneal lin- ings black. Gill arches blackish, but rakers and filaments pallid. We found two color variants. One variant, confined to specimens trawled on the Sala y Gomez Ridge, is very pale with almost unpig- mented pectoral fins and small melanophores scattered over naked areas of the head. The other variant, represented by all specimens from the Nazca Ridge and those taken in traps on the Sala y Gomez Ridge, have darkly pigmented pectorals and large melanophores densely covering naked areas of the head. The darkly pigmented head areas contrast strongly with the light adjacent scaled and ridged areas. The holotype and two paratypes (ZMMGU 1 8 1 1 6 and 1 7693 [2 1 3 mm TL]) attributed to the dark variant, however, have somewhat intermediate pigmentation patterns. (A 220+ mm paratype from ZMMGU 17643 is a more typical dark variant.) SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 43 Food.— Parin et al. (1990:42) found a high percentage of polychaet worms (Polinoidae and Tomopteridae) and copepods (the harpacticoid Cervicornia sp. and pelagic calanoids Clausocal- anus sp., Chirundia streetse, Bradyidius sp., Ae- tideus sp., Pseudocalanus sp., and Oncaea conif- era) in the alimentary canal of 20 individuals examined. Also present were shrimps, isopods, amphipods (Gammaroidea, including Oedice- rotidae), foraminiferans, and ophiurans (Ophi- ura). Size.— To about 32 cm. Distribution.— Nazca and Sala y Gomez ridges and off central Chile in 340-780 m, its primary depth distribution probably around 400- 550 m. Etymology.— From the Latin, immaculatus, not spotted; in reference to the lack of a distinc- tive color pattern in the species, in contrast to two other NSG species, C. spilonotus and C. mul- tifasciatus, which have prominent blotches on the body. Comparisons and Remarks. — Caelorinchus immaculatus is close in all diagnostic characters to C. karrerae Trunov, 1 984, a species originally described from material collected in the south- eastern Atlantic off South-West Africa (Namib- ia), Valdivia Seamount, Discovery Seamount, and also reported from the southwestern Indian Ocean. (Other specimens of C. karrerae, depos- ited in CAS, IOAN, ZMMGU, and ZIN, have been examined by us from the Saya de Malha Bank, Broken Ridge [West Australian Ridge], and Madagascar Ridge.) Differences between the two species are slight and may represent populational differences, but we think they are sufficient to justify our current treatment. In our comparison of specimens of the two species, C. immaculatus specimens have a somewhat more upturned snout, the orbit shape is less "squared off' an- teriorly than in C. karrerae, the count of diagonal scale rows below the second dorsal is slightly higher (18.5-24 vs. 16-20 in C. karrerae), the snout is shorter (37-43% HL vs. 42-46%), the posterior nostril is somewhat smaller (6.9-9.1% HL vs. 7.7-1 1.5%), and the upper jaw is longer (22-26% HL vs. 19-23%). Furthermore, there are two or three rows of non-spinulated scales (rarely with rudimentary ridges) lateral to the median nasal ridge in C. immaculatus, whereas in C. karrerae, there is a single row of large scales bearing 4-8 well-developed subparallel ridges with numerous small reclined spinules. 18 20 22 24 Orbit to angle of preopercle (mm) Figure 1 2. Scatter diagram comparing relationship of dis- tance orbit to angle of preopercle with orbit diameter in Cae- lorinchus innotabilis (plus signs) and C. immaculatus (circles). Diagonal line represents 1:1 ratio of the two measurements. The new species is also closely similar to the New Zealand and Australian species C. innota- bilis McCulloch, 1907, with which the single Chilean specimen was identified by Iwamoto (1978). It differs from that species, however, in having ( 1 ) an overall blackish first dorsal (C. in- notabilis has a pale first dorsal base and blackish tips); (2) no scales on underside of head above and lateral to mouth (C. innotabilis has patches of small scales there; (3) a shorter trunk and tail (HL about 3.7-4.2 in TL cf. about 4.2-5.5 TL in innotabilis); (4) a more prolonged preopercle (distance orbit to angle of preopercle less in C. innotabilis; see Fig. 12); (5) a longer upper jaw (22-26% HL cf. 1 9-21%); (6) a longer postorbital distance (27-29% HL cf. 25-26%); (7) usually more pectoral rays (i 1 8— 12 1 cf. i 1 5-i 1 9); (8) more scale rows below 1 D. and 2D; and (9) a somewhat deeper body (about 50% HL vs. less than 50%). The two color variants of C. immaculatus present a bit of a problem. Differences between the variants are not restricted to color, but also include some morphometric features. The dark variants have on average a shorter snout and shorter preoral length, and they are somewhat larger (all >59 mm, cf. <68 mm in pale vari- ants.). The different capture localities and cap- ture gear suggest some geographical and ecolog- ical separation. The differences may be related 44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 13. Caelorinchus multifasciatus new species; holotype, ZMMGU 18117 (32.7 mm HL, 135 mm TL), from Sala y Gomez Ridge in 439-500 m. Photograph by Sergei Dudarev. to ontogenetic changes in body shape and hab- itat. Certainly, the proportional-measurement differences are interrelated and attributable to the longer snout of the light variant. More spec- imens and study are necessary before we can firmly establish the taxonomic status of the two variants. Material Examined. — (dark variants noted with an aster- isk) Holotype: ZMMGU 17692* (69 mm HL, 268+ mm TL); Sala y Gomez Ridge, 560 m; Prof. Shtokman cr. 18, sta. 1955/ 62. Paratypes (85 spec, from 15 stations). CHILE. CAS 38314 (50.6 HL, 215 TL); 32°17'S, 71°39.5'W; 580 m; Anton Bruun cr. 18 A, sta. 702. Sala y Gomez Ridge: ZMMGU 17690 (2: 47.5-50 HL, 180-224 TL) and CAS 50895 (37.9 HL, 161 + TL); 535-574 m; Ichthyandr cr. 5. tr. 54. ZMMGU 17691 (39 HL, 177 TL); 25°02'S, 88°35'W; 550 m; Astronomer no trawl no. ZMMGU 17693* (2: 59.3-59.5 HL, 213+-220 TL); 580 m; Prof. Shtokman sta. 1956/61. ZMMGU 17694 (26: 27.4- 63.3 HL, 117-249 TL); 580-564 m; Prof. Shtokman cr. 18, sta. 1964. ZMMGU 17695 (13: 40.5-51.8 HL, 162-220 TL); 562-545 m; Prof. Shtokman cr. 18, sta. 1965. ZMMGU 17698 (61 HL, 243 TL); 550-630 m; Hercules Xt. 74. ZMMGU 181 10 (3: 53.5-57.5 HL, 214+-231+ TL); 540 m; Ichthyandr cr. 5, tr. 53. ZMMGU 18111 (37.7 HL, 151 TL) and CAS 50896 (2: 39.4-39.9 HL, 137+-139+ TL); 420 m; Ichthyandr cr. 5, tr. 57. ZMMGU 18112(48.2HL, 180+ TL); 550-560 m; Prof. Mesiatzevcr. 13, tr. 2. ZMMGU 181 13(18: 38-50.7 HL, 135- 2 14 TL); 563-590 m; Prof. Shtokman cr. 18, sta. 1976. ZMMGU 181 14 (32 HL, 147 TL); 545-600 m; Prof. Shtokman cr. 18, sta. 1977. Nazca Ridge: ZMMGU 17696* (5: 64-70 HL, 254- 315 TL) and CAS 75975 (4: 64-68 HL, 245+-307 TL); 340- 780 m; Prof. Mesiatzev cr. 13, tr. 44. ZMMGU 17697* (69.5 HL, 310 TL); 940-960 m; Prof. Mesiatzev cr. 13, tr. 94. ZMMGU 18115* (62.7 HL, 282 TL); 730-720 m; Prof. Shtok- man cr. 18, sta. 1828/35. ZMMGU 181 16* (63.6 HL, 261 TL); 505 m; Prof. Shtokman cr. 18, sta. 1879/82. Caelorinchus multifasciatus new species (Figures 13, 14) "Coelorinchus sp. nova 4 Sazonov et Iwamoto": Parin 1990: 1 6 (listed from Sala y Gomez Ridge). Diagnosis. — Body terete and shallow, width across pectoral bases about equal to greatest depth. Anterolateral margins of snout incom- pletely supported by bone. Underside of head Figure 14. Lateral view (a) and dorsal view (b) of holotype, ZMMGU 18117 (32.7 mm HL) of Caelorinchus multifasciatus new species. Drawn by Tomio Iwamoto. Scale bar equals 25 mm. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 45 completely naked except for a few scales over- lapping anterior edge of snout. Spinules on body scales in 5-6 discrete parallel rows, those on scales atop head between lateral occipital ridges in a single ridgelike longitudinal row on each scale. Second dorsal fin close behind first dorsal, its rays about as high as opposites of anal fin. A series of about 7 dark broad bands or saddles on body. A long light organ extending from anus almost to isthmus, the opposite ends expanded, each recessed in a shallow fossa. Counts and Measurements.— D. 11,10 + 85 + ; IP. il7-il8; GR-I (inner) 0+1+5, GR- II (outer/inner) 0+1+6; scales below ID. 6, mid-ID. 5.5, 2D. 6.5, lat.l. 35. Measurements in mm, percent HL in paren- theses: postrostral 18.3 (56.0); snout 14.6 (44.6); preoral 14 (42.8); internasal 6.5 (19.9); interor- bital 6.9 (21.1); orbit 8.2 (25.1); suborbital 4.4 (13.5); postorbital 9.7 (29.7); orbit-preop. 9.5 (29.1); upper jaw 7 (21.4); barbel 3.5 (10.7); gill slit about 3.0 (9.2); pre-D. 36 (1 10); pre-A. 51.3 (157); V.-A. 15.6 (48); body depth 15 (46); 1D.- 2D. 4.3 (13.1); base ID. 6.8 (20.8); height ID. 15.5 (47); IP. 14 (43); V. 14 (43). Description.— Body and head long, slender, terete; width over opercles about equal to body depth, width over pectoral base slightly less. Snout sharply pointed, a slender scute at tip; profile of snout viewed from above a slender convex cone. A prominent bony suborbital ridge, strengthened with thickened coarsely spined scales, runs from snout tip to preopercle, its posterior tip pointed, the ridge dividing the head into upper and lower parts. Orbits about 4 in head, shorter than snout, about 1.2 in postorbital length of head. Inter- orbital space flat, width slightly less than orbit diameter. Mouth small, inferior, rictus restricted; upper jaw less than orbit; barbel rather thick at base but tapers rapidly distally. Subopercle forms a narrow short tip posteroventrally. Branchiostegal membranes broadly attached to isthmus; gill openings strongly restricted, ex- tend anteriorly only to level of posterior edge of preopercle. Gill rakers few, tubercular; outer gill slit restricted, length about equal to barbel length. Scales of body with needlelike spinules aligned in 5 or 6 parallel rows, spinules in middle row slightly if at all larger than those in adjacent rows. Scales atop head somewhat elongated, with spi- nules erect and aligned in single, longitudinal, sharp, ridgelike rows. Dorsally on each side of median nasal ridge and behind leading edge of snout with scattered small scales, each covered with few erect spinules, and extensive naked ar- eas. A squarish naked area medially at end of occipital region and a few boundary areas on head with naked margins. Whether these naked areas become more extensively scaled in larger specimens is unknown. Ridges of head with modified scales; those on suborbital in 2 rows, heavily thickened and beset with stout spinules. Tip of snout with a slender sharp scute. Under- side of head entirely naked, although some scales overlap anterolateral snout edges onto ventral surface; whether these become more extensive or better defined in larger specimens is uncertain. Teeth in both jaws small and scarcely visible above thick gum papillae; those in upper jaw in short broad band that falls well short of posterior extent of rictus; those in lower jaw in a long tapered band roughly half width of premaxillary band. Fins rather small; first dorsal short, its second spinous ray slender with 3 fine needlelike spines near distal tip; second dorsal well developed and about equal in height to anal fin. Interspace be- tween dorsals short, much less than length base of first dorsal. Outer pelvic ray slightly thickened and prolonged, extending just past vent. Light organ large and prominently developed along ventral midline of abdomen and chest. Posterior end abuts anus, which lies immediately before anal fin; anterior end in a shallow fossa on chest, just behind isthmus and before pelvic fin bases, the two ends connected by a broad black median line. Pigment pattern highly distinctive, marked most prominently by a series of about seven broad saddles or bands, the anteriormost one at ante- rior end of nape and separated (by a narrow pale area that runs from hind margin of gill cover over nape) from a second band that extends over 9 or 10 scale rows to near the posterior border of the first dorsal. A faint diagonal pale strip leads to another more diffuse band that subtends only about 6 scale rows and ends under the 2nd or 3rd ray of second dorsal. A much darker band (9-10 scales wide) begins below the 5 th or 6th ray of second dorsal. Broad pale areas separate the remaining bands, which are dark and subtend 8-9 scale rows each; the last band blending im- perceptibly into darkened tail tip. Abdomen dark, somewhat bluish, covered with large melano- phores having silvery middle. Interorbital region with a dark narrow band; another across lateral 46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 15. Caelorinchus nazcaensis new species; paratype, CAS 50949 (70 mm HL, 280 mm TL), from Nazca Ridge in 340 m. Photograph by Susan Middleton. occipital ridges at posterior margin of naked area at head of nape. Dorsal aspects of head gray, underside pale, virtually white except for faint punctations over gill membranes, gular region, posteroventrally on preopercle, and along as- cending process of premaxilla. Anterior parts of mouth white, but inner walls dark. Lips and bar- bel white. Long spinous ray of first dorsal black on basal one-third or so, remainder dusky; fin somewhat darker distally and anteriorly. Other fins pale to sparsely flecked with small melano- phores. Lining of gill cavity, the gill arches, and rakers heavily punctate. Size. — Probably a small species, attaining at least 14 cm. Distribution. — Known only from a single small specimen taken on the Sala y Gomez Ridge in 439 m. Etymology.— From the Latin multo, many, and fasciatus, with bands, for the distinctive col- or pattern on the new species. Comparisons and Remarks.— The new spe- cies is one of the more striking members of the genus, with its slender, terete body, sharply pointed snout, and multiple bold saddle marks. It appears to be most closely related to C. cin- gulatus and C. spilonotus, the three being equally characterized by the combination of long light organ extending almost to the isthmus; antero- lateral margin of snout incompletely supported by bone; underside of head naked; long, slender, sharply pointed snout; spinules on scales nee- dlelike and arranged in relatively few parallel rows; rays of second dorsal about equal in height to those of anal; orbit about four in HL, longer than upper jaw; and saddle marks on body. Cae- lorinchus multifasciatus is easily distinguished from the other two species by its more numerous bands and more terete body; its median nasal process not dark as in the others; its first dorsal lacking a black tip; its snout somewhat longer and more slender; and its spinules on scales atop head in single, longitudinal, ridgelike rows. The species appears to be unavailable to trawl gear because of its rocky-bottom habitat; only a single small specimen was captured in a torn bottom trawl. The holotype was captured on one of the westernmost seamounts sampled during the 1 8th cruise of the Prof. Shtokman in spring of 1987. No other Caelorinchus was collected there, but specimens of Mataeocephalus were captured at depths of 739-763 m in the area surrounded by rocks near the peak of the sea- mount. Material Examined. — Holotype: ZMMGU 18117 (32.7 mm HL, 135 mm TL); Sala y Gomez Ridge, 439-500 m; Prof. Shtokman cr. 18, sta. 2023. Caelorinchus nazcaensis new species (Figures lie, 15, 16) Coelorinchus sp.: Parin et al. 1981:11 (brief description, 12 spec. Nazca Ridge, here reported). "Coelorinchus sp. nova 1 Sazonov et Iwamoto": Parin 1990: 15 (listed from Nazca Ridge). Parin et al. 1990:42 (stomach contents). Kotlyar and Parin 1990:104, fig. 3a (otolith). Diagnosis.— Snout shorter than orbit, antero- lateral margin not supported by bone, terminal scute sharply pointed; orbit 2.5-3.0 in HL, 0.9- 1.2 into postorbital length. Free end of suboper- cle variably prolonged into a flap. Mouth rather SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 47 Figure 16. Caelorinchus nazcaensis new species; paratype, CAS 50898 (77 mm HL, 322 mm TL), from Nazca Ridge in 340 m. Scale bar for (a) and (b) equals 25 mm; that for (c) equals 1.0 mm. Drawn by Amy Pertschuk. large for genus and unrestricted at posterior an- gle, upper jaw about one-third HL. Upper jaw teeth in narrow bands extending to posterior an- gle of mouth. GR-I about 12-14 total. Underside of head (including lower jaws) naked except for small patch of scales above end of mandible. Anus separated by 1-3 scale rows from anal fin; light organ very small, length slightly more than length posterior nostril; no pedicel. Counts and Measurements. — (data for ho- lotype in square brackets). D. 11,9-11 [11,10] + 118-129; IP. il9-i23 [i20/i21]; A. about 123- 138; GR-I (outer/inner) 0/(1-3) + 1 + (9-11) (total 12-14), GR-II ( 1 -2) + (0- 1 ) + (9- 1 1 ) (total 11-13) [1 + 1 + 10]/(l-3) + (0-1) + (8-11) (total 11-13) [3 + 1 + 10]; scales below ID. 7- 10 [8], below mid- ID. 5.5-7 [5], below 2D. 6.5- 8 [6], lat.l. 35-43 [42]; pyloric caeca 30-44. Head length 46-94 [67.7] mm; other dimen- sions given as percent HL: snout 28.2-32. 1 [31.3]; preoral 23.6-29.2 [29.0]; orbit 33.8-39.7 [37.4]; interorbital 18.9-24.5 [21.9]; suborbital 10.4- 13.3 [11.8]; postorbital 34.9-41.4 [33.5]; orbit- preop. 34.2-40.7 [35.6]; upper jaw 30.0-37.8 [33.8]; gill slit 14.6-22.3 [18.9]; barbel 10.9-18.1 [17.1]; body depth 58-79 [67]; pre-D. 100-111 [107]; pre-V. 104-116 [106]; pre-A. 130-156 [139]; V.-A. 32-51 [37]; height ID. 44-56 [52]; ID. base 23.2-28.0 [25.1]; 1D.-2D. 27.8-52.1 [38.7]; IP. 38-50 [51]; V. 29-42 [39]. Description. — Head large, about 4 in total length, broad, greatest width slightly less than greatest depth of head. Orbits huge, more than one-third HL, much longer than snout, 0.9-1.2 postorbital length, about 0.95-1 .2 of length from orbit to angle of preopercle. Posteroventral end of subopercle variously developed into a short flap directed ventrally or posteroventrally. In- teropercle completely hidden behind preopercle. Snout rather short, sharply pointed, tipped with a stout, narrowly conical scale. Median and lat- eral processes of nasal bones separated by a broad gap, leaving anterolateral margin of snout un- supported by bone. Mouth of moderate size, up- per jaw about one-third HL; maxilla extends to below middle of orbits. Dentition consists of small, conical, slightly recurved teeth in moderately wide bands, the mandibular band narrower than that of premax- illary. Premaxillary band about 6-7 teeth rows wide, tapering posteriorly to a few teeth wide and extending along most of mouth gape, but falling just short of end of rictus. Mandibular band sim- ilarly shaped but longer than premaxillary band, extending beyond end of rictus. Squamation characterized by rather large scales covering most of body and dorsal surfaces of head, but almost entirely absent on ventral head surfaces (a few small scales ventrally near junc- tion of infraorbital and preopercular ridges in 48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 some specimens). A narrow half-moon-shaped naked area on each side of leading edge of snout. Gill and nasal membranes naked. Suborbital shelf formed of two rows of stout, spiny, scutelike scales, the lower margin forming a distinct ridge that extends from snout to just short of pre- opercle angle. Angle formed by dorsal and ven- tral planes of suborbital region obtuse. Median nasal ridge low, formed by a row of about 8 broad scales, each covered with about 12 low spinule ridges that radiate from an anteromedian focus. Two rows of smaller scales on either side of me- dian nasal ridge, these separated laterally by a shallow naked groove from a row of scales along mesial side of supranarial ridges. Surpanarial ridges incompletely scaled in smaller paratypes. A narrow spiny ridge one scale wide separating orbit from nostril region. Head ridges other than those of suborbital not endowed with markedly heavy, enlarged, and spiny scutelike scales. No supraoccipital ridge. Scales of trunk below in- terspace of dorsal fins beset with 5-8 slightly di- vergent, ridgelike rows of short, sawtooth-shaped spinules (Fig. 1 lc). As with most species of ma- crourids, spinule rows on scales probably be- come more numerous with size. A small scaleless black fossa of light organ im- mediately preceding anus (fossa may be covered by scales in fully scaled specimens, but in our study specimens, scales missing from area). Anus separated from anal fin by 1-3 scale rows. Swim bladder in 72.2 mm HL male about 40 mm long, broader anteriorly, with two short horns directed anteriorly; closely attached to ribs on either side of vertebral column. Drumming mus- cles well developed in ventrolateral position be- hind each horn. A long oval window on dorsal surface. Five short broad retia, each terminating in a small globular gas gland (but a smaller, 70 mm HL male, CAS 50949, had 4 retia and gas glands). Swim bladder in a female of 88.9 mm HL similar in shape, but drumming muscles ru- dimentary and 4 retia. Fins all short; height first dorsal much less than postrostral length of head; pectorals slightly more than postorbital length of head and barely ex- tending to level of vent; pelvic rays fall well short of that level except for outermost ray, which just reaches to vent in some specimens. Color in alcohol overall swarthy to grayish brown, paler ventrally on head, but anteroventral margin of snout markedly darker than more pos- terior areas; opercular region darkish; abdomen bluish. Fins generally dusky, outer ray of pelvic fins whitish. Branchial, oral, and peritoneal lin- ings bluish to brownish black. Lips and barbel pale. Food.— Parin et al. (1990:42) recorded mostly pelagic food items in the stomachs of 10 speci- mens examined. These included the shrimp gen- era Gennada, Sergestes, Plesionika, the mysid Paralophogaster glaber, the copepod Pleuro- mamma, squid, and fish (Chauliodus sp., Lam- panyctus sp., Myctophidae gen., sp.). Size.— To about 40 cm. Distribution. — Known only from the type- specimens taken in 225-530 m on the Nazca Ridge. Etymology.— Derived from the type locality, the Nazca Ridge. Comparisons and Remarks. — Caelorinchus nazcaensis is closely similar to C. aconcagua Iwamoto, 1978, from the Pacific coast of Chile, sharing with that species a rather deep head (deeper than broad); relatively large mouth (al- most one-third HL); essentially completely na- ked underside of head (note exception in new species); naked areas on either side of leading edge of snout; small light organ without pedicel immediately before anus; rather wide outer gill slit (more than one-half orbit diameter); numer- ous gill rakers (12 or more total on first arch) compared with other members of genus; snout much shorter than orbit diameter; gill mem- branes narrowly joined across isthmus and form- ing a deep "V"; and similar-shaped opercular bones. The two species are readily differentiated by (1) orbit diameter (somewhat larger in C. aconcagua, 37.4-43.5% HL cf. 33.8-39.7%); (2) barbel length (shorter in C. aconcagua, 6.2-10.8% HL cf. 1 0.9-18.1%); (3) pectoral fin length (5 1 .9- 66.3% HL in C. aconcagua cf. 37.8-51.1%); (4) dark bluish color of trunk (extends around entire body in C. aconcagua, but restricted ventrally on belly below level of pectoral fin origin in new species); (5) oral cavity (white in C. aconcagua, blackish throughout in new species); (6) pyloric caeca (more than 30 in new species cf. 1 6-20 in C. aconcagua); (7) scale spinulation; and (8) head covering (thinner, and scales dorsally on head not as dense as in C. aconcagua). Caelorinchus nazcaensis also resembles C.fas- ciatus (Giinther, 1878), sharing with that and related southern hemipshere species (including C. aspercephalus Waite, 1 9 1 1 , C. australis (Rich- ardson, 1839), C. biclinozonalis Arai and Mc- SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 49 Figure 17. Caelorinchus spilonotus new species; paratype, ZMMGU 18124 (52.8 mm HL), from Sala y Gomez Ridge in 545-600 m, Prof. Shtokman cr. 18, sta. 1977, 1-2 May 1987. Photograph by Sergei Dudarev. Millan, 1982, C. bollonsi McCann and Mc- Knight, 1980, C. cookianus McCann and McKnight, 1980, and C. minis (McCulloch, 1926) the following features: (1) snout short, rather blunt; (2) orbits huge, much longer than snout, about equal to or greater than postorbital length; (3) anterolateral margin of snout not supported by bone; (4) light organ rather small, not ex- tending to level of bases of pelvics; (5) postero- ventral angle of subopercle rounded to variably produced, but not into a long narrow point. The new species differs from C. aspercephalus, C. australis, and C. biclinozonalis in lacking scales on the underside of the head (except for a few isolated ones in some individuals). Features dif- ferentiating C. nazcaensis from C. mirus, C. fas- ciatus, C. cookianus, and C. bollonsi include: lon- ger premaxillary tooth bands (not extending to end of rictus in others); mouth opening not no- tably restricted as in other species; first dorsal fin shorter than postrostral length of head (equal to or longer than in others); pectoral and pelvic fins shorter; spinules on body scales broadly saw- tooth-shaped (cf. smaller, finer, greatly reclined and imbricate); and terminal snout scute sharply pointed (blunt in others). Material Examined.— (all from Nazca Ridge) Holotype: ZMMGU 18094 (67.7 mm HL, 288 mm TL); 235-250 m; Prof. Mesiatzev cr. 12, tr. 23. Paratypes (33 spec): CAS 50949 (2:65-70 HL, 270-280 TL); 340 m; Ichthyandr cr. 6, tr. 1 . CAS 50948 (54 HL, 240 TL); 320 m, Ichthyandr cr. 6, tr. 2. CAS 50898 (77 HL, 322 TL); 340 m; Ichthyandr cr. 6, tr. 3. CAS 50897 (84 HL, 350 TL) and ZMMGU 17699 (8:46-89 HL, 197-383 TL); 330 m; Ich- thyandr cr. 5,tr. 13. ZMMGU 17700(59.5 HL, 242 TL); 330- 340 m; Ichthyandr cr. 5, tr. 14. ZMMGU 17701 (3:60-93 HL, 270-402 TL) and CAS 50899 (69 HL, 265+ TL); 330 m; Ichthyandr cr. 5, tr. 15. ZMMGU 17702 (2:69.5-75 HL, 270+- 307 TL); 310-330 m; Ichthyandr cr. 5, tr. 17. CAS 50900 (2: 70-76 HL, 290-305 TL); 330 m; Ichthyandr cr. 5, tr. 33. ZMMGU 17703 (6:71-82.5 HL, 289-357 TL); 330 m; Prof. Shtokman cr. 18, sta. 1845. ZMMGU 17704 (4:53.5-86 HL, 225-370 TL); 330-350 m; Prof Shtokman cr. 18, sta. 1851. ZMMGU 1 7705 (82 HL, 3 10 TL); 530-500 m; Prof. Shtokman cr. 18, sta. 1864/76. Non-type material (47 spec): ZMMGU 18093 (2:62.2-70 HL, 239+-295 + TL); 300-330 m; Astronomer trawl without no. (23°30.5'S, 81°45'W). ZMMGU 18095 (78.4 HL, 336 + TL); 235-225 m; Prof. Mesiatzev cr. 13, tr. 25. ZMMGU 18096 (59 HL, 255 TL); 340-325 m; Prof. Mesiatzev cr. 13, tr. 35. ZMMGU 18097 (72.7 HL, 329 TL); 235-230 m; Prof. Mesia- tzev cr. 13, tr. 36. ZMMGU 18098 (70.5 HL, 294+ TL); 230- 240 m; Prof. Mesiatzev cr. 13, tr. 29. ZMMGU 18099 (2: 58- 59.5 HL, 247-253 TL); 320-325 m; Prof. Mesiatzev cr. 13, tr. 43. ZMMGU 18100 (3:63-80.5 HL, 267+-324+ TL); 340- 780 m; Prof. Mesiatzev cr. 13, tr. 44. ZMMGU 18101 (68.5 HL, 286 TL); 225-240 m; Prof Mesiatzev cr. 13, tr. 109. ZMMGU 1 8 1 02 (9 1 HL, 399 TL); 225-240 m; Prof Mesiatzev cr. 13, tr. 31. ZMMGU 18103 (2:72-74.5 HL, 295-315 TL); 230 m; Prof. Mesiatzev cr. 13, tr. 120. ZMMGU 18104 (81.8 HL, 342 TL); 320-340 m; Ichthyandr cr. 6, tr. 10. ZMMGU 18105 (72.3 HL, 278+ TL); 230-250 m; Odisseycr. 2, tr. 15. ZMMGU 1 8 106 (2:65.5-7 1 .3 HL, 273-297 TL); 235 m; Odis- sey cr. 2, tr. 11. ZMMGU 18107 (80 HL, 348 TL); 230 m; Odisseycr. 2, tr. 14. ZMMGU 18108 (75 HL, 320 TL); 225- 247 m; Prof. Shtokman cr. 18, sta. 1867. ZMMGU 18109 (5: 68-87.5 HL, 292-366 TL); 235 m; Prof Shtokman cr. 18, sta. 1873. IOAN uncat. (21:61-92 HL, 252+^100 TL); 340 m; Akademik Kurchatov cr. 34, sta. 3594. Caelorinchus spilonotus new species (Figures lid, 17, 18) Coelorinchus cingulatus: Parin et al. 1981:11 (12 spec, from Sala y Gomez Ridge). "Coelorinchus sp. nova 3 Sazonov et Iwamoto": Parin 1990: 15 (listed from Sala y Gomez Ridge). Kotlyar and Parin 1990:104, fig. 3b (otolith). Diagnosis. — Snout acutely pointed, tipped with a narrow, pointed scute; median nasal bone blackish; anterolateral margins of snout not sup- ported by bone; ventral surfaces of snout entirely naked. Subopercle posteroventrally produced into a short triangular flap. Mouth small, restricted at posterior angle; upper jaw 20-26% of HL. Up- per jaw teeth in broad short bands that fall short of posterior angle of mouth. Scales on body with slender, conical, reclined, imbricate spinules ar- ranged in 5-10 parallel to slightly divergent rows (more rows in larger specimens). Anterior rays of second dorsal fin as long as opposite members of anal fin. Interspace between dorsal fins short, 50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 v^V\^ v - Figure 18. Caelorinchus spilonotus new species; paratype, CAS 50950 (50.3 mm HL), from Sala y Gomez Ridge in 440 m. Drawn by Tomio Iwamoto. Scale bar equals 25 mm. less than base of first dorsal. Two prominent sad- dle marks, one immediately before first dorsal fin connecting to broad pectoral blotch, second saddle separated from first by a distance about equal to snout length. First dorsal fin black tipped. Anus immediately before anal fin. Light organ extends forward between pelvic fins and onto middle of chest, both ends expanded into a tear- drop shape and lying within a shallow fossa. Counts and Measurements (from 26 speci- mens; mean, x, in parentheses, holotype in square brackets).- ID. 11,8-10 (11,9.35) [11,9]; IP. i 1 7— i22 (i 19.73, n = 52) [il9]; GR-I (outer/inner) total 6-8 [0/0 +1 + 5], GR-II total 5-7 [0 + + 5/1 + 1 + 5]/total 6-8; scales below ID. 6.5- 8 [7.5], below 2D. 5.5-7.5 [7], below mid-ID. 4.5-5.5 [5], lat.l. 37-44 [43]. Total length of specimens examined +96-206 mm [184+ mm], HL 29.5-60 mm [60 mm]. The following in percent HL: postrostral 49.7-60.5 (55.5) [54.5]; snout 41.8-51.4 (46.1) [46.7]; preoral 35.4-44.8 (39.1) [40.5]; internasal 17-22 [19.3]; orbit 21.2-27.9 (24.7) [24.2]; interorbital 18.0-24.1 (20.6) [21.0]; suborbital 10.8-14.8 (12.5) [12.8]; orbit-preop. 25.3-32.8 (29.8) [29.2]; upper jaw 20.3-26.0 (24.0) [24.2]; gill slit 8.6- 13 [13.3]; barbel 4.8-9.0 (6.9) [7.8]; body depth 37.6-47.5 (41.7) [44.2]; light organ 47.3-60.7 (51.7) [53.3]; pre-D. 102-115 (107) [107]; pre- V. 100-115 (107) [106]; pre-A. 139-161 (150); V.-A. 38-55.8 (46.6) [46.7]; height ID. 34-49 (41.3) [44]; ID. base 13.8-21.6 (18.7) [17.8]; 1D.- 2D. 4.2-13.3 (9.2) [9.5]; IP. 37-45 (40) [38]; V. 27-39 (33) [33]. Description. — Head about as deep as wide. Snout sharply pointed; anterolateral margins not supported by bone; a broad gap between median and lateral processes of nasal bone. Head ridges distinct but not especially strong, except for sharp suborbital ridge, which divides ventral and dor- sal portions of head. Supraoccipital ridge weakly developed. Interopercle completely hidden be- hind preopercle; subopercle developed postero- ventrally into a short triangular flap. Gill mem- branes broadly attached to isthmus, without a free posterior fold. Gill slits somewhat restricted, outer slit slightly longer than pupil diameter. Gill rakers low, epibranchial rakers shaped like flat- tened disks; remainder tubercular. Chin barbel short, almost rudimentary, usually shorter than length of posterior nostril. Scales of body (Fig. 1 Id) moderate sized, cov- ered with parallel to slightly divergent rows of small, sharp, conical, imbricate spinules, the pos- teriormost tips extending beyond margin of scale. About 5 or 6 spinule rows in smallest specimens examined (about 30 mm HL); 9 or 10 in largest specimens (about 60 mm HL). Scales on head generally like those of body, but with shorter, more erect spinules in fewer, more divergent rows. Nostril membranes completely naked, but sub- orbital shelf below and behind nostrils complete- ly scaled. Narrow strips on either side of median nasal ridge naked. Terminal snout scute sharp, rather small, narrow, and weak (tip broken off in several specimens); protrudes directly forward or slightly upwards; upcurve of tip more highly developed in smaller specimens. Light organ large, long, extending from anus to chest just behind isthmus. Luminescent glands as described and illustrated for the subgenus Quincuncia by Okamura ( 1 970b:figs. 43, 81). Al- imentary canal about like that illustrated for Caelorinchus longissimus by Okamura (1970b: fig. 64D). Extent of anteriormost bends could not be accurately determined because all specimens SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 51 had everted stomachs that distorted bending pat- tern. Pyloric caeca short, thick; 9-12 in 9 spec- imens. Swim bladder oval, blunter end anteriad, with 4 or 5 (in 5 specimens) short retia, each tipped with a broad, flat gas gland. Eggs in the large ovaries of CAS 50951 (58.8 mm HL) dis- tinct, the largest about 1.0 mm in diameter. Fins generally small, height first dorsal about as long as snout, second spinous ray not pro- duced. Pectoral fin short, about equal to snout length. Outer pelvic ray about equal to or slightly longer than postorbital length of head, falls short of anal fin. Second dorsal relatively well devel- oped for genus, height of rays anteriorly about equal to opposite members of anal fin. Interspace between dorsal fins short, less than length base of first dorsal. Color in alcohol light brown to tawny overall, paler ventrally on head and on tail; darker ven- trally on trunk. Saddles and blotches darker brown. Two prominent saddles, one before first dorsal fin, the second posteriorly about one snout length behind first and originating at position of 9th ray of second dorsal fin. The first saddle blending in larger specimens with large blotch above pectoral fin base. This pectoral blotch 5 or 6 scale rows wide, posterior saddle 6 or 7 rows wide. A faint blotch in larger specimens between dorsal-fin interspace and lateral line. Underside of head entirely lacking pigmentation except for fine scattered punctations posteriorly. Median process of nasal bone blackish or dusky. Mouth pale except for scattered punctations posteriorly on roof of mouth and in gullet. Gill chamber and gill membranes pale with scattered punctations; gill arches and filaments pale. Peritoneal mem- brane blackish; stomach pale to dusky near esophagus. Fin rays whitish except for black- tipped first dorsal, and uppermost pectoral ray often darker. Food. -One female (58.8 mm HL, CAS 5095 1) had a large squid beak in its otherwise empty stomach; all other specimens had everted stom- achs. Size. — A small species, maximum length slightly more than 20 cm. Distribution.— Known only from the Sala y Gomez Ridge and Hawaiian Islands in 330- 600 m. Etymology.— From the Greek, spilos (spot), and notos (back), in reference to the peculiar col- oration of this species. Comparisons. —The species is most similar to, and may be conspecific with, C. cingulatus Gil- bert and Hubbs, 1 920, a species known only from the two small type-specimens taken in the Phil- ippines and one 232 mm specimen recently re- corded from the Okinawa Trough by Okamura (in Okamura and Kitajima 1984:229, 366, fig. 161). The holotype (USNM 7822 1 , 40.2 mm HL) and small paratype (41.4 mm HL, 148+ mm TL; USNM 78233) of C. cingulatus were ex- amined for comparison. Most meristic and mor- phometric features of the two species are similar, but their pigmentation patterns differ. (Pigmen- tation in the paratype is almost totally lost from long preservation; thus, color-pattern compari- sons were made from the original description and from the description and illustration by Oka- mura in Okamura and Kitajima 1 984.) Caelorin- chus cingulatus lacks the large blotch above the pectoral fin base that is so prominent in C. spi- lonotus. Furthermore, the first saddle mark be- gins on the nape in front of the first dorsal fin and extends ventrally to join the pectoral blotch. None of this saddle lies below the soft rays of the first dorsal fin, as described and illustrated by Okamura for C. cingulatus. The second saddle mark does not extend below the lateral line in the new species, but does so in C. cingulatus, and the second saddle begins under the 7th- 10th rays of the second dorsal in the new species, but under the sixth in C. cingulatus. Gilbert and Hubbs (1920:483) describe brown streaks that "radiate backward from the eye . . . the upper one, more conspicuous, extends hor- izontally backward, just below the postorbital scaly ridge, to the upper angle of the branchial aperture, where it is continuous with the dusky opercular blotch." This upper streak is promi- nent in Okamura's (in Okamura and Kitajima 1984, fig. 161, p. 228) photograph of his speci- men but completely lacking in our specimens of C. spilonotus. In C. cingulatus a diagonal streak extends anteriorly and ventrally from the second saddle mark, joining the darker abdominal re- gion. This streak is also absent in C. spilonotus. The new species may have fewer gill rakers on the inner series of the second arch (7 or 8 cf. 9 in C. cingulatus) and fewer pyloric caeca 9-12 (cf. about 14 in the holotype and 15 in the para- type of C. cingulatus, although Gilbert and Hubbs [1920:482] reported 21 for the paratype). With so few specimens of C. cingulatus, these enu- merated characters cannot be adequately as- sessed to determine their value as species char- 52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 acters. The possibility exists that they may simply reflect individual or geographic variation within one species. The three Hawaiian specimens of C. spilonotus have noticeably longer, more attenuated snouts, with the terminal scute narrow, long, and sharply pointed. Naked areas laterodorsally behind the leading edges of the snout are more extensive than in the NSG specimens. Punctations on the underside of the head are lacking in the NSG specimens but noticeably present in the Hawai- ian specimens (and in C. cingulatus). Head col- oration of the Hawaiian specimens is slightly darker than that of the NSG specimens and fol- lows more closely the color description of C. cin- gulatus. We do not consider these differences as sufficient to recognize the Hawaiian population as distinct. The elongation of the snout is rem- iniscent of the situation in C. caribbaeus (Goode and Bean, 1885), where the snout length is re- markably variable within a single population in the northern Gulf of Mexico (see Marshall and Iwamoto 1973). Among other Pacific members of the genus, the new species and C. cingulatus share many important characters with C. gladius Gilbert and Cramer, 1897 from the Hawaiian Islands. The chief differences are: C. gladius lacks saddle marks but has a prominent black blotch over the pec- toral base; the snout is sharp and narrow, with the terminal scute remarkably long and devoid of spinules dorsally. The second spinous ray of the first dorsal fin has a few minute denticula- tions distally along the anterior margin in the two C. gladius specimens examined (47 and 66 mm HL, CAS-SU 8517), and surprisingly, also in the Hawaiian specimens of C. spilonotus. Cer- tain proportions differ notably as a result of the extremely elongated snout of C. gladius (e.g., the snout length is considerably greater than the postrostral length of the head in C. gladius but is much shorter than that measure in C. spilono- tus from the NSG, and about equal in the Ha- waiian representatives). Material Examined. — Holotype: ZMMGU 18125. (60 mm HL, 187+ mm TL); Sala y Gomez Ridge, 545-600 m; Prof. Shtokman cr. 18, sta. 1977. Paratypes (82 spec). Sala y Gomez Ridge: CAS 50950 (50.3 mm HL, 182 mm TL); 440 m; Ichthyandr cr. 5, sta. 40. CAS 50952 (2:30.6-38.4 HL, 108+-1 15 + TL); 480 m; Ichthyandr cr. 5, sta. 50. ZMMGU 17706 (4:42.5-56.5 HL, 138+-204 + TL); 540 m; Ichthyandr cr. 5, tr. 53. ZMMGU 17707 (6:29.5- 58.5 HL, 96+-201 TL) and CAS 50951 (6:43.5-58.8 HL, 131+-202 TL); 410 m; Ichthyandr cr. 5, tr. 56. CAS 50953 (56.2 HL, 192+ TL); 420 m; Ichthyandr ex . 5, tr. 57. ZMMGU 17708 (12:30.5^3.6 HL, 116-162 TL); 380 m; Prof. Shtok- man cr. 18, sta. 1940. ZMMGU 17709 (5:35-52 HL, 133-188 TL); 410-385 m; Prof. Shtokman cr. 18, sta. 1941. ZMMGU 18123 (15:42.7-52.3 HL, 130+-172 TL); 563-590 m; Prof. Shtokman cr. 18, sta. 1976. ZMMGU 18124 (30:25-58 HL, 98-198 TL); 545-600 m; Prof. Shtokman cr. 18, sta. 1977. Hawaiian Islands: LACM 45409-1 (3:52.7-58.2 HL, 176+- 181+ TL); Townsend Cromwell tr. 57; Mar-Apr 1972. Non-type material (7 spec). Sala y Gomea Ridge: ZMMGU 18118 (30.5 HL, 97+ TL); 360-400 m; Ichthyandr cr. 5, tr. 51. ZMMGU 181 19 (2:48.5-56.5 HL, 162+-182+ TL); 535- 575 m; Ichthyandr cr. 5, tr. 53. ZMMGU 18120 (44.5 HL, 157 TL); 550 m; Astronomer trawl without no. (25°02'S, 88°35'W). ZMMGU 18121 (48.8 HL, 174 TL); 565 m; Prof. Mesiatzevcr. 15, tr. 50. ZMMGU 18122 (2:44.7-54 HL, 138+- 170+ TL); 400 m; Prof. Mesiatzev cr. 15, tr. 52. Coryphaenoides Gunnerus, 1765 Type species: Coryphaenoides rupestris Gunnerus, 1765, by monotypy. Diagnosis. — Macrourine grenadiers with 6 branchiostegal rays. Snout shape variable but never greatly produced beyond mouth. Subor- bital ridge variously and usually weakly devel- oped, not connected to preopercular ridge, which in turn is never sharply angular at its posterior end. Dentition variable among species, arranged in broad bands to 1 or 2 rows, but teeth never few and fanglike. Barbel present. Second spinous ray of first dorsal finely serrated along leading edge, although serration occasionally obsolete in large individuals of some species. Gill rakers tu- bercular to short and tablike, those of outer series of first arch rudimentary. Anus immediately in advance of anal fin or slightly anterior to it; no light organ. Pyloric caeca simple, unbranched, usually fewer than 20. Retia and gas glands 4-7. (Adapted from Iwamoto and Sazonov 1988.) Coryphaenoides paradoxus (Smith and Radcliffe, 1912) Macrourus paradoxus Smith and Radcliffe in Radcliffe 1912: 115-116, pi. 25, fig. 1 (off eastern Palawan, Philippines, 2,021 m). Coryphaenoides (Nematonurus) paradoxus: Gilbert and Hubbs 1916:143. INematonurus macrocephalus Maul, 1951:17 (type locality Madeira). ^Coryphaenoides macrocephalus: Marshall and Iwamoto 1973: 575-578 (North Atlantic). Coryphaenoides sp.: Parin et al. 1981:1 1 (Nazca Ridge). Coryphaenoides paradoxus: Wilson et al. 1985:1,243-1,254 (Darwin Guyot, central North Pacific, about 1,600 m). Iwa- moto and Sazonov 1988:72-75, fig. 3c, 24, 28 (se. Pacific). Parin 1990:16 (listed from Nazca ridge and area between Nazca and Sala y Gomez ridges). SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 53 Diagnosis.— A large species (120+ cm) in subgenus Coryphaenoides with ID. 11,9-11, IP. il6-i21, V. 9-11 (rarely 8). Snout low, scarcely protruding. Orbits relatively small, about 1.5 into snout, 1.8-1.9 into interorbital space, 5.0-6.0 into HL. Interopercle barely exposed as a broad fleshy naked tab. Suborbital region vertical; no strong sharp ridges on head. Mouth large, almost ter- minal, upper jaw extends to below posterior orbit margin. Teeth prominent, an outer enlarged se- ries on premaxillary behind which a narrow vil- liform band; mandibular teeth in about 3 irreg- ular series near symphysis, becoming uniserial posteriorly. Head fully scaled except lips, gill membranes, interopercle. (After Iwamoto and Sazonov 1988.) Size.— A large species attaining more than 1 20 cm. Distribution. — Broadly distributed in the Pa- cific (Kermadecs, Hawaiian Islands, Nazca Ridge), Indian Ocean [our data], and Atlantic (if C. macrocephalus is a synonym). Remarks.— The two specimens from NSG were fully described by Iwamoto and Sazonov (1988). In the eastern Pacific, the species is likely to be confused only with C. bulbiceps, which at- tains a similar large size. The stronger teeth, lon- ger barbel, smaller orbit, lower snout, and higher pelvic fin ray count readily distinguish C. par- adoxus from C. bulbiceps. Although we (Iwa- moto and Sazonov 1988:75) previously alluded to the similarity of C. paradoxus and C. rudis Gunther, 1 878, we now strongly suspect that the two are the same. We have examined the type specimens of C. paradoxus and C. rudis and found no characters that would suggest specific differ- ence. The scale spinules in the C. rudis lectotype appeared longer and more densely placed, but these features of the spinules tend to be highly variable in most grenadiers. If this synonymy proves correct, C. rudis has priority. We are re- luctant at this point, however, to make this syn- onymy because of the lack of comparative ma- terial from near the type locality. As far as we can determine, only the lectotype of C. rudis is known (the paralectotype is a species of Nezu- mia). McCann and McKnight's (1980:35) record of the species from off New Zealand stems from misidentification of a specimen representing a perhaps-undescribed species. Brauer's (1906:246) record from the western Indian Ocean is based on a specimen (ZMB 1 764 1) representing an un- described species of Kuronezumia. Material Examined. -ZMMGU 16528 (155 HL, 765 TL); Nazca Ridge, 980 m; Ichthyandrct. 5, sta. 1. ZMMGU 16527 (166 HL, 820 TL); Sala y Gomez Ridge, 1,070-1,100 m; Prof. Mesiatzev cr. 13, tr. 10. Hymenocephalus Giglioli, 1884 Type species: Hymenocephalus italicus Giglioli, in Giglioli and Issel, 1 884, by original designation (also monotypic). Diagnosis. — Branchiostegal rays 7. Head bones weak, often paper-thin. Head covering membranous. Ventral striae on abdomen, chest, and shoulder girdle; light organ with two lens- like bodies, one before the anus, the second on the chest anterior to pelvic fin bases, both con- nected externally by a long median-ventral line. Distribution.— The genus is primarily trop- ical to warm temperate in distribution and gen- erally absent in high-latitude waters. With five species, the genus (sensu lato, including Spico- macrurus Okamura, 1970a and Hymenogadus Gilbert and Hubbs, 1920) is surprisingly well represented on the Sala y Gomez and Nazca ridg- es. Four species are found off the Hawaiian Is- lands. The genus is absent, however, along the entire mainland Pacific coasts of the Americas (although a single, apparent stray of undeter- mined species was reported by Iwamoto 1979: 140). In contrast, the genus is well represented in the western Pacific. Gilbert and Hubbs (1920) re- corded seven species and three subspecies from the Malay Archipelago (to include their terms "Philippine Islands and East Indies"); Okamura {in Okamura and Kitajima 1984) recorded five species from the Okinawa Trough, and six spe- cies and two subspecies (Okamura 1970a) from waters off Japan. The number of species in the Indian Ocean is as yet undetermined, but prob- ably ranges between two and four in the western and central parts, and more from the Malay Ar- chipelago and Australia. No Hymenocephalus species is known from New Zealand; only two have been reported from Australia, although rep- resentatives of other species are deposited in IOAN, ZMMGU, and AMS from warm-water regions of that continent. The Atlantic has only four representatives. Some of the species of Hymenocephalus have wide distributions; others appear to be rather narrowly confined. Hymenocephalus gracilis is an example of a widely distributed species, hav- ing first been described from off the Philippines and subsequently recorded from Japan (Oka- 54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 1 9. (a) Hymenocephalus sp. cf. aterhmus. (b) H. neglectissimus, holotype ZMMGU 18219(18.5 mm HL, 1 23 + mm TL) from offSala y Gomez Ridge in 580-564 m. Drawn by Y. I. Sazonov. mura 1970a), the Atlantic (Marshall and Iwa- moto 1973), the western Indian Ocean (Iwamoto 1982), and now the southeastern Pacific. The Hawaiian species H. aterhmus has also been re- corded from the western Atlantic (Marshall and Iwamoto 1973) and western Indian Ocean (Shcherbachev 1987); a related form is here re- ported from NSG. Hymenocephalus striatulus, originally described from Hawaii, is now known from Soviet collections from the NSG, and its closest relative appears to be H. billsamorum Marshall and Iwamoto, 1973, from the Gulf of Mexico and Caribbean Sea. Hymenocephalus longiceps Smith and Radcliffe, 1912 and H. stria- tissimus Jordan and Gilbert, 1904, are wide- spread in the western Pacific, from Japan south to Indonesia; the latter is represented by four geographic subspecies, and its closest relative ap- pears to be the new species from NSG. Most of the remaining species have relatively confined distributions, although available collections are still inadequate to circumscribe their limits with certainty. Remarks.— A distinctive group of about 22 or more small species, few attaining lengths of more than 25 cm. Hymenocephalus sp. cf. aterrimus Gilbert, 1 905? (Figure 19a) ^Hymenocephalus aterrimus Gilbert, 1905:666, pi. 93 (Holo- type: USNM 51649; off Kauai, Hawaii, 385-500 fms; Al- batross sta. 3989). Parin 1990:16 (listed from Nazca Ridge and area between Nazca and Sala y Gomez ridges). Diagnosis.— A species of subgenus Papyro- cephalus, with orbit diameter 21-23% HL; in- terorbital width 24-35% HL; outer gill slit 27- 30% HL. Barbel absent. Pelvic fin rays 13. Color mostly blackish. Counts and Measurements. — 1 D. 11,9-10; IP. il3-il5; GR-I (outer/inner) 17-19/4 + 17- 19; GR-II 3 + 17-18/3 + 15-16. Total length 84-136 mm; HL 26.5-35 mm. Following di- mensions in percent HL (3 spec): postrostral 71.4-75.5; snout 26.4-34.3; suborbital 20-21.9; postorbital 48.5-51; orbit-preop. 52.8-55.4; up- per jaw 45.4^48.7; body depth 69-76; light organ 59-68; pre-lD. 104; pre- V. 103-1 10; pre- A. 147- SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 55 160; V.-A. 50-59; base ID. 31.4-37.9; 1D.-2D. 32.1-40.8; IP. 70; V. 60. Description.— Trunk and tail strongly com- pressed laterally, dorsal and ventral profiles taper rapidly behind first dorsal fin for about one HL distance. As typical for subgenus, head large and deep, bones generally paper-thin, scale bones over occipital region large and thin. Orbits small, more than 4 into head. Snout high, median nasal ridge notably elevated. Subopercle deep; propercle large, forming most of gill cover, outer edges crenulated; opercle commensurately small; a deep subopercular notch. Preopercular ridges forming a large triangular process at posteroventral cor- ner. Teeth in moderately wide bands in both jaws, the individual teeth uniformly short and stoutly conical with rather blunt tips. Scales all missing, but scale pockets large. Anterior lens of light organ small, round, dif- ficult to see; posterior lens slightly larger, oval, within a broad, teardrop-shaped, black, naked area immediately before anus, as typical of ge- nus. Black line connecting the two lenses poorly defined. Ventral striae generally as described for others of genus, but not as well defined and much more difficult to see. Pectoral and pelvic fins about on same vertical; first dorsal origin slightly behind that; anal fin begins midway below long interspace between dorsal fins. Pectoral and pelvic fins rather long, both extending slightly past anal origin. Color overall black or swarthy, darkest over trunk and ventral aspects offish; tail mostly dark brown. A broad midlateral band of heavier me- lanophores beginning above midlateral portion of trunk, extending posteriorly onto and even- tually completely including tail. Paired and first dorsal fins with dark rays and pale interradial membranes; anal fin pale but bases of rays black. Floor of buccal cavity below tongue forming a black triangle; tongue, however, pale dorsally with only a few scattered large melanophores on ven- tral surface. Roof of mouth with splotches of black. Size.— To at least 130 mm. Distribution.— Known only from the Nazca and Sala y Gomez ridges. Comparisons and Remarks. — The NSG specimens are similar in many characters to H. aterrimus and H. papyraceus. Using the key pro- vided by Gilbert and Hubbs ( 1 920:520), our four specimens key out to H. aterrimus, and most meristic and morphometric features agree with the original and Marshall and Iwamoto's (1973: 607) descriptions of that species. However, com- parison of the four with Hawaiian paratypes (CAS-SU 8513) of H. aterrimus, four Atlantic representatives (CAS 14514), and others from the Atlantic and Indian oceans deposited in IOAN and ZMMGU suggests that the head is too deep and narrow, and the median nasal ridge is too high to be that species. The poor condition of the specimens, however, might account for the apparent differences. We also compared our ma- terial with a 32.4 mm HL specimen of//, papyra- ceus (ZMMGU 18258) taken in the East China Sea off southern Japan. Among other characters, H. papyraceus differs in having a smaller barbel, fewer pelvic fin rays (11), larger orbit, narrower suborbital, and longer upper jaw. With more and better specimens, it should be possible to better compare and quantify these apparent differences, but for now, it seems best to leave the identity as questionably H. aterrimus. Material Examined. -ZMMGU 17729 (2:33-34.3 mm HL, 1 25-1 3 1 + mm TL) and CAS 75792 (34 HL, 1 34 TL); Sala y Gomez Ridge, 1,070-1,100 m; Prof. Mesiatzev cr. 13, tr. 10. ZMMGU 18128 (26.5 HL, 84+ TL); Nazca Ridge, 340-780 m; Prof. Mesiatzev cr. 13. Hymenocephalus gracilis Gilbert and Hubbs, 1920 (Figure 20a) Hymenocephalus gracilis Gilbert and Hubbs, 1 920:522, fig. 3 1 (Holotype: USNM 78227, Albatross sta. 5292, 162 fms, off Luzon, Philippines). Marshall and Iwamoto 1973:602, fig. 31 (N. Atlantic spec). Parin 1990:16 (listed from Nazca and Sala y Gomez ridges). Hymenogadus gracilis: Okamura 1970a:61, pi. 17, fig. 27 (Ja- pan); 1984:201, 359, fig. 143 (E. China Sea). Diagnostic Description. — Body low, terete, gradually tapering posteriorly; tail laterally com- pressed. Head low, its length 18-22% TL; ridges low and poorly developed. Snout low, rather long, 1 .3-1 .7 in orbit (usually 1 .3-1 .4, relatively short- er in juveniles), pointed and protruding well be- yond mouth. Orbit oval, 29-41% HL, upper margin reaching upper profile of head. Maxillary extends to vertical through posterior margin of orbit. Nasal and medial rostral ridges conspic- uous; suborbital ridge developed, dividing sub- orbital region into upper and lower portions, the former very narrow. Counts and Measurements.— (30 NSG spec.) ID. 11,9-1 1; IP. il5-il8; V. 8 (1 spec, with 8/7); 56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 ;—*■<--'•■• Figure 20. Hymenocephalus species from Sala y Gomez Ridge, collected on the 1 8th cruise of Prof. Shtokman. (a) Hy- menocephalus gracilis, ZMMGU 17740 (22 mm HL), sta. 1941, in 410-385 m, 28 April 1987. (b) Hymenocephalus striatulus, ZMMGU 17715 (27.8 mm HL), sta. 1970, in 540-560 m, 1 May 1987. Photographs by Sergei Dudarev. GR-I (outer/inner) 7-11/3-4 + 14-16 (17-18 total), GR-II 14-18 total. Total length 58+-125 mm; HL 1 1.8-24 mm. Following in percent HL: postrostral 73-82; snout 22.6-27.9; preoral 11.1-17.3; orbit 29.2-40.7; interorbital 14.7-22.6; suborbital 7.5-10.6; post- orbital 28.8-36.5; orbit-preop. 32.6-43.3; upper jaw 42.3-49.6; gill slit 20.8-29.8; barbel 23.7- 32.4; body depth 47-67; light organ 54-70; pre- 1D. 98-1 13; pre-V. 93-108; pre-A. 143-163; V.- A. 49-66; ID. height 47-77 (6 spec.); base ID. 31^15; 1D.-2D. 30-87; IP. 35-56; V. 44-55. Size.— To 130 mm. Distribution.— South China Sea off Luzon, Japan, East China Sea, southeastern Pacific, tropical western North Atlantic, eastern Atlantic off" Morocco, and Indian Ocean off Zanzibar (our unpublished data). Comparisons and Remarks.— This species seems to have a preference for island areas. The capture of three specimens in midwater trawls fished well off bottom suggests occasional forays into the bathypelagic realm. Hymenocephalus tenuis Gilbert and Hubbs, 1917, is a closely re- lated species known only from the immature ho- lotype collected in the Hawaiian Islands. Differ- ences between the two species are slight (see Gilbert and Hubbs 1920:520) and should be re- evaluated when more specimens of//, tenuis be- come available. Material Examined. — (39 spec, 7 sta.) Sala y Gomez Ridge (eastern part): ZMMGU 17735 (20.7 mm HL, 98+ mm TL); 390-385 m; Prof. Mesiatzev cr. 15, tr. 53. ZMMGU 17736 (10.9 HL, 56+ TL); 300-0 m over bot. depth >2,000 m; IKMT; Prof. Shtokman cr. 18, sta. 1887. ZMMGU 17737 (about 6.2 HL, 28 TL); 300-0 m over bot. depth > 2,000 m; IKMT; Prof. Shtokman cr. 18, sta. 1925. ZMMGU 17738 (2: 11.8-16.8 HL, 58+-93+ TL); 380 m; Sigsbee trawl; Prof. Shtokmancr. 18, sta. 1938. ZMMGU 17739 (10:19-23.2 HL, 80+-120 TL) and CAS uncat. (10:1 1.3-23 HL, 64-124 TL); 380 m; Prof. Shtokman cr. 18, sta. 1940. ZMMGU 17740 (8: 18-24 HL, 84-120 TL) and CAS uncat. (5:19-23 HL, 97-1 16 TL); 410-385 m; Prof Shtokman cr. 18, sta. 1941. IOAN uncat. (12.0 HL, 58.5 TL); 200-0 m over bot. depth > 2,000 m; IKMT; Prof. Shtokman cr. 18, sta. 1949. Hymenocephalus neglectissimus new species (Figures 19b, 21) No literature applies to this species. Diagnosis.— Orbits large, greatest diameter about 44-55% HL; interorbital 20.8-27.4%; bar- bel thin, 1 5-28%; GR-I (lower limb, inner series, including raker at angle) 13-17, rarely 18. Body depth over anal origin 47-58% HL. Pectoral rays i 1 2— i 1 4 (rarely il5); V. 8. Black blotch on dor- sum with clear-cut outlines; few isolated or no pigment cells above its posterior projection, few or none behind it, and few small widely spaced dots below (on sides of abdominal region). Counts and Measurements.— (see also Di- agnosis) (data on holotype in square brackets) ID. 11,7-8 (rarely 9) (x = 11,7.9, n = 50) [11,8]; SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 57 Q $ 6 + + t *4f + * A A A + + + + A + • + + °s HEAD LENGTH (mm) Figure 2 1 . Scatter diagram showing relationship of interorbital width to head length in H. semipellucidus (solid circles), H. neglectissimus (squares), H. s. striatissimus (plus signs), H. s. aeger (circled plus signs), and H. hachijoensis (triangles). Data for Hymenocephalus s. aeger from Gilbert and Hubbs (1920), for H. hachijoensis from Okamura (1970a). IP. il2-il4 (rarely il5) (jc = 12.8, n = 92); GR-I (outer) 11-15 (x = 12.9, n = 51) [14]; GR-I (inner) (2-3) + (0-1) + (13-17, rarely 18), total 16-21 (X = 18.2, n = 51) [3 + + 15]; GR-II (outer) (2, rarely 3) + (1, rarely 0) + (13-16), total 16-19 (Jc = 17.4, n = 51) [2 + 1 + 15]; GR-II (inner) (2-3) + (0-1) + (13-15), total 16- 19 {X = 17.1, n = 51) [3 + 1 + 14]; caeca 10- 11 (Jc= 10.3, n = 12). Total length 78 +-1 25 mm; HL 12.5-1 9.2 mm (x = 16.5, n = 29) [18.5]. The following in per- cent of HL: postrostral 77.9-89.6 (Jc = 85.4, n = 28) [86.5]; snout 16.4-23.8 (Jc = 19.6, n = 27) [17.3]; preoral 10.4-18.9 (Jc = 14.8, n = 15) [13.0]; postorbital (greatest length) 37.5-43 (Jc = 40.8, n = 8) [37.8], (least length) 26.7-38.6 (Jc = 33.8, n = 28); suborbital 5.2-9.5 (Jc = 6.9, n = 29) [7.0]; orbit-preop. 28.1-37.3 (Jc = 32.3, n = 28) [28.6]; orbit (max.) 44.8-54.8 (Jc = 49.9, n = 29) [49.7], (horizontal) 44.6^19.7 [49.7]; upper jaw 53.3-58.8 (Jc = 56.1, n = 28) [55.7]; interorbital 20.8-27.4 (Jc = 23.9, w = 26) [2 1.1]; gill slit 31.1- 36.9 (Jc = 33.3, n = 16) [32.4]; barbel 15.3-28.0 (Jc = 21.5, n = 28) [18.9]; pre-A. 146-166 (Jc = 155, n = 28) [162]; pre-V. 92-113 (Jc = 105, n = 28) [109]; V.-A. 47.5-68 (Jc = 587, n = 28) [60.5]; body depth 66-88 (Jc = 77, n = 46) [78], depth over A. origin 47-58 (Jc = 54, n = 24) [55]; light organ 65-89 (Jc = 80, n = 24) [76.8]; height ID. 75-92 (Jc = 83, n = 9); 1D.-2D. 60.1-90.4 (Jc = 72.4, n = 26) [73.5]; length IP. 63-92 (Jc = 77, n = 24) [80]; length V. 71 (n = 1). Description. — Head 6.5-7.6 into TL (21 specimens); greatest body depth usually some- what less than postorbital length of head, the trunk tapering more rapidly to origin of anal fin 58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 than beyond in adults, but tapering gradually in smaller fishes. Head moderately broad, its great- est width notably less than depth of head over midorbit. Orbit circular (rarely greatest diameter oblique); both maximum and horizontal diam- eters of orbit exceed maximum postrostral length of head. Snout short, rather low, bluntly round- ed, not produced beyond anterior tip of pre- maxillary. Dentition consists of small, conical, slightly recurved teeth in 2-4 irregular rows on premax- illary and 1-3 on dentary (one row on short pos- teriormost part of tooth band adjacent to coro- noid process). Luminescent organ as illustrated for H. s. stria- tissimus by Okamura (1970b, fig. 74A), but ex- ternally both lenses seem somewhat larger and lack pigmentation on surface. Alimentary canal short and simple with only 2 bends (see Okamura 1970b, fig. 62A). Pyloric caeca short, 10-11. Scales on all specimens now lost. Fins weak, often broken, especially pelvics. Pectoral and pelvic (in one specimen only) fins slightly produced beyond anal fin origin, extend- ing to about 4th to 8th anal ray. Spinous second ray of first dorsal fin with a short threadlike pro- longation; its overall length about equal to post- rostral length. Second dorsal rudimentary over almost entire length, its origin above 1 3th to 1 6th anal ray. Color superficially resembles that of H. semi- pellucidus (see below) or H. striatissimus (as il- lustrated by Okamura 1984), but with numerous constant differences. Blackish blotch on dorsum darker and with clear-cut outlines (see Fig. 1 9b); its posterior projection rather short with isolated (if any) small pigment cells above, very few be- hind (posterior end of projection appears some- what eroded), and few pigment cells below pro- jection (some in short midlateral series); pigment rarely passes from trunk to tail beyond vertical of 5th to 7th anal fin ray. Tail appears transparent over almost entire length. Dorsalmost 5-9 cer- atobranchial rakers on second gill arch blackish; those more ventral whitish; lateral rakers on first arch all unpigmented. Stomach light to dark gray. Size.— To at least 130 mm. Distribution. — Known only from the Sala y Gomez Ridge in 525-600 m. Etymology. — Latin neglectus (neglected or unnoticed) is used in the superlative, neglectis- simus, to reflect the late discovery of the species. Hymenocephalus semipellucidus and H. neglec- tissimus were first collected together in 1983, but the latter was not recognized as being distinct from the former until 1990. Comparisons and Remarks.— Hymenoceph- alus neglectissimus is very similar to H. semipel- lucidus from NSG and to the species in the H. striatissimus complex from the tropical western Pacific. It is partially sympatric with H. semipel- lucidus, the two having been collected together at four localities bounded by latitudes 25°00'S and 25°30'S and longitudes 88°30'W and 89°00'W. Superficially, the two NSG species of Hymeno- cephalus may be easily and faultlessly separated by their color patterns. Hymenocephalus neglec- tissimus differs from members of//, semipelluci- dus and also the western Pacific H. striatissimus complex in having a more contrasting outline of the dark dorsal blotch, with a rather short pos- terior projection that is somewhat eroded cau- dally and surrounded by few or no isolated pig- ment cells. In contrast, H. semipellucidus and H. striatissimus have a dorsal blotch with eroded outlines surrounded by numerous, relatively large pigment cells and a relatively long posterior pro- jection to the blotch that extends well back (pos- teriorly) onto the tail. Hymenocephalus neglec- tissimus also differs in having unpigmented lateral gill rakers on the first arch. It further differs from H. semipellucidus in having fewer pectoral fin rays (il2-il4 vs. i 14— i 1 8) (Table 3), a somewhat shorter snout, wider interorbital (Fig. 21), nar- rower suborbital, slightly longer upper jaw, and longer barbel, but considerable overlap is seen in these mensural features. Body depth in //. neglectissimus appears to decrease more rapidly posteriorly than it does in H. semipellucidus; the difference between the maximum body depth in, and the depth at anal origin of, 23 specimens varied from 1 1 .2-33.1% (jc = 22.7) in the former, compared with 5.5-27.9% (x = 16.7) in 18 spec- imens of the latter. In this respect, H. neglectis- simus more closely compares with H. striatissi- mus (14-29%, usually 22-26%). The new species differs from H. striatissimus (including subspecies striatissimus, aeger, and torvus, but excluding H. hachijoensis) in having a somewhat shorter snout, larger orbit, shorter postorbital, and narrower interorbital (see Fig. 2 1 ). Gill rakers are also fewer in H. neglectissi- mus, but this character shows considerable over- lap (see Table 3). Hymenocephalus neglectissimus is a smaller SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 59 Table 3. Selected counts of pectoral fin rays and gill rakers of the Hymenocephalus striatissimus complex. Pectoral fin rays 12 13 14 15 16 17 SD H. semipellucidus H. neglectissimus H. striatissimus: sp. Tasman Sea Timor and Coral seas Molucca Sea Sulu Sea S. China Sea Japan 29 13 14 Inner gill rakers, first arch, lower limb 15 16 17 19 SD H. semipellucidus H. neglectissimus H. striatissimus: sp. Tasman Sea Timor and Coral seas Molucca Sea Sulu Sea S. China Sea Japan 15 16 Total gill rakers, second arch 17 19 20 21 22 23 SD 15 16 Total inner gill rakers, second arch 17 19 20 21 22 SD H. semipellucidus H. neglectissimus H. striatissimus: sp. Tasman Sea Timor and Coral seas Molucca Sea Sulu Sea S. China Sea Japan species (maxima at about 130 mm TL and 20 mm HL) than H. semipellucidus (130-160 mm TL and 20-26.5 mm HL) and H. striatissimus (150-180 mm TL, 25-28 mm HL; data from Okamura et al. 1982, Okamura and Kitajima 1984, and our material). It may well be the small- est macrourid species. Material Examined.— (all from Sala y Gomez Ridge) Ho- lotype: ZMMGU 18219 (18.5 mm HL, 123+ mm TL); 580- 564 m; Prof. Shtokman cr. 18, sta. 1964. 60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 22. Hymenocephalus semipellucidus new species; paratype, CAS 75978 (26 mm HL), from Sala y Gomez Ridge in 565-555 m, Prof. Mesiatzev cr. 13, tr. 1, 1 Sept. 1983. Drawn by Tomio Iwamoto. Scale bar equals 10 mm. Paratypes (marked with an asterisk *) and other material (270 specimens from 8 localities). ZMMGU 18220 (30* + 94: 15.5-19.6 HL, 60+-130 TL); data as for holotype. ZMMGU 17724 (55:12.6-18.2 HL, 90-128 TL); 562-545 m; Prof. Shtokman cr. 18, sta. 1965. ZMMGU 17726 (25:14.2-18.5 HL, 85-1 1 5 TL); 545-600 m; Prof Shtokman cr. 1 8, sta. 1977. ZMMGU 18 140 (20* + 20: 12.5-18.7 HL, 78+-125TL); 525- 530 m; Prof. Mesiatzev cr. 15, tr. 49. ZMMGU 18217 (17.5 HL, 99+ TL) and CAS 75976 (16.3 HL, 1 16+ TL); 565-555 m; Prof Mesiatzev cr. 13, tr. 1. ZMMGU 18218 (7:16.5-19 HL, 86+-1 15 TL); 550-560 m; Prof. Mesiatzev cr. 13, tr. 2. ZMMGU 18221 (27:14.2-18.7 HL, 80+-123 TL); 563-590 m; Prof Shtokman cr. 18, sta. 1976. Hymenocephalus semipellucidus new species (Figures 21-23) "Hymenocephalus sp. nova Sazonov et Iwamoto": Parin 1 990: 17 (listed from Sala y Gomez Ridge). Hymenocephalus striatissimus (non Jordan and Gilbert, 1 904): Parin et al. 1990:42 (stomach contents). Kotlyar and Parin 1990:106 (otolith). Diagnosis.— Orbits large, greatest diameter about 43-51% HL; interorbital 16.5-22.1% HL; barbel thin, 1 3-22% HL; GR-I (lower limb, inner series) 14-16 (rarely 13). Body depth over anal fin origin 48-64. Pectoral rays i 1 4— i 1 8 (rarely i 1 9); V. 8. Counts and Measurements. —(holotype data in square brackets) ID. 11,7-9 (x = 8.1, n = 51) [11,8]; IP. i 1 4-i 1 9 (x = H6.5, n = 98) [il7]; GR-I (outer) 9-15 (jc = 11.6, n = 51) [13]; GR-I (inner) (2-4) + (0-1) + (13-15), total 16-20 (jc = 17.9, n = 51) [3 + + 14]; GR-II (outer) (1-4) + (0- 1) + (12-16), total 15-19 (x = 16.7, n = 51) [2 + 1 + 13]; GR-II (inner) (2-3) + (0-1) + (12- 15), total 15-18 (x= 16.5, n = 51) [2 + 1 + 13]; caeca 9-13 (x = 11.4, n= 17). Total length 102+-165 mm; HL 17.7-26.5 mm. The following in percent of HL: postrostral 76.9-86.8 (x = 81.9, n = 25) [79.6]; snout 20.9- 27.0 (jc = 24.6, n = 25) [24.9]; preoral 1 1.8-17.4 (x = 14.1, n = 21) [15.1]; orbit max. 42.9-50.5 (jc = 45.7, n = 26) [49.0]; orbit horiz. 37.0-48.2; interorbital 16.5-22.1 {x = 19.6, n = 25) [20.4]; postorbital (greatest) 38.3-45.1 (x = 42.0, n = 24) [40.8]; postorbital (least) 33. 1-44.9 (x = 37.9, n = 24) [37.1]; suborbital 8.5-1 1.8; orbit-preop. 30.2-39.6 (jc = 35.5, n = 25) [35.9]; upper jaw 49.2-56.8 (jc = 52.6, n = 25) [52.2]; gill slit 26.7- 34.5 (jc = 31.4, n = 16) [31.4]; barbel 13.0-22.0 (jc = 17.7, n = 26) [15.5]; body depth (max.) 61- 85 (jc = 72, n = 26) [66]; depth over A. orig. 48.2-63.9 (jc= 54.9, n= 18) [53.1]; pre-A. 152- 173 (jc = 162, n = 26) [165]; pre-V. 97-111 (jc = 103, n = 26) [97]; V.-A. 53.6-81.8 (jc = 69.0, n = 26) [80.8]; height ID. 66-96 (jc = 77, n = 17) [78]; 1D.-2D. 67.6-88.9 (jc = 77.5, n = 24) [71.8]; IP. 58-78 (jc = 70, n = 25) [76.7]; V. 62- 78 (jc = 71, n = 22) [67]; light organ 71.9-92.6 (jc= 80.8, n= 18) [77.6]. Description. — Head 6-7 into TL; greatest body depth usually somewhat less than postros- tral length of head, the trunk tapering gradually to and beyond origin of anal fin. Head rather broad, its greatest width slightly less than depth of head over midorbit. Greatest diameter of orbit oblique and about equal to or somewhat more than postorbital length of head (taken from orbit margin to uppermost posterior angle of opercle). Snout rather low, bluntly rounded, not produced beyond anterior tip of premaxillary. Dentition consists of small, conical, slightly recurved teeth in 1-3 irregular rows in both jaws in most specimens (a few individuals have pre- maxillary bands 4 or 5 teeth wide). Luminescent organ as illustrated for H. s. stria- tissimus by Okamura (1970b, fig. 74A). Alimen- tary canal short and simple, with only 2 bends (see Okamura 1970b, fig. 62A). Pyloric caeca short, thick; 9-13 in 17 specimens. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 61 Few scales remain, almost all of these lacking spinules. A few isolated scales with short conical spinules as illustrated for genus by Okamura (1970b, fig. 2). Fins rather weak, tips of first dorsal broken off in all specimens; pectoral and pelvic fins extend posteriad to level of 4th to 9th anal fin ray. Sec- ond dorsal rudimentary over almost entire length, its origin above 1 4th to 1 6th anal ray. Color description given by Gilbert and Hubbs (1920:533-534) for H. striatissimus aeger accu- rate for new species as well— this and other char- acters suggest close affinity of the two species. Black blotch on dorsum with eroded outlines, surrounded by large pigment cells with numerous branched projections; posterior projection of blotch variable in length (extends either to level of anal-fin origin or to vertical of origin of 13th- 1 8th anal ray). Pigment cells surrounding blotch numerous below and behind, fewer (rarely ab- sent) above. Arrangement of these cells varies greatly: from irregularly scattered to grouped along myosepta (more densely along those going dorsad and posteriad above lateral line, and ven- trad and anteriad below blotch). Some cells may be arranged in a line going posteriad along mid- lateral myosepta. Dark pigment attains level of 9th to 25th anal ray. Dorsalmost 9 or so cera- tobranchial rakers on second gill arch blackish, lateral rakers on first arch also blackish (some- times pale on anteriormost few); those more ven- tral pale. Stomach blackish. Hymenocephalus semipellucidus and Russian specimens from Japan of//, striatissimus do not show the silvery sheen and black striations as prominently as in Albatross specimens of latter species, but differences probably an artifact of preservation. Alcohol-fixed specimens retain sil- very appearance, whereas those fixed in formalin lose silvery reflections. Thick translucent skin covers abdominal region of NSG specimens, but careful teasing and stripping away of epidermis reveals striations underneath. Black melano- phore pattern over body of new species promi- nent (see Fig. 22). Similar dark pigment patterns may have been present in H. striatissimus spec- imens examined, but specimens currently faded and show only faint traces of such patterns. Food. — Parin et al. (1990:42) found the spe- cies to feed predominantly on copepods (includ- ing Pleuromamma sp., Gaussia scotti, Phyllopus mutatus, Chirundina streetsi, Arietellus sp., and Aetideidae sp.). Benthic polychaets (including Polinoidae) and the gonostomatid fish Cyclo- thone were also important. Size.— To at least 165 mm. Distribution. — Known only from the Sala y Gomez Ridge in 550-800 m. Etymology. — From Latin semi, half, and pel- lucidus, clear or transparent, in reference to the partially transparent head covering, and the translucent caudal region where the vertebra can be seen in fresh specimens. Comparisons and Remarks.— The new spe- cies keys out to H. striatissimus using Gilbert and Hubbs's (1920:520) key, but it differs from that species in a number of important features. The gill-raker counts, notably the inner series in the first (outermost) arch, show the best sepa- ration of the two species (see Table 3) in our material. Note from the table that gill raker counts in our H. striatissimus specimens suggest a clinal difference in populations of that species, with the Molucca Sea specimens, representing H. s. aeger, completely separated from the typical subspecies H. s. striatissimus of Japan, and the Sulu Sea and South China Sea specimens intermediate in their counts. South China Sea specimens were consid- ered by Gilbert and Hubbs (1920:531) to be an intergrade between //. 5. torvus and H. s. stria- tissimus. In gill-raker counts, ZMMGU specimens of//. striatissimus from the Coral and Timor seas ap- pear closest to H. striatissimus specimens from the Sulu Sea. A small specimen (ZMMGU 1 8260) from the Tasman Sea, however, is almost indis- tinguishable from H. semipellucidus and H. ne- glectissimus, differing only in having a distinct color pattern similar to that of H. striatissimus and slightly different morphometry from that of H. semipellucidus. Pectoral ray counts in the new species tend to be higher than those of//, striatissimus, but there is considerable overlap in this feature (Table 3). The suborbital region appeared to be narrower in the new species, and a scatter diagram plotting measurements of the least width against the greatest diameter of the orbit (Fig. 23) showed a good separation between the Malay Archipelago specimens of H. striatissimus on the one hand, and Japan specimens of that specimens, and //. semipellucidus specimens on the other hand. The separation of the Japanese specimens of//, stria- tissimus from other specimens of that species lends additional support for continued recogni- tion of subspecies. A more extensive study may 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 4 5 6 7 ORBIT DIAMETER (mm) Figure 23. Scatter diagram showing relationship of suborbital width to orbit diameter in Hymenocephalus semipellucidus (circles), H. s. striatissimus from Japan (triangles), and H. striatissimus aeger from the Malay Archipelago (plus signs). necessitate elevation of two or more of the sub- species to species level. The interorbital width is distinctly narrower in H. semipellucidus, ranging 16.5-22% HL, as compared with about 28% HL in Japanese spec- imens of//, striatissimus, and 28-39% in Malay Archipelago specimens. Although there is con- siderable overlap in this character between H. semipellucidus and H. neglectissimus, the scatter diagram (Fig. 21) comparing the interorbital width of five populations of Hymenocephalus shows a distinct separation between the two spe- cies. The new species also appears to have a more gradually tapered body than does H. striatissi- mus. This is best reflected in a comparison of the body depth below the origin of the first dorsal fin (greatest body depth) and the depth over the anal fin origin. In the new species, the first mea- surement ranged about 61-85% HL, the second 48-64% HL, with a difference in each specimen of 5.5-27.9% between the two proportions. In contrast, the H. striatissimus specimens had body depths of 55-81% HL, and 37-56% HL, with a difference of 14-29% (most examples at 22-26%). Differences between H. semipellucidus and H. neglectissimus were discussed under the descrip- tion of the latter species. Hymenocephalus semi- pellucidus differs from the three subspecies of//. striatissimus (//. 5. stratissimus, H. s. aeger, and //. 5. torvus) more than each differs one from another. The fourth subspecies, //. 5. hachijoen- sis, from Japanese waters, may represent a sep- arate species intermediate (as Okamura 1970a suggested) between H. striatissimus and H. lon- giceps. We have examined three specimens of//. s. hachijoensis (ZMMGU 18243 and 18244 from the North-West (=Emperor) Ridge and ZMMGU uncat. from the Kyushu-Palau Ridge) and found that they differ from the three other subspecies of//, striatissimus (as well as the two NSG spe- cies) in color pattern and barbel length, which would seem sufficient to elevate H. hachijoensis to species level. Material Examined.— (103 spec, from 7 sta.) Sala y Gomez Ridge: Holotype: ZMMGU 18129 (24.5 mm HL, 152 mm TL); 750 m; Prof. Shtokman cr. 18, sta. 2019. Paratypes(40 spec): ZMMGU 17723 (7:20.0-24.8 HL, 138- 1 62 TL); 580-564 m; Prof. Shtokman cr. 1 8, sta. 1 964. ZMMGU 17725 (14:17.7-23.3 HL, 106+-150 TL); 563-590 m; Prof. Shtokman cr. 18, sta. 1976. ZMMGU 17728(3:22.0-22.8 HL, 1 04 +-1 38 TL); data as for holotype. ZMMGU 17721 (4:19.5- 23 HL, 91+- 149 TL) and CAS 75978 (2:20.0-26.5 HL, 1 37- 165 TL); 565-555 m; Prof. Mesiatzev cr. 13, tr. 1. ZMMGU 17722 (6:18-21 HL, 105+-132+ TL) and CAS 75977 (4:22- 23 HL, 108+-142 + TL); 550-560 m; Prof. Mesiatzev cr. 13, tr. 2. Other material: ZMMGU 17727 (53:19.5-26.5 HL, 96+- 1 64 TL); 730-790 m; Prof. Shtokman cr. 1 8, sta. 20 1 8. ZMMGU 18205 (9:22.5-26 HL, 121-184 TL); 750-800 m; Prof. Shtok- man cr. 18, sta. 1996. Specimens of Hymenocephalus striatissimus used for com- parisons. -Coral Sea: ZMMGU 18259 (2:23.7-26.5 HL, 125+- 147 TL); 18°48'S, 149°58'E; 660 m; Lyra tr. 20; 14 Jun. 1968. Timor Sea: ZMMGU uncat. (3: ca. 22-25.2 HL, 110+-146 TL); 9°00'S, 130°38.8'E; 445-520 m; Akademik Berg tr. 553. Tasman Sea: ZMMGU 18260 (questionably identified as H. striatissimus) (1 5. 1 HL, 97.5 TL), 34°17.8'S, 171°30.9'E; 670- 630 m; Dmitry Mendeleev cr. 16, sta. 1265; 5 Jan. 1976. JA- PAN. CAS-SU 8549 (paratype of//, striatissimus: 25 mm HL, 1 10+ mm TL); Suruga Bay; Albatross (no other data). Molucca Sea: CAS-SU 25463 (8 paratypesof H. s. aeger: 1 1.2-26.5 HL, 69-1 46 TL);00°15'N, 127°24'36"E: 545 m; Albatross sta. 5621; 28 Nov. 1909. CAS 57180 (3:25.1-28.1 HL, 160-178 TL); Suruga Bay, off Heda [Heta]; shrimp trawl; 18 Feb. 1969. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 63 PHILIPPINES. CAS-SU 25620 (18.8 HL, 1 1 1 TL); near Jolo, 6°02'55"N, 120°53'E; 186 fms [340 m]; Albatross sta. 5173, 5 Mar. 1908. CAS uncat. (14.7 HL, 93 TL); off n. Luzon, 18°29'45"N, 121°39'E; 150 fms [274 m]; Albatross sta. 5328, 19 Nov. 1908. CAS-SU 25464 (9.9-13.5 HL, 50-88 TL); be- tween Jolo and Tawi Tawi, 5°48T2"N, 120°30'48"E; 224 fms [410 m]; Albatross sta. 5563, 21 Sep. 1909. Specimens of H. hachijoensis used for comparisons: Em- peror Seamounts: ZMMGU 18243 (32.7 HL, 205 TL); ca. 32°N, 173°E; depth unknown; Mys Vnony tr. 86; Sep. 1979. ZMMGU 18244 (26.2 HL, 163+ TL); 41°04'N, 170°32'E; 1,050-1,060 m; Mys Vnony tr. 103; 22 Sep. 1979. Kyushu- Palau Ridge. ZMMGU uncat. (ca. 29.5 HL, 160+ TL); no exact data on catch; Prof. Deryugin; 1971. Hymenocephalus striatulus Gilbert, 1905 (Figure 20b) Hymenocephalus striatulus Gilbert, 1905:665-666, fig. 259 (type-locality Hawaii, off SW coast Oahu; 192-352 fathoms [351-644 m]). Parin 1990:16 (listed from Sala y Gomez Ridge). Parin et al. 1990:41—42 (stomach contents). Kotlyar and Parin 1990:104, fig. 3d (otolith). Hymenocephalus sp.: Parin et al. 1981:11-12 (67 specimens from Sala y Gomez Ridge). Diagnosis.— A species of subgenus Hymeno- cephalus, with snout rather low, pointed, pro- jecting beyond mouth, about as long as inter- orbital or longer; orbit diameter 35-43% HL, much greater than interorbital width; suborbital narrow, 9-15% HL; barbel rudimentary (1.5 or more into least suborbital width) or obsolete. Pectoral rays il4-i20; V. 14-15. Counts and Measurements.— (from more than 100 Sala y Gomez specimens; mean values in square brackets) ID. 11,8-11 [jc = 11,9.5; IP. il4-i20 [il6.9]; V. 14-15 [14.5; GR-I [outer/in- ner] 17-23 [20.2]/(4-6) + (21-25) [5.0 + 1 + 22.5], GR-II (3-5) + 1 + (20-24) [4.4 + 1 + 22.0]/(3-5) + 1 + (19-23) [4.2 + 1 + 20.9]. Total length 103-180; HL 28.5-34.5 mm. The following in percent of HL: postrostral 73.3-8 1 .2 [76.9]; snout 23.2-30.5 [27.5; orbit 34.7^2.8 [39.1]; interorbital 21.2-29 [24.9]; postorbital 33- 42 [37.0]; orbit-preop. 34.2-40.6 [37.2]; subor- bital 9.1-14.9 [10.9]; upper jaw 47.2-58.8 [51.6]; barbel 3.1-12; gill slit 29.2-37 [33.3]; pre-D. 95- 106 [102]; pre- A. 138-152 [146]; pre-V. 95-1 13 [102]; V.-A. 43.8-59.1 [50.8]; ID. base 28.6- 36.8 [32.3]; ID. height 52-75 [63]; 1D.-2D. 35.3- 73.4 [55.2]; IP. 48-61 [53]; V. 55-84 [70]; light organ 51.5-62.5 [56.7]; body depth 58-70 [65]. Description.— Body slender, subcylindrical, greatest width over pectoral bases about three- fourths greatest depth; the trunk tapering grad- ually posteriorly to tail tip. Head low and broad, greatest width about equal to or more than its depth. Head bones somewhat stouter than most others of genus, and mucous cavities less devel- oped—these conditions somewhat intermediate between those of subgenus Hymenogadus and Hymenocephalus. Head covering transparent. Orbit large, its greatest diameter on a diagonal and slightly shorter than postorbital length. In- terorbital space moderately broad, although sharply narrowed forward of midorbit level. Pre- opercle ridges form a large triangular process at postero ventral corner. Snout moderately pointed and protruding beyond large mouth. Upper jaws extend posteriorly to hind edge of orbit. Teeth in moderately wide bands in both jaws, the individual teeth uniformly short and stoutly conical, with rather blunt tips. Small round anterior lens of luminescent organ in middle of chest, but difficult to discern; pos- terior lens slightly larger, oval, within a broad teardrop-shaped black naked area immediately before anus, as typical for genus. Black line con- necting the two lenses poorly defined. Ventral striae not especially prominent and less extensive than in H. striatissimus. Pyloric caeca 11 or 12, short (less than orbit diameter or interorbital width) and thick. Two slender retia terminate in 2 flattened half-moon- shaped gas glands. First dorsal fin long-based, its height some- what less than postrostral length; second dorsal scarcely developed except near tail tip; pectoral fins slender, extending to about level of anal or- igin; pelvic fins moderate in size, the outermost ray slightly prolonged, extending to about 5 th to 8th anal ray. Color in alcohol overall grayish or blackish except over transparent head covering. Abdo- men from base of pectoral fins ventrally to pel- vies and back to vent dark violet to blue; chest and bases of paired fins black. Scattered mela- nophores of different sizes cover all of trunk and tail, the melanophores generally larger and more widely spaced on ventral half of body. A broad, dense midlateral band of melanophores begins about midlateral portion of trunk and extends posteriorly onto, and eventually completely in- cludes, tail. Anterior edge of nape and leading edge of snout black. Large scattered melano- phores cover most of suborbital and lower por- tion of preopercle, all of opercle, and parietal region. Jaws anteriorly, and gular and branchios- tegal membranes black. Floor of mouth below tongue forming a black triangle; tongue, how- 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 ever, pale dorsally with only a few scattered large melanophores on ventral surface; roof of mouth with splotches of black. Gill filaments pale; arch- es and rakers lightly peppered with small mela- nophores or pale. Paired and first dorsal fins with dark rays and pale interradial membranes; anal fin pale, but bases marked by small black dots. FooD.-Parin et al. (1990:42) found H. stria- tulus to feed predominantly on pelagic organ- isms, especially copepods (Pleuromamma sp., Oncaea conifer a, Xanthocalanus sp., Aetideidae spp.), but also small fish, chaetognaths, gam- marids (Lysianassidae), shrimp (Bentheogenne- ma pasithea), and mysids (Paralophogaster gla- ber). Benthic polychaets (including the family Polinoidae) were also important food items. Size. —A small species, attaining about 1 80 mm TL. Distribution. — Known only from the Ha- waiian Islands and the Sala y Gomez Ridge in depths of about 350 to 640 m. Comparisons and Remarks. —Gilbert's ( 1 905) original description and illustration of the species are excellent; they should be referred to for ad- ditional details. Specimens from the Sala y Go- mez Ridge show one notable difference from those collected off Hawaii; they have a rudimentary but distinct mental barbel, the length of which goes about 1.5 times into the least suborbital width, in contrast to the almost obsolete barbel in Hawaiian specimens. We do not feel this single difference is sufficient to recognize the species as distinct from H. striatulus, especially knowing that barbel length in species of Hymenocephalus can vary widely. A notable example is the wide- ranging species H. striatissimus of Japan, South China Sea, Philippines, and East Indies. Gilbert and Hubbs (1920:527) recognized three subspe- cies of//, striatissimus, and Okamura (1970a: 50) described a fourth, based in part on geo- graphical differences in barbel development. Per- haps the Sala y Gomez specimens should also be recognized as a subspecies of the Hawaiian H. striatulus, but we choose not to recognize them as such at this time. Gilbert and Hubbs's ( 1 920:52 1) key to the sub- genera and species of Hymenocephalus can be used to distinguish H. striatulus from all other known species of the genus except H. billsamo- rum Marshall and Iwamoto, 1973, from the trop- ical western Atlantic. The two species share in common a relatively slender body, low pointed snout, high pelvic fin ray counts (13-15), and a rudimentary or obsolescent barbel. Hymeno- cephalus striatulus differs from H. billsamorum in having larger orbits, narrower suborbital and interorbital regions, and slightly more pectoral fin rays (i 1 6— i 1 8 vs. il4-il5). Material Examined. —Sala y Gomez Ridge: ZMMGU 17711 (33 spec.:24-34.5 mm HL, 117+-150+ mm TL); 540 m; Ichthyandrcx. 5, tr. 53. ZMMGU 17712 (34:25-37 HL; 1 12+- 180+ TL); 345-540 m; Ichthyandr cr. 5, tr. 55. ZMMGU 1 77 1 3 (275: 1 7.8-34 HL, 90-1 70 TL); 580-564 m; Prof. Shtok- man cr. 18, sta. 1964. ZMMGU 17714 (185:19.2-36.5 HL, 91-164 TL); 562-545 m; Prof. Shtokman cr. 18, sta. 1965. ZMMGU 17715 (2:27.8-29.3 HL, 116+-U8+ TL); 540-560 m; Prof. Shtokman cr. 18, sta. 1970. ZMMGU 17716(4:23.5- 30.8 HL, 80+-144 TL); 570-580 m; Prof. Shtokman cr. 18, sta. 1971. ZMMGU 17717 (37:18.8-32.5 HL, 95-154 TL); 563-590 m\ Prof. Shtokman ct. 18, sta. 1976. ZMMGU 17718 (100:15.3-33.5 HL, 75-135 TL); 545-800 m; Prof. Shtokman cr. 18, sta. 1977. ZMMGU 17719 (3:30.6-32.2 HL, 125-142 TL); 750-800 m; Prof. Shtokman cr. 18, sta. 1996. ZMMGU 17720 (24:16-35 HL, 62.5+-165 TL); 730-790 m; Prof. Shtokman cr. 18, sta. 2018. ZMMGU 18212 (7:28-34.5 HL, 103+-180 TL); 565-555 m; Prof. Mesiatzev cr. 13, tr. 1. ZMMGU 18213 (10:29-34 HL, 128+-185 TL); 550-560 m; Prof. Mesiatzev cr. 13, tr. 2. ZMMGU 18214 (2:30.7-32 HL, 122+-141 TL); 410-420 m; Prof. Mesiatzev cr. 13, tr. 4. ZMMGU 18215 (13:20-32.3 HL, 100-158 TL); 530 m; Prof. Mesiatzev cr. 15, tr. 49. ZMMGU 18216 (2:28.9-32.5 HL, 125-134+ TL); 400 m; Prof. Mesiatzev cr. 15, tr. 52. Kuronezumia Iwamoto Kuronezumia Iwamoto, 1974 [as subgenus of Nezumia Jordan, 1904]. Type species: Nezumia (Kuronezumia) bubonis Iwamoto, 1974. Diagnosis. — Macrourines with 7 branchios- tegal rays. Body and head compressed and deep, greatest depth below origin of first dorsal fin about 90-1 10% HL. Snout rounded in profile, almost entirely covered (except narrow median ventral and ventralmost margin) with small uniform, finely spinulated scales. Suborbital region ver- tical, without an angular midlateral ridge, the region covered with small scales without a row of enlarged, scutelike scales. Mouth moderate in size, upper jaw extends posteriorly to below an- terior half of orbit, length 30-44% HL. Dentition in broad villiform bands in both jaws; lower jaw band broad and short; outer series on premax- illary slightly enlarged. Gill rakers 8-1 1 (total) on outer side of second arch. Small patches of scales on branchiostegal membranes in most spe- cies. Body scales rather small, adherent, densely covered with long slender spinules; transverse ridges in most adults (but variously absent in some). Anus far removed from anal fin, closer to pelvic bases. Anterior dermal window of light SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 65 Figure 24. Kuronezumia pallida new species; holotype, ZMMGU 17730, 98.5 mm HL, from Sala y Gomez Ridge in 550 m. Photograph by Susan Middleton. organ usually small, situated between pelvic fin bases and separated from anus by a broad scaly area, which is greatly swollen in two species. Py- loric caeca about 35^0 except K. leonis, which has 15-18. Color overall light gray or brown to swarthy, fins dusky to blackish, naked mem- branes blackish to dark gray. Abdominal verte- brae 12-13. Remarks.— The genus, here elevated from subgeneric status, includes K. bubonis, K. pallida, K. dara (Gilbert and Hubbs, 1916), K. leonis (Barnard, 1 925), K. macronema (Smith and Rad- cliffe, 1912), and two undescribed species. The definition of the genus has had to be expanded to accommodate K. leonis and one of the un- described species. The peculiarly enlarged, bul- bous swelling housing the light organ in K. bu- bonis was used to diagnose the subgenus Kuronezumia, but K. pallida and K. dara lack this swelling. The combination of other features, especially the head physiognomy and squama- tion, nonetheless serve to unite the seven species and justify recognizing them as representatives of a distinct taxon. The genus is now known from the tropical western Atlantic, throughout the Pa- cific (NSG, Hawaii, Japan, off southeastern Aus- tralia [AMS specimens provided by J. R. Pax- ton], off New Zealand [specimen of K. leonis examined in LACM and NMNZ, and K. bubonis in NMNZ], and the Indian Ocean (Madagascar Ridge, West Australian Ridge [Shcherbachev 1987]; Kerguelen Plateau [one juv. of K. leonis, ZMMGU uncat.]). Kuronezumia pallida new species (Figures 24, 25a) Nezumia sp.: Parin et al. 1981:12 (brief descr. of specimens here described as new). "Kuronezumia sp. nova Sazonov et Iwamoto": Parin 1990:16 (listed from Sala y Gomez Ridge). Diagnosis.— No large pores of sensory canals on head. Length upper jaw 34-42 HL, smaller in juveniles. GR-I (inner) 8-9 total; scales below ID. 1 1-14; V. 1 1. No bulblike swelling of light organ; anterior dermal window represented by a small, lenticular scaleless area between midbases of pelvic fins. Color pale brown, except bran- chiostegal membranes and fins darker. Counts and Measurements.— (data for ho- lotype followed in parentheses by 79 mm para- type and 25 mm juvenile) ID. 11,1 1 (11,13, 11,1 1); 2D. about 140+ ; IP. i24/i25 (i25/i24, i25/i27); V. 11/11 (11/11, 11/11); GR-I [outer/inner] 7 (8, 7)/0 + 9 (0 + 8, 1 + 8), GR-II + 7(1 + 8, 1 + 9)/0 + 9, (0 + 9, 1 + 9); scales below 1 D. 1 1 [about 1 4 on right side] (12-13,12), below 66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 2D. 1 1 [about 14 on right side] (11,12). Abdom- inal vertebrae 12; anal pterygiophores anterior to first haemal spine 12. The following in percent of HL: postrostral 74. 1 (73.4, 64.0); snout 28.4 (28.5, 28.8); preoral 17.3 (15.8, 17.6); orbit 23.9 (25.3, 34.8); inter- orbital 24.4 (26.6, 25.2); orbit-preop. 46.3 (42.4, 34.8); suborbital 19.8(20.2, 18.0); upper jaw 40.1 (42.4, about 34); barbel 26.4 (23.4, 22.8); gill slit 17.8 (17, -); body depth about 95 (94, 86); pre- D. 120 (110, 111); pre-V. 104 (104, 94); pre-A. 144 (137, 131); pre-vent 119 (114, 114); V.-A. 46.3 (38.6, 37.2); ID. height-(82, 101); ID. base 28.9 (29.1, 31.2); 1D.-2D. 35.0 (38.6, 29.2); IP. -(61,68); V.-(73, 101). Description.— A large, deep-bodied species, greatest depth about equal to HL in adults, about 5.5 in TL; head deep, relatively compressed. Snout high, blunt, with vertical anterior profile. Orbits moderate in size, circular, diameter less than snout length, about equal to interorbital width. Mouth large, jaws subterminal; posterior end of maxillary extends to vertical through mid- dle of orbit. Head ridges inconspicuous; infra- orbital region deep, vertical, without modified scutelike scales, about 1 3 scales wide, 6 or 7 in upper portion. Scaled areas of opercle and sub- opercle together form a deep, inverted triangle. Interopercle narrowly exposed and scaled along ventral and posterior margins. Free margin of preopercle smooth. Gill membranes broadly united (at level of hind border of orbits), almost without free margin behind their connection with isthmus. Mental barbel moderately thick, long, about equal to orbit diameter, tapered to a fine tip. Sensory canals on head not broadened or swol- len; no enlarged open pores. Free neuromasts serially arranged along surfaces of supraorbital, infraorbital, postorbital, and mandibular canals, and on anterior surface of snout. Olfactory cavity rather small (about diameter of pupil), nasal ro- sette occupies small portion of cavity; anterior nostril rounded, much smaller than posterior nostril, which is semi-elliptical and about 2.1 into least suborbital width; internarial mem- brane narrow, with flap length equal to diameter of anterior nostril. Premaxillary teeth in broad, abruptly tapered bands that fall short of rictus (about 5 rows wide anteriorly); outermost series notably enlarged, straight, conical. Dentary teeth band short, broad, 3-5 rows wide; all teeth uniformly small. Lips, interdental spaces, and adjacent portions of oral cavity covered with numerous, branched (at tips), hairlike papillae that almost cover teeth. Similar papillae on anteriormost portion of snout above upper jaw. Head scales densely covered with erect to sub- erect needlelike spinules in 2-1 divergent series; those bordering orbits with 1-2 prolonged crests radiating from orbit (most pronounced on sub- orbital). Spinules on scales more densely placed in paratype, whereas in holotype spinulated sur- faces of adjacent scales separated from each other by smooth interspaces. No scutelike scales at tip of snout and anterior end of nasal bones; scales around tip scarcely stouter than more peripheral ones. Head almost completely scaled except over gular membranes, anterior end of mandible, and along lower part of snout and suborbital im- mediately above upper lip. Scales present over lowermost branchiostegal rays in two patches on each side in holotype; upper patch much smaller than lower one. Body scales (Fig. 25a) small, densely covered with conical retrose spinules forming 7-15 radiating series. Transverse ridges (reticulate pattern) absent in holotype, but this may be size related— in a 56 mm HL paratype ofK. bubonis (CAS 27874), transverse ridges were sparse but present around the focus (Fig. 25b). Scales cover proximal part of pel vies between fin rays. Light organ represented by black, scaleless, rather small anterior dermal window situated be- tween midbases of pelvics and separated from periproct by convex scale-covered area. Peri- proct rather narrow, preceded by posterior der- mal window of size about equal to anterior win- dow. Internally, light organ consists of large bulblike black luminous gland separated from body cavity by peritoneum, without muscular tissue over upper surface; smaller reflector situ- ated between luminous gland and pelvic bones immediately above anterior dermal window. In- ner layer of skin between both dermal windows uniform in consistency, becoming thinner ante- riorly, blackish with slight opalescence, without indication of separate lenses. Swim bladder moderately large, oval, with blunt end anteriad. Two gas glands well devel- oped, connected to two relatively long retia. Stomach everted in paratype; number of pyloric caeca not determinable. Origin of first dorsal fin slightly behind, origin of pelvic fins well before, vertical through pec- SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 67 toral origins. Spinous second ray of first dorsal scarcely extended beyond longest branched rays; serrations on leading edge relatively small and low. Interspace between dorsals rather short, about one-third longer than base of first dorsal. Pectoral fin moderately long, originating well be- low level of top of gill opening. Pelvic fin with long filamentous outer ray extending to about 8th- 10th anal ray. Origin of anal fin somewhat before vertical of posterior end of first dorsal. Color in alcohol: body and head grayish with yellow tinge. Gular membrane light gray, bran- chiostegal membrane blackish. Fins dark gray to blackish, but anal fin blackish on anterior portion only. Oral and branchial cavities unpigmented. Size.— To about 55 cm TL. Distribution. — Known only from the Sala y Gomez Ridge in 540-800 m. Etymology. — From the Latin pallida, light or pale, in reference to the pale coloration of the species, contrasted with the dark-colored K. bu- bonis. Comparisons and Remarks.— The new spe- cies appears most closely related to K. dara (Gil- bert and Hubbs, 1916) from Japanese waters. Previous to Okamura's report (in Okamura and Kitajima 1984:217, 363, fig. 153) of two speci- mens (220-318 mm TL) from the Okinawa Trough, the three known specimens of that spe- cies were too small to make comparisons with other species of Kuronezumia. Okamura's de- scription and excellent color figure of the larger Okinawa specimen show the close similarity of that species to other members of the genus. The similarities lie not just in the physiognomy of the head, but also in the structure of the fins, the squamation, and the overall color. Kuronezumia pallida appears to differ from K. dara in having smaller orbits (24-25% HL cf. 25-36%), longer upper jaw (34-42% HL cf. 30-35%), interorbital space about equal to or slightly more than orbit diameter (cf. about equal to or less than orbit diameter), and barbel about equal to orbit (cf. much shorter than orbit, at least in small K. dara; condition not stated for large specimen by Oka- mura). The 171 mm juvenile (ZMMGU 18066) in our material agrees closely in these propor- tional measurements with K. dara. Allometric growth probably accounts for the differences be- tween juveniles and adults of the species. It thus seems that juveniles of K. dara and K. pallida are indistinguishable by these measurements. The new species differs from K. bubonis and an un- Figure 25. Scales from below interspace between dorsal fins of (a) Kuronezumia pallida (holotype, ZMMGU 17730, 98.5 mm HL) and (b) K. bubonis (paratype, CAS 27874, 56 mm HL). Scale bars represent 0.5 mm. Drawn by Tomio Iwa- moto. described species from the South China Sea (see Iwamoto 1974) primarily in lacking the bulbous swelling before the periproct and in having a paler coloration. Material Examined.— Sala y Gomez Ridge: Holotype: ZMMGU 17730 (98.5 mm HL, 548+ mm TL); 550 m; As- tronomer sta. 104. Paratype: ZMMGU 17731 (79 mm HL, 379+ mm TL); 540 m; Ichthyandr cr. 5, tr. 53. Non-type material: ZMMGU 18066 (25 HL, 171 TL); 750- 800 m; Prof. Shtokman cr. 16, sta. 1996. 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Malacocephalus Giinther, 1862 Type species: Macrourus laevis Lowe, 1843, by monotypy. Diagnosis. — Macrourines with 7 branchios- tegal rays. Mouth large, usually greater than 45% HL; GR-I (inner series) usually less than 1 2 total. Teeth large, widely spaced, in 1 row in lower jaw, usually larger posteriorly; in 2 rows to narrow band in premaxillary. Head completely and uni- formly scaled, lacking enlarged, modified scales; suborbital area vertical or nearly so and covered with small, finely spinulated scales; lowermost branchiostegal rays scaled; scales without trans- verse ridges. Anus removed from anal fin and closer to pelvic fin, preceded by 2 naked fossae (dermal windows of light organ), one round to bean-shaped fossa between pelvic fin bases, the other immediately before anus at anterior end of periproct region. Pyloric caeca 50-100, multiply branched. Distribution.— Worldwide in warm to tem- perate seas. Remarks. —The genus is divided into two sub- genera by some authorities: subgenus Pawnurus Parr, 1 946 (two species, with serrated first dorsal fin ray) and subgenus Malacocephalus (two to four species, lacking serrated first dorsal ray). Malacocephalus boretzi Sazonov, 1985, de- scribed from the central North Pacific, has char- acters that lessen the distinction between the two nominal subgenera. Sazonov (1985:17), how- ever, supported their continued recognition and also provided evidence (in the internal structure of the light organ) that showed a close relation- ship of Lucigadus Gilbert and Hubbs, 1920 (which he elevated to full genus) to Malacocepha- lus. A question still remains whether or not there is more than one species in the Malacocephalus laevis complex (see Remarks in following de- scription). Malacocephalus laevis (Lowe, 1843) Macrourus laevis Lowe, 1843:92 (off Madeira). Malacocephalus laevis: Giinther 1862:397. Parin et al. 1981: 12 (Sala y Gomez Ridge). GolovarT and Pakhorukov 1987: 73 (Nazca and Sala y Gomez ridges). Parin 1990:16 (Sala y Gomez Ridge). Parin et al. 1 990:43 (stomach contents). Kot- lyar and Parin 1990:106 (otolith). Macrurus (Malacocephalus) laevis: Giinther 1887:148. Diagnosis. — Upper jaw 45-54% HL. Premax- illary teeth in 2 distinct rows, mandibular teeth in 1 row. A smooth spinous second ray of first dorsal; fin rays V. 9 (occasionally 8 or 10); IP. il6-i21. Remarks.— This widespread species has been more than adequately described and illustrated by numerous recent authors (e.g., Marshall 1973; Okamura 1970a), and the reader is referred to other sources for a complete description. Iwa- moto (1979) reported a single individual from off southern California, the first record of the species from the eastern Pacific. Subsequent to that report, numerous specimens were collected in 1979 on seamounts off the Baja California peninsula by the Japanese fishery research vessel Kaiyo Maru (Eichii Fujii, Tokai Regional Fish- ery Research Laboratory, Tokyo, per. comm. with TI, Dec. 1979) and by Soviet vessels off the Sala y Gomez Ridge (Parin et al. 1 98 1 : 1 2). The Kaiyo Maru specimens were examined (by TI) in 1980 at the Far Seas Fishery Research Laboratory in Shimizu, Japan. The Soviet collections consti- tute the primary basis for this account. We identify the eastern Pacific specimens with Malacocephalus laevis, a species originally de- scribed from the North Atlantic. The status of three other nominal Pacific species of the genus has not been completely resolved. The three in- clude M. nipponensis Gilbert and Hubbs, 1916, M. hawaiiensis Gilbert, 1905, and M. luzonensis Gilbert and Hubbs, 1920. Okamura (1970a:69) placed M. nipponensis into the synonymy of M. laevis, but later (Okamura in Okamura et al. 1 982: 145, 347-348) recognized M. nipponensis based on its lack of "scaled patch on the gular mem- brane and in a few other characters." Iwamoto (1970:41 1) recognized all three species and pro- vided a key to the genus based on the literature and examination of relatively few specimens. Marshall (1973:650-652) also recognized the three species but noted that "Examination of a good series of individuals may show that nip- ponensis is identical with hawaiiensis." In Table 4, selected counts and measurements of speci- mens of Malacocephalus from three oceans are compared. Although the data were not treated statistically, a casual inspection suggests that they do not support taxonomic recognition of the pop- ulations. (The raw data for this table are depos- ited in the library of the Department of Ichthy- ology, California Academy of Sciences; photocopies are available to any interested party.) Size.— To at least 52 cm. Distribution. — Probably worldwide in tem- perate to tropical waters, but absent on the con- SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 69 Table 4. Comparison of selected measurements and counts of Malacocephalus laevis from the Sala y Gomez Ridge, the Atlantic Ocean, and the Indian Ocean. tinental slopes of the eastern central and South Pacific. Depth range generally between 300 and 700 m, but often encountered shallower or deep- er. Material Examined. —(84 spec, 1 9 sta.) Sala y Gomez Ridge: ZMMGU 18074 (2:63.7-73.3 mm HL, 392+^120 + mm TL); 550 m; Astronomer trawl without no.; 25°02'S, 88°35'W; 24 Jul. 1975. ZMMGU 1 8075 (6:39.5-78.5 HL, 23 1 +^63 + TL); 345-540 m\ Ichthyandrcr. 5,tr. 55. ZMMGU 18076(14:41.7- 53.3 HL, 255-322 TL); 410 m; Ichthyandrcr. 5, tr. 56. ZMMGU 18077 (3:37.5-66.8 HL, 230+^03+ TL); 540 m; Ichthyandr cr. 5, tr. 53. ZMMGU 18078 (2:50.5-61 HL, 288+-359+ TL); 535-575 m; Ichthyandrcr. 5, tr. 54. ZMMGU 18079 (72 HL, 440+ TL); 420 m; Ichthyandrcr. 5, tr. 57. CAS uncat. (8:42.5- 79 HL, 248^43 TL); 430 m; Ichthyandrcr. 6, tr. 56. ZMMGU 18080 (4:40-74.5 HL, 257-410 TL); 565-555 m; Prof. Mesi- atzev cr. 13, tr. 1. ZMMGU 18081 (4:61.5-67 HL, 281 +- 443+ TL); 550-560 m; Prof. Mesiatzev cr. 13, tr. 2. ZMMGU 18082 (38 HL, 239 TL); 410-420 m; Prof. Mesiatzev cr. 13, tr. 4. ZMMGU 18083 (2:39-53.5 HL, 238-295 TL); 285-300 m; Prof. Mesiatzev cr. 13, tr. 7. ZMMGU 18084 (3:40-49.5 HL, 242-288 TL); 400 m; Prof. Mesiatzev cr. 15, tr. 52. ZMMGU 18085 (5:37.8-44.1 HL, 239 +-264+ TL); 530 m; Prof. Mesiatzev cr. 15, tr. 49. ZMMGU 18086 (43.5 HL, 271 TL); 304 m; Prof. Mesiatzev cr. 1 5, tr. 54. ZMMGU 1 8087 (6: 42.8-86.2 HL, 255^75 TL); 410-385 m; Prof. Shtokman cr. 18, sta. 1941. ZMMGU 18088 (5:39.5-88 HL, 214+^63 TL); 580-564 m; Prof. Shtokman cr. 18, sta. 1964. ZMMGU 18089 (5:40.5-84 HL, 237-490 TL); 562-545 m; Prof. Shtokman cr. 18, sta. 1965. ZMMGU 18090 (5:38.3-57.7 HL, 238+-333 + TL); 563-590 m; Prof. Shtokman cr. 18, sta. 1976. ZMMGU 18092 (7:45.5-100 HL, 237+^80 TL); 545-600 m; Prof. Shtokman cr. 18, sta. 1977. Mataeocephalus Berg, 1898 Type species: Coelocephalus acipenserinus Gilbert and Cra- mer, 1897, by original designation. Diagnosis. — Macrourines with 7 branchios- tegal rays. Snout long, pointed, somewhat de- pressed, tipped with a two-pronged scute. Mouth inferior, upper jaw less than 30% HL. Teeth in most species in short, broad bands, confined to anterior end of jaws. Outer gill rakers on first arch rudimentary or absent. Spinous ray of first dorsal fin serrated or smooth. Periproct at or close to anal fin origin. Distribution.— Tropical waters of Pacific and Indian oceans in upper to mid-slope depths of about 550-1,700 m (according to our unpubl. data). Remarks.— A small genus, comprising five or more species, in need of revision. See Iwamoto (1979:144) for a discussion of the genus. 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Mataeocephalus acipenserinus (Gilbert and Cra- mer, 1897) Coelocephalus acipenserinus Gilbert and Cramer, 1 897:422- 423, pi. 42, fig. 1 (Kaiwi Channel, Hawaiian Is.; 572-728 m; Holotype: USNM 47721). Mataeocephalus acipenserinus: Berg, 1898:41. Parin 1990:16 (listed from Sala y Gomez Ridge). Kotlyar and Parin 1990: 106 (otolith). Diagnosis.— Teeth in lower jaw confined to a short, broad, lunate patch at tip of jaws, none laterally. Underside of head anterior to lower jaw angle naked; scales below mid- ID. 6-7.5, below 2D. 8 or 9 (rarely 7). Some denticulations on leading edge of dorsal spine; V. 8; IP. i20-i25. Anus at posterior end of a broad, oval, naked area situated close to anal fin origin and far re- moved from pelvic fin insertions. Counts and Measurements.— (of 21 Sala y Gomez spec.) (see also Diagnosis) ID. 11,8-10; V. 8 (7/8 in one spec); GR-I (outer/inner) 3-5 total/6-8 total, GR-II 5-8 total/6-9 total; scales below ID. 8-9. Total length 1 18-250 mm, HL 26.5-54 mm. The following in percent HL: postrostral 58.1- 64.8; snout length 38.5-44.0, width at lateral an- gles 34.1^3.2; preoral 37.8-46.2; orbit 28.9- 34.4; interorbital 19.7-23.0; postorbital 26.3- 33.0; suborbital 1 3.9-1 7. 1 ; orb.-preop. 27.7-35.0; upper jaw 20.0-27.5; barbel 3.5-8.6; gill slit 7.4- 10.4;pre-lD. 105-1 13; pre-A. 126-140; pre- vent 116-135; pre-V. 96-119; V.-A. 22.6-35; body depth 47-60; ID. base 17.3-22; ID. height 50- 63; 1D.-2D. 13.5-32.2; IP. 41-57; V. 45-70. Description. — Entire head profile sharply conical in lateral view, the profile continuing smoothly to dorsal fin base dorsally and to pelvic or anal fin bases ventrally; thereafter profile ta- pers to slender, attenuated tail tip. Orbit large, about 1.5 into snout, almost equal to postorbital; sides of snout viewed dorsally gently curved from orbits to bifid scute on snout tip; interorbital about 1.3 into orbit, equal to width between su- pranarial ridges, space slightly concave; mouth "U" shaped, small, inferior, the rictus greatly restricted by lip folds laterally; upper jaw extends to a vertical slightly posterior to midorbit; barbel very small, length much less than large posterior nostril; suborbital forming a rounded triangle at posteroventral corner; interopercle exposed along posteroventral margins. Gill membranes broadly attached to isthmus; gill openings extend only to below preopercular ridge. Gill slits restricted, the first (outermost) slit about 10% of HL; other slits restricted by narrow fold of membrane between lowermost parts of arches. Rakers on outer side of first arch small, scarcely visible as tiny pimples that show no contrast with dark arches; rakers elsewhere short and tubercular. Scales below origin of second dorsal in 5 1 mm HL specimen with fine needlelike spinules ar- ranged in 5-8 parallel rows, fewer rows in smaller specimens. Underside of head naked forward of jaw angle except for overlapping series of mod- ified scales along leading edge of snout; some scale patches ventrally on preopercle. Suborbital strongly angular in cross section; heavy, modi- fied, scutelike scales in 2 rows on dorsal part leading forward to rather sharp edge of snout. Tip of snout with a pair of spiked, conical tu- bercles, as characteristic of genus. Ridges other than suborbital not strongly supported by heavy scales. A prominent naked groove dorsally be- hind broad anterolateral margins of snout. Jaw teeth all small, in a broad short band in premaxillae, in a short tapered band only at an- terior end of lower jaw. Disposition of teeth as illustrated for M. tenuicauda by Iwamoto (1979, fig. 9). Lengths of first dorsal and pectoral fins about half HL. Serrations on spinous dorsal ray weakly and sparsely developed, but present in all ex- amined specimens. Outer pelvic ray slightly thickened and prolonged, extending past anal or- igin to level of 1 0th anal ray or beyond in some specimens. Anal fin well developed, origin about on vertical through first dorsal insertion; pectoral and pelvic origins about on same vertical, that of first dorsal well behind. Interspace between dorsals 1-2 times length base of first dorsal. Periproct region broad, the region black, na- ked, and separated by a narrow gap from anal fin. Anus at posterior end of periproct, much closer to anal fin than to pelvic fin insertions. A large lenslike structure forming anterior portion of periproct extending forward towards, but fall- ing short of, pelvic girdle. Photophore length 6.2- 8.5% HL in 12 Sala y Gomez specimens (this compares with 6.7-8.1% in 3 Hawaiian para- types and 3.1-6.7%, usually less than 6%), in 21 specimens of M. microstomus from the Indian Ocean. Pyloric caeca short and thick: 14, 14, and 1 5 counted in 3 specimens. Overall color variable, but Sala y Gomez spec- imens generally rather light grayish overall, whit- ish on underside of head anterior to lower jaw SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 71 angle, and deep bluish over opercles and abdo- men. Fins clear to light dusky. Rim of nostrils narrowly blackish. Mouth black; gill cavity black dorsally and along gill slits and arches, but pale ventrally and on gill filaments. Size.— Attains at least 25 cm TL. Distribution.— Our Sala y Gomez Ridge specimens were collected at two closely adjacent stations on the westernmost seamount surveyed during the 1 8th cruise of the Prof. Shtokman. Until this cruise, M. acipenserinus was known only from off Hawaii. Its presence on the Sala y Gomez Ridge was unexpected, although several slope-dwelling Hawaiian species of other fami- lies have been captured on the ridge (Parin et al. 1981:5, and N. V. Parin, pers. comm.), suggesting a closer affinity of the two faunas than previously realized. Comparisons and Remarks.— A revision of the genus Mataeocephalus is in preparation by one of us (YS) and Y. N. Shcherbachev. Prelim- inary findings show the genus to comprise two species groups— M. adustus group and M. mi- cros tomus group— each with several species of uncertain status. Our Sala y Gomez Ridge spec- imens of Mataeocephalus belong to the second group, which includes M. microstomus (Regan 1908), M. nigrescens Smith and Radcliffe, 1912 (probably a synonym of the former species), M. tenuicauda (Garman 1899), and M. acipenseri- nus. The species in this group lack scales on the underside of the snout, and the mandibular teeth are in short lunate patches (as illustrated by Iwa- moto 1979, fig. 9a). Proportional measurements and counts of all species in the M. microstomus group are similar. Differences between species are slight, and the main differentiating character appears to be the degree of development of the light organ. In this regard, our specimens agree most closely with the Hawaiian species, M. acipenserinus, in hav- ing the light organ anteriorly prolonged. In con- trast, M. microstomus has a small subtriangular gland with a tiny anterior dermal window (ADW) just in front of the periproct, and M. tenuicauda has a rudimentary gland and no ADW. Our Sala y Gomez specimens differ slightly from para types of M. acipenserinus in that the anterior margin of the ADW in the paratypes is much closer to the insertion of the inner pelvic rays than is the case in our specimens. This re- flects the shorter distance between pelvic and anal fins in the paratypes (22.6-31.9% HL, usu- ally less than 26% cf. usually 26%-34.9% [but 22.6% in one specimen]). Considering the great variability and consequent low taxonomic value of this character, and in the absence of other discernible differences, we recognize the Sala y Gomez populations as M. acipenserinus. Material Examined.— Sala y Gomez Ridge: ZMMGU 1 7732 (8:26.5-51.5 mm HL, 1 17-250 mm TL) and CAS uncat. (5: 40-50 HL, 166+-182 + TL); 750-800 m; Prof. Shtokman cr 18,sta. 1996. ZMMGU 17733 (8:32.3-54.7 HL, 155-240 TL) 730-790 m; Prof Shtokman cr. 18, sta. 2018. Hawaiian Is- lands: CAS-SU 3142 (4 paratypes, 44-51.5 HL, 161-177 TL): 21°08'30"N, 157°49'W; 627 m; Albatross sta. 3470. Nezumia Jordan, 1 904 Type species: Nezumia condylura Jordan and Gilbert, 1904, by original designation. Diagnosis. — Macrourines with 7 branchios- tegal rays. Teeth small, in narrow to broad bands in both jaws; those on premaxillary do not occur past maxillary process. Gill rakers < 1 2 on inner side of first arch in most species. Snout variously naked on ventral surfaces, anteriorly tipped with spiny tubercles in most species. Suborbital shelf with 2 rows of modified scutelike scales forming a prominent ridge. Body scales covered with nee- dlelike to shield-shaped spinules; transverse ridges present. Spinous second ray of first dorsal fin serrated in most species. Anus removed from anal fin origin and situated within an oval-shaped periproct. A small naked fossa of light organ be- tween pelvic fins. Distribution.— Worldwide in temperate to tropical seas; most species found at upper to mid- dle continental-slope depths (about 200-1,500 m). Nezumia convergens (Garman, 1899) (Figure 26) Macmrus convergens Garman, 1899:210, pi. 48, fig. 1 (Gulf of Panama, 695-1,020 fm [1,271-1,865 m]; Albatross sta. 3353, 3357, 3393). Nezumia convergens: Parin and Sazonov 1 982:86 (5 spec; Peru). Parin 1990: 16 (recorded from area between Nazca and Sala y Gomez ridges). See Iwamoto (1979:171) for synonymy. Diagnosis. — Body slender, greatest depth 7- 8 in TL in large adults. Upper jaw 26-33% HL; barbel 8-20% HL (usually 1 .5-2.0 in orbit); GR-I and GR-II (inner series) (1-2) + (5-8) (usually 5-7 on lower limb). Spinules on body scales con- ical to narrowly lanceolate, aligned in 10-12 slightly convergent longitudinal rows in large 72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 vwKwyWJ!** '^m^^^^ 1 ^^^•^^■^j^,. Figure 26. Nezumia convergens. From Iwamoto (1979:fig. 15b) adults, middle row often slightly enlarged. Al- most all of ventral surfaces of snout and antero- ventral surface of suborbital naked; mandibles naked anteriorly. Second spinous ray of first dor- sal fin longer than postrostral length of head, its leading edge beset with closely spaced teeth; first dorsal fin uniformly dusky; V. 10-11. Anus in middle half of space between pelvic fin insertion and anal fin origin. Light organ not well devel- oped externally; scaleless fossa not present be- tween pelvic fin bases. Counts and Measurements. — (of NSG spec- imen only) ID. 11,10; IP. i23; V. 11; total GR-I [outer/inner] 9/10, GR-II 10/10; scales below ID. 11, below mid- ID. about 10, below 2D. 9.5; lat.1.46. The following in mm, percent HL in paren- theses: snout 14.1 (29.1); preoral 1 1.6 (23.9); or- bit 14.5 (29.9); interorbital 9.9 (20.4); postorbital 20.8 (42.9); orbit-preop. 18.8 (38.8); suborbital 7.4 (1 5.3); upper jaw 16.2 (33.4); barbel 7.5(15.5); gill slit 6.8 (14.0); pre-A. 75 (154); pre-vent 66 (137); 1D.-2D. 14.5 (30); height ID. 49 (101); IP. 29 (59); V. 30 (62); body depth 36 (73). Size.— To about 30 cm. Distribution. — Broadly distributed along continental slopes of eastern Pacific, from Gulf of California and northern Mexico south to Chile (lat. 35°S) and also in Galapagos Archipelago. Depth range 600-1,865 m. Remarks.— See Iwamoto (1979: 171, figs. 15b, 1 8f) for a description and illustration. This fine specimen taken near the Sala y Gomez Ridge agrees well in most respects with specimens of TV. convergens from the continental margins, al- though a few characters fall outside the ranges established for the species by Iwamoto (1979). Most notable is the gill raker count, which was high by one raker in the inner series of the first and second arches. Scale spinules also appeared to be more broadly lanceolate. The lateral line scale count over a distance equal to the predorsal length differed by two (46 vs. 36-44), the post- orbital length was slightly longer (43% HL vs. 34-40%), and the height of first dorsal was great- er (101% HL vs. 68-93%). Unfortunately, only the single specimen was collected, thus preclud- ing a more meaningful analysis of these differ- ences. Material Examined. -ZMMGU 17734 (1:48.5 mm HL, 272+ mm TL); area between the Nazca and Sala y Gomez ridges, 1,050 m; Prof. Mesiatzev cr. 13, tr. 14. Nezumia propinqua (Gilbert and Cramer, 1897) (Figures 27, 28a, b) Macwurus propinquus Gilbert and Cramer, 1897:424, pi. 42, fig. 2 (type-locality Kaiwi Channel, Hawaiian Islands; 642 m). Gilbert 1905:667. Lionurus propinquus: Gilbert and Hubbs 1916:144 (list). Nezumia propinquus: Marshall and Iwamoto 1973:625 (list). Okamura in Okamura et al. 1982:163, 350, color fig. 97 (6 spec, 216-250 TL; Kyushu-Palau Ridge; 695-219 m). Nezumia propinqua: Iwamoto 1983:8; 1986:339 (2 spec; Mo- zambique; 740 m). Parin 1 990: 1 6 (listed from Sala y Gomez Ridge). Kotlyar and Parin 1990:106 (otolith). Diagnosis. — Body scales covered with slender conical spinules in 5-8 parallel rows (in adults to about 20 cm TL); scales below 2D. 10-13. Mandibles and underside of head to posterior angle of mouth naked. A distinct black tip on first dorsal fin. Length pectoral fin about 60% HL. Pelvic fin rays 14-18; IP. rays il8-i22 (count data included for specimens from other areas). Counts and Measurements.— (from 17 Sala y Gomez specimens) ID. 11,1 1-13; IP. il9— i22; V. 15-17 (rarely 14 or 18); total GR-I (outer/ inner) 7-12/9-12; total GR-II 9-11/8-11; scales below ID. 12-14, below mid- ID. 8-11, below 2D. 10-13, lat. 1 . 43-47; caeca 24 (1 spec). SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 73 m&r- Figure 27. Nezumia propinqua (approximately 30 mm HL) from Sala y Gomez Ridge, collected on Prof. Shtokman cr. li Photograph by Sergei Dudarev. Total length 125+-200 + mm; HL 19.3-34 mm. The following in percent of HL: snout 26- 32; preoral 20-29; orbit 31^10; interorbital 22- 28; postorbital (greatest) 41-45, (least) 34-39; orbit-preop. 32-40; suborbital 12-15 (19); upper jaw 31-38; barbel 17-28; gill slit 14-19; pre-A. 131-159; pre-V. 96-121; pre-vent 119-142; V.- A. 36-53; 1D.-2D. (17) 24-57; height ID. 84- 108; IP. 58-69; V. (74) 80-99; body depth 78- 100. Description.— Head about 6 in TL, body depth slightly less than head length; trunk short, length abdominal cavity much less than HL. Snout bluntly pointed, tipped with a broad coarse tubercle; mouth subterminal, small, upper jaw about one-third HL, extends to below hind edge of pupil; barbel short but prominent, about one-half orbit diameter; gill membranes rather narrowly restricted across isthmus with a mod- erately free posterior fold, the gill openings ex- tending forward to below hind edge of orbits. Anus within a broad periproct, somewhat closer to anal fin origin than to pelvic origins. A small oval dermal window between pelvic bases. Ab- dominal cavity terminates above 9th or 1 0th anal ray. First dorsal fin with a small keellike first spine; second spinous ray prolonged, armed along lead- ing edge with sharp and widely spaced serrations, length of spine about equal to HL. Second dorsal fin rudimentary over entire length. Pectoral and pelvic fins rather short; outer ray of pelvics slight- ly prolonged into a filament that extends to end of abdominal cavity. Origin of pelvics below pos- terior margin of opercle and in advance of pec- toral base, which in turn is in advance of first dorsal origin; anal origin below midbase of first dorsal. Scales with about 7-10, more or less parallel rows of small, reclined, needlelike spinules (Fig. Figure 28. Scales from dorsum below interspace between dorsal fins of: (a) Nezumia propinqua (CAS-SU 8538; Hawaii); (b) N. propinqua (Sala y Gomez Ridge); (c) N. condylura (CAS-SU 22925; Japan). Scale bar represents 0.5 mm. Drawn by Tomio Iwamoto. 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 28b). Suborbital shelf formed of a double row of stout scales. Ventral aspects of snout, most of suborbital, and lower jaw completely naked. Ce- phalic lateral-line pores rather prominent along lower margins of snout, suborbital, preopercle, and lower jaws. Teeth in broad bands in both jaws; outer series of upper jaw slightly enlarged. Large papillae densely interspersed between teeth and along mesial side of jaw, giving superficial appearance of mandibular band being much broader than it actually is. A 29-mm HL specimen from CAS 75990 had 24 short, thick pyloric caeca, their lengths about equal to pupil diameter. Ovaries in that speci- men moderately developed, but no eggs yet vis- ible. Largest Sala y Gomez specimen (ZMMGU 17749, 34 mm HL) a mature female with well- developed ovaries containing eggs about 1.0 mm in diameter. Color in alcohol pale yellowish brown overall; abdomen bluish, turning to blackish ventrally on trunk. Fins all pale except distally black on first dorsal. Mouth pale; broad outer margins of gill cavity dark; gill rakers and gular membranes dark or blackish. Underside of head pale. Size.— To about 25 cm. Distribution.— Nezumia propinqua has been reported from Hawaii, the Kyushu-Palau Ridge, off Vietnam, off western Australia, and off Mo- zambique, in 523-870 m. Comparisons and Remarks.— Nezumia pro- pinqua and N. condylura are extremely close and may be conspecific. Okamura (in Okamura et al. 1982:350) used the distance anus to anal fin as the primary character to distinguish the two spe- cies. From measurements we have made, that distance ranges from 1 5-20% HL in N. condylura compared with 12-32% HL (most greater than 20% HL) in N. propinqua. On that basis, the Sala y Gomez specimens would fall into N. condylura. The length of pelvic fins also suggests a separa- tion of the two: 79-100% in N. propinqua vs. 68- 81% in N. condylura. Based on this measure- ment, the Sala y Gomez specimens fall into N. propinqua. Another character that may offer a means of separating the two species is the num- ber of spinule rows on body scales. In specimens of 27-28 mm HL, N. propinqua specimens had 5-8 spinule rows on scales below the interspace of the dorsal fins, whereas N. condylura speci- mens had 8-12 rows (see Fig. 28). No other char- acter we have examined suggests a separation, but a good size-series of fresh, well-preserved specimens of each have not been compared. For now, its seems best to treat the current material as ./V. propinqua, recognizing, however, that fu- ture workers making a more thorough study may arrive at another conclusion. Nezumia evides Gilbert and Hubbs, 1 920, rep- resents another closely related species that can be distinguished by its slightly fewer scale rows below the second dorsal fin (9-10). Material Examined.— (17 spec.) Sala y Gomez Ridge: ZMMGU 17741(31. 5 HL, 188+ TL); 565-555 m; Prof. Mesi- atzevcr. 13, tr. 1. ZMMGU 17742 (2:22.3-3 1.5 mm HL, 151+- 191.5 mm TL) and CAS 75990 (2:29-32 HL, 164+-200 + TL); 550-560 m; Prof. Mesiatzev cr. 1 3, tr. 2. ZMMGU 1 7749 (34 HL, 175+ TL); 540-550 m; Prof. Shtokman cr. 18, sta. 1970.ZMMGU 17750(4:20-29.5 HL, 125+-169+ TL); 750- 800 m; Prof. Shtokman cr. 18, sta. 1996. ZMMGU 18070 (6: 19.3-33 HL, 121+-174.5+ TL); 730-790 m; Prof. Shtokman cr. 18, sta. 2018. ZMMGU 18126 (26 HL, 157+ TL); 580- 564 m; Prof. Shtokman cr. 18, sta. 1964. Pseudocetonurus Sazonov and Shcherbachev, 1982 Type species: Pseudocetonurus septifer Sazonov and Shcher- bachev, 1982, by original designation. Diagnosis. — Macrourines with 7 branchios- tegal rays. Head notably large and broad, pre- opercle and suborbital bones deep and large, opercle commensurately small; orbit small, 1 9- 30% HL, diameter much less than broad inter- orbital; snout high, slightly projecting beyond mouth. Mental barbel small, 10% or less of HL. Gill opening wide, extending forward to below hind end of maxilla; gill membranes loosely and narrowly attached to isthmus. Gill rakers usually 16 or 17 total on inner series of first arch. Teeth small, close-set, in narrow tapered band on pre- maxilla, uniserial on dentary. Scales with nu- merous small, awl-shaped spinules; no reticula- tions on anterior field; lateral line scales absent, a series of dark papillae in its place. Vent about halfway between pelvic fin insertion and anal fin origin (usually closer to pelvic insertion), sur- rounded by a black, oval to teardrop-shaped na- ked area and preceded by a small, round dermal window of light organ between pelvic fin bases. Pyloric caeca short, 22-34. Color black to dark brown overall. Distribution. — Known from Hawaii and the Nazca and Sala y Gomez ridges. Remarks.— Only the single species known. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 75 Figure 29. Pseudocetonurus septifer. (a) Juvenile (ZMMGU 18127, 26.7 mm HL), Professor Shtokman sta. 1856, IKMWT in 900-0 m, 20 April 1987. (b) Adult (ZIN uncat., 72 mm HL, 393+ mm TL) from Nazca Ridge in 950 m; Hercules sta. 40. Photograph by Susan Middleton. Pseudocetonurus septifer Sazonov and Shcher- bachev, 1982 (Figure 29) Pseudocetonurus septifer Sazonov and Shcherbachev, 1 982:7 1 2, fig. 2 (type locality Sala y Gomez Ridge, 850-860 m). Parin 1990:16-17 (listed from NSG). Iwamoto in Cohen et al. 1990, fig. 229 (in key). Diagnosis.— As for the genus. Counts and Measurements. — 1 D. 11,8-12 (usually 9 or 10); IP. il6-i20; V. 9 or 10, rarely 8; GR-I (outer/inner) 7-12 (usually 9-1 l)/(l-2) + (12-16) (total 13-18, usually 16 or 17), GR- II (1-3) + (12-15) (total 14-18, usually 15 or 16)/(l-3) + (12-15) (total 14-17, usually 15 or 16); scales 1D.-A. 33-39 (n = 18), midbase 1D.- A. 25-34; 2D.-A. 18-24 (n = 17); caeca 22-34 (n = 10). Total length 1 1 7 +-393 + mm; HL 24-72 mm. The following in percent HL: postrostral 72.3- 79. 1 ; snout 25. 1-34.4; preoral 1 8.8-27.0 (usually 20-25); orbit 18.8-31.2; interorbital 32.8^4.4; postorbital 46.6-58.6; orbit-preop. 52.5-64.4; suborbital 19.3-25.5; upper jaw 40.9-48.5; bar- bel 6.5-10.3; outer gill slit 22.6-30.6; body depth 75.4-94.5; pre-D. 95-1 1 1; pre-V. 102-127; pre- vent 109-134; pre- A. 119-146; V.-A. 12-38.1; ID. height 52-66; ID. base 22.8-30.2; 1D.-2D. 24-38; IP. 68-81; V. 41-63. Description. — Head large, 5.0-5.5 into TL, relatively broad and deep, greatest width about 1.5 into greatest depth; trunk compressed, grad- ually tapering into a long straplike tail (the tip often missing). Orbits small, semi-elliptical to oval, its greatest diameter usually oblique. In- terorbital broad, irregularly convex, 2.3-3.0 into HL. Suborbital region deep, almost vertical, con- vex, without division into upper and lower parts, and without stout scutelike scales. Preopercle no- tably large, with posterior margin oblique, more strikingly so in smaller specimens. Subopercle posterior margin forming a deep notch; sub- opercle and interopercle broadly exposed beyond preopercle in holotype and 393 mm specimen 76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 (ZIN uncat.), but interopercle narrowly exposed in smaller specimens. Free margins of opercular bones crenulated. Snout short, bluntly pointed, and high, forming a pronounced hump in dorsal profile, its length usually greater than orbit di- ameter; anterior profile subvertical. Mouth large, jaws subterminal; posterior end of maxillary ex- tends to vertical through midorbit. Mental barbel very small, almost a rudiment, length less than half orbit diameter. Gill membranes narrowly attached to isthmus below hind end of upper jaws (opercular opening consequently wide). Sensory canals on head large. Free neuromasts on head indistinct and poorly developed. Lateral line canal absent on body, being replaced by small black papillae (free neuromasts?) situated some- what irregularly along normal course of lateral line. Olfactory capsules small; posterior nostril slightly larger than anterior; internarial mem- brane forming a prominent flap equal to diam- eter of anterior nostril in 393 mm ZIN specimen, but flap inconspicuous in smaller specimens. Teeth in jaws small, conical, closely spaced; uniserial on dentary; in narrow band 3-4 irreg- ular rows wide anteriorly on premaxillary, nar- rowing to about 2 rows wide posteriorly, outer series slightly enlarged. Squamation in 393 mm specimen overall forming a smooth, velvety cover without coarse ridges or scutes. Head almost completely scaled, including underside of snout, suborbital region, and mandibular rami. No scaly ridges on head; no scutes at terminal and lateral tips of snout. Scales on posterior margins of opercular bones lacking spinules (or absent?). Scales absent on gular and branchiostegal membranes, over su- pranasal ridge, dorsal edge of orbit, on posterior portion of median nasal ridge, and in rather wide band surrounding postorbital sensory canal. Scales greatly enlarged over postorbital canal and posterior portion of interorbital space to level of anterior margin of pupil, but very small anteri- orly on snout, especially along leading edges, with some scales nonimbricate. Elsewhere on head, scales intermediate and about equal in size. Body scales large, covered with numerous slender, sub- erect, slightly curved spinules in quincunx order. Light organ externally consists of a teardrop- shaped anterior dermal window between mid- base of pelvic fins, connected by a narrow isth- mus to posterior dermal window at anterior end of periproct. Anus much closer to pelvic fin bases than to origin of anal fin. Swim bladder well developed. Pyloric caeca short, unbranched. Ra- diographs of two specimens showed abdominal vertebrae 12; anal pterygiophores before first haemal spine 12. Origin of first dorsal fin opposite that of pelvic fin; pectoral fin situated slightly anterior to these; anal fin origin slightly behind vertical through last ray of first dorsal. Second unbranched ray of first dorsal high, not especially stout, terminating in a short filament; leading edge smooth except for distal one-fifth or so where numerous small, inconspicuous denticles present. Rays of second dorsal moderately developed, but much finer and shorter than opposites of anal fin. Interdorsal space about equal to snout length. Pectoral fins long, more than two-thirds HL, situated well be- low level of top of gill opening; pelvic fin long, outer ray filamentous and extending well past anal fin origin. Color in alcohol overall black to brownish black. Scale pockets on head and body of largest specimens dark blue with black margins; in smaller specimens most dorsal surfaces of head, snout, and suborbital grayish. Opercle, gular and gill membranes, and jaws black; in small speci- mens leading edge of snout and rim of olfactory capsule faintly blackish; oral cavity mostly black, but floor anteriorly whitish; branchial cavity black except pale anteriorly on isthmus and gill fila- ments. All fins black. Holotype generally lighter in color than above description suggests, but this probably a result of fading, which was not so evident in other specimens. Size.— To more than 39 cm. Distribution.— Nazca and Sala y Gomez ridges and Hawaiian Islands, in 340-950 m. Remarks.— The collection of many small ad- ditional specimens of this species, originally de- scribed from a single large individual, has shown its similarity to members of the genus Ventri- fossa. This supports moving Pseudocetonurus even farther from Cetonurus, with which it was initially loosely grouped, but "without ... in- dicating a close kinship" (Sazonov and Shcher- bachev 1982:6). In fact, aside from the lack of a developed lateral line, the expanded interorbital, suborbital and preopercular regions, the rela- tively large head, and the high snout, there is little to suggest that Ventrifossa and Pseudoce- tonurus are much different. The dark color, small eyes, and enlarged sen- sory canals suggest a bathypelagic existence, and the capture of many specimens in a midwater SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 77 trawl over the Nazca Ridge lends support to this idea. The swim bladder is rather well developed, nonetheless, and in large adults it is enveloped in a tough tunica externa and filled with spongy tissue. The rete-gas gland complex of a small 1 50 mm specimen was well developed and consisted of a straplike rete that terminated posteriorly in a tightly appressed pair of flattened, horse-hoof- shaped gas glands. The rete bundle did not ap- pear to be separated into two bundles, although closer inspection may show them to be so, as it is in other members of the tribe Malacocephalini. The discovery of four specimens in good con- dition from the Hawaiian Islands was surprising and suggests the possibility of a much wider dis- tribution for this species. The specimens were dip netted at the surface in 1950 after having been killed by the lava flow from a volcanic erup- tion of Mauna Loa on the Island of Hawaii. Gos- line et al. (1954) and Gosline (1954) reported on the kill and collection of the fishes from this erup- tion. Material Examined.— Sala y Gomez Ridge: Holotype: ZMMGU P16011 (54 mm HL, 257+ mm TL); 25°20.2'S, 93°35.5'W; 850-860 m. Other Material: Nazca Ridge: ZMMGU 17743 (19:24- 41.5 HL, 93+-212 + TL) and CAS 67409 (10:29.5-38.5 HL, 125+-184 + TL); 340-780 m; IKMWT; Prof. Mesiatzev cr. 13, sta. 44. ZIN uncat. (72 HL, 393+ TL); 950 m; bottom trawl; Hercules tr. 40. ZMMGU 18127 (26.7 HL, 126+ TL); 900-920 m over bottom depth of 1,270-1,200 m; IKMWT; R/V Prof. Shtokman cr. 1 8, sta. 1 856. Hawaiian Islands: LACM 45410-1 (4:39-45 HL, 209-231 TL); lava flow kill, Kona, Ha- waii; Y. Yamaguchi, collector; 7. VI. 1950. Trachonurus Giinther, 1887 Type species: Coryphaenoides villosus Giinther, 1877, by monotypy. Diagnosis. — Macrourines with 7 branchios- tegal rays. Teeth in narrow bands in both jaws. Scales bristly, covered with short to moderately long, erect, conical spinules; head entirely scaled, including patches on gular and (in some) bran- chiostegal membranes; scales along second dor- sal and anal fins somewhat enlarged and thick- ened in most species. Second spinous ray of first dorsal fin slender, flexible, without serrations; 6- 8 segmented first dorsal rays; pelvic fin with 6 or 7 rays, its origin below base (usually midbase) of first dorsal. Anus in middle of broad oval periproct spanning most of distance between anal and pelvic fins, a broad, naked anterior extension ending at pelvic fin bases. Abdominal vertebrae 12 or 13. Swim bladder with 2 retia mirabilia. Pyloric caeca 6-14, stubby to moderately long and thick. Color blackish to dark brown. Size.— To at least 64 cm. Distribution. — Worldwide in tropical to warm-temperate waters, but not recorded from continental slopes of eastern Pacific. Remarks.— The genus constitutes a small group of four nominal species, all of which were synonymized by Marshall (1973) with Tracho- nurus villosus. A closer re-examination of spec- imens from Hawaiian and Atlantic waters may require a re-evaluation of the status of T. sen- tipellis Gilbert and Cramer, 1897, and T. sulcatus (Goode and Bean, 1885). Study (by TI) of old Albatross collections from the Philippines and recent collections from Australia and New Zea- land suggests the possibility of as many as four undescribed species represented. Shcherbachev et al. (1979) reported numerous specimens of T. villosus from the Indian Ocean; our subsequent examination of this material suggests that some of the specimens may represent undescribed spe- cies. Trachonurus villosus (Giinther, 1877)? Coryphaenoides villosus Giinther, 1877:441 (south of Yeddo [=Tokyo]). Macrurus (Trachonurus) villosus: Giinther 1887:142, pi. 36, fig. B (lectotype, BMNH 1887.12.7.105. here designated; s. of Yeddo, Challenger sta. 232 in 345 fm [631 m]. Paralec- totype, BMNH 1887.12.7.106; s. of Philippine Is., Chal- lenger sta. 214 in 500 fm [914 m]). Trachonurus villosus: Parin 1990:17 (listed from vicinity of Nazca and Sala y Gomez ridges). Diagnosis.— Orbit about 15-16% HL; sub- orbital width about half orbit diameter. Outer premaxillary teeth slightly enlarged. A well-de- veloped grooved lateral line; scale spinules rel- atively short and stout; diagonal scale rows from hind edge of first dorsal to anal fin more than 25; gular and branchiostegal membranes heavily scaled. Rays IP. il6-il8. Pyloric caeca moder- ately long, not stubby. Remarks.— The two specimens from the Sala y Gomez Ridge appear quite similar to speci- mens of Trachonurus from the Gulf of Mexico and Caribbean Sea, and some specimens from Australia. A more thorough study is necessary to determine if they are the same. They differ from a syntype of T. villosus (BMNH 1 887. 1 2.7. 105; 26 mm HL, 1 77 + mm TL), taken south of Yeddo (Tokyo) in having a smaller or- 7S PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 bit, broader suborbital, longer distance orbit-pre- opercle, longer upper jaw, and more pectoral fin rays. It cannot be discounted that the exception- ally large size of these Sala y Gomez specimens may account for these differences in proportional measurements. A second syntype (BMNH 1887.12.7.1 06) from south of the Philippines ap- pears to be a different species with a longer upper jaw, longer barbel, longer first gill slit, wider 1 D- 2D. interspace, and larger scales. (This specimen was in poor shape when examined by TI in Oct. 1 986.) The features characterizing T. villosus have not been properly assessed, and the relationships of the species with others of the genus have not been adequately determined. Because two spe- cies are probably represented in the syntypic se- ries of T. villosus, the Yeddo syntype is here des- ignated as the lectotype. It was obviously the one used by Gunther in his original description, which listed only "south of Yeddo" as the type locality, and it was the specimen figured in his report on the Challenger fishes (Gunther 1 887: 142, pi. 36B). The Sala y Gomez specimens differ from Ha- waiian specimens of the genus (nominally T. sen- tipellis) in having a deeper body, a more rounded, less conical snout, more diagonal scale rows be- low the hind margin of first dorsal, more nu- merous pectoral fin rays, smaller orbit, narrower suborbital, and more heavily scaled gular and branchiostegal membranes. Material Examined.— (2 spec, both Prof. Mesiatzev) Sala y Gomez Ridge: ZMMGU 17744 (1:124 mm HL, 637+ mm TL); 1,070-1,100 m; cr. 13, tr. 10. ZMMGU 17745 (1:65 HL, 303+ TL); 1,050 m; cr. 13, tr. 14. Ventrifossa Gilbert and Hubbs, 1920 Type species: Coryphaenoides garmani Jordan and Gilbert, 1904, by original designation. Diagnosis. — Macrourines with 7 branchios- tegal rays. Head smoothly rounded, without coarse, angular ridges, and lacking stout, modi- fied scales; snout without a terminal scutelike scale except in subgenus Sokodara. Mouth mod- erate to large, upper jaw 35-55% HL. Jaw teeth in narrow to moderately broad bands in upper jaw, outer series enlarged; lower jaw teeth small, none enlarged, in 1 to several irregular series laterally. Gill rakers 1 2-20 total on inner series of first arch. Branchiostegal membranes naked. Body scales small, densely covered with slender conical or triangular spinules; no transverse ridg- es on anterior field. Spinous second ray of first dorsal finely serrated along leading edge or smooth. Anus removed from anal fin origin and closer to pelvic fin; periproct oval-shaped, ex- tending forward to small fossa between pelvic fin bases. Pyloric caeca about 34-75. Size.— To at least 53 cm. Distribution.— Tropical waters of the Indo- Pacific, but with an isolated pocket of two species in the tropical western Atlantic (Gulf of Mexico and Caribbean Sea). Species generally confined to upper continental slope depths ranging from about 200 m to more than 1,000 m, but most often captured at 500-800 m. Remarks.— A medium-sized group with 24 named species and several more known to us, yet to be described. Sazonov (1985) removed the subgenus Lucigadus Gilbert and Hubbs, 1920, with its five species, and elevated it to the status of genus. Iwamoto (1979) has recently treated the group in some detail. Ventrifossa johnboborum Iwamoto, 1982 (Figure 30) Ventrifossa johnboborum Iwamoto, 1982 (in part):55-61, fig. 1 (type locality Bismarck Sea; non-type specimens are V. fusca Okamura, 1982). Shcherbachevetal. 1986:202 (3 spec. from western Indian Ocean on Freda Seamount, 810 m). Shcherbachev 1987:7 (listed from Indian Ocean). Parin 1990: 17 (listed from Sala y Gomez Ridge). Kotlyar and Parin 1990:107, fig. 4a (otolith). Malacocephalus sp.: Parin et al. 1981:13 (1 spec, ZMMGU 17746, here reported, from Ichthyandr cr. 5, tr. 53). Diagnosis.— A broad-headed species of sub- genus Sokodara (see Iwamoto 1 979). Interorbital width 27.8-33.4% HL, slightly less than orbit diameter, which goes about 3 in head; upper jaw 39-45% HL; barbel 13-19% HL. Scales small, 65-7 1 from lateral line origin over distance equal to pre- ID. length. Dorsal spines weakly serrated in adults; ID. 11,10-11 (13); IP. i21-i23; V. 9- 10. Counts and Measurements.— (5 specimens; see also Diagnosis) GR-I (outer/inner) 7—10/12— 14 total; GR-II (outer/inner) 10-13/1 1-14 total; caeca 65-73 (2 spec); vertebrae 14-16. Total length 280+-391 + mm, HL 66.5-85.1. The following in percent of HL: postrostral 73.8- 76.7; snout 27.8-31.6; preoral 19.2-23.8; orbit 31.4-35.7; suborbital 10.5-14.0; postorbital 38.3-43.6; orbit-preop. 39.1-42.7; gill slit 20.4- 28.7; pre-lD. 101-110; pre-V. 97-114; pre-A. 127-137; V.-A. 33-49; base ID. 23.8-31.0; 1D.- 2D. 33.1^15.1. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 79 — v~> Figure 30. Ventrifossa johnboborum. Sala y Gomez Ridge in 540 m, Ichthyandr cr. 5, sta. 53. Drawn by Tomio Iwamoto. Scale bar equals 25 mm. Description.— Greatest body depth slightly more than postrostral length of head, about 1.3- 1.4 in HL, more than 6 in TL. Head width more than half its length. Snout broad, width at lateral angles (anterior to nostrils) slightly less than both interorbital width and snout length; moderately protruding beyond large subterminal mouth, length less than orbit diameter. Orbits large, 2.7- 3.2 in HL, about 1.2-1.3 in postorbital length. Suborbital region gently rounded, separated into upper and lower regions by a ridge-line that runs close to orbital rim anteriorly but diverges widely posteriorly; scales on suborbital not modified into heavy tubercular scutes. Preopercle broadly rounded; interopercle exposed along posterior and ventral margins of preopercle. Upper jaw 2.2- 2.4 into HL; maxillary extends to below posterior one-third of orbits; rictus not restricted poste- riorly. Barbel small, slender, about 1.8-2.5 in orbits. Premaxillary dentition in holotype in narrow band 4-5 irregular rows wide near symphysis, narrowing posteriorly. In Sala y Gomez speci- mens, teeth in 2 or 3 irregular rows, outer series slightly enlarged. A wide toothless gap at pre- maxillary symphysis. Mandibular dentition small, in 2 or 3 irregular series that taper to 1 row posteriorly; inner series larger than outer series. Scales all small, densely covered with small, slender, conical, relatively erect spinules ar- ranged in irregularly quincunx or widely diver- gent, V-shaped rows. Scales uniformly cover all of head except gill membranes, lips, and nostril membranes. A row of small scales on anterior orbital rim between posterior nostril and orbit. Scale pockets present on exposed interopercle surfaces, although no scales remain there on ex- amined specimens. No stout spiny ridges or heavily modified scales on head or body, except tip of snout has small, blunt, tubercular median scale (not noticeably enlarged or protruding). Fins unsealed. Pyloric caeca about 65 in holotype, 73 in a male 325+ mm TL. Caeca short, about equal to suborbital width, branched at base. Retia 2, long, slender; gas glands small. Paired drumming mus- cles in 325 mm male large, covering almost half antero ventral surface of swim bladder. Stomach in that specimen contained remains of cepha- lopods, including beaks, eyes, and body parts. Light organ with 2 dermal windows, the anterior one between pelvic bases small and round. First dorsal and paired fins moderately large for genus. Height of first dorsal fin about equal to postrostral length of head; spinous second ray weakly serrated distally, scarcely prolonged be- yond succeeding rays; when laid back, longest rays barely reach or extend slightly beyond origin of second dorsal fin. Base of first dorsal about 1.1-1.6 into space between first dorsal and sec- ond dorsal. Origin of pelvic fin anterior to that of pectoral, which is anterior to that of first dor- sal. Outer pelvic fin ray slightly produced into a filament that extends to about 3rd or 4th anal ray. Origin of anal fin under hind edge of first dorsal. Length of pectoral about equal to length of snout plus orbit. Color in alcohol overall grayish brown to swar- thy, bluish over abdomen; fins black. Gill mem- branes and entire oral cavity black. Gill chamber pallid over epihyal, ceratohyal, anteroventral 80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 31. Ventrifossa macwdon new species; non-type material, ZMMGU 18139, from Sala y Gomez Ridge, in 730-790 m, Prof. Shtokman cr. 18, sta. 2018, 7 May 1987: (a) specimen 67.5 mm HL; (b) 69 mm HL. Photograph by Sergei Dudarev. portion of cleithrum, and gill filaments; remain- der dark dusky to blackish. Gill arches and rak- ers, leading edge of snout, and periproct region blackish. Peritoneum and stomach black. Anal rays blackish anteriorly fading to dusky poste- riorly. Barbel overall pallid but base blackish. Lips edged with thin black margin. Comparisons and Remarks.— Ventrifossa johnboborum is most closely related to V. fusca Okamura, 1982, described from the Kyushu-Pa- lau Ridge, and V. misakia Jordan and Gilbert, 1904, from Japan. Ventrifossa johnboborum can be distinguished from the other two species by its blackish lining of the oral cavity (compared with pallid or white) and pelvic fin ray count (9- 10 cf. 8). It is further distinguished from V. fusca by its larger orbit (2.9-3.1 in HL, cf. 3.4-3.6), shorter interspace between dorsal fins (2.3-3.0 in HL cf. 1.8-2.1), and longer barbel (1.8-2.7 in orbit diameter, cf. 3.8—4. 1). The original description of V. johnboborum was based on the holotype and three other spec- imens not designated as type-specimens. These other specimens, one from offBatag Island in the Philippines, the other two from the South China Sea, were sufficiently different from the holotype so as to cast doubt as to their conspecificity, but their differences (V. 8, barbel length 11-1 2% HL, upper jaw length 36-39% HL) did not justify description of a second species. The specimens otherwise agree closely with V. johnboborum and also V. fusca, especially concerning the pelvic ray count, and proportional measurements of the barbel and upper jaw. They disagree with V. fusca in having dark mouth linings (which questions the validity of that character in V. johnboborum), but otherwise seem so similar that they are ten- tatively identified with that species. (Dr. Osamu Okamura, BSKU, kindly loaned two paratypes of V. fusca for comparison with southeastern Pa- cific specimens of V. johnboborum.) The subgenus Sokodara is represented by three Pacific species (two in the western Pacific, one in the South Pacific), and two Indian Ocean spe- cies. Ventrifossa misakia is known only from off Japan; V. fusca from the western North Pacific south of Japan; V. johnboborum from the Bis- marck Sea, the western Indian Ocean, and on the Sala y Gomez Ridge; and V. nasuta Smith, 1935, from South Africa and Mozambique. The dis- tributions of V. fusca and V. johnboborum will undoubtedly be extended with more collecting, and it should be interesting to see if and where the two distributions abut or adjoin. Size.— To at least 47.5 cm (Shcherbachev et al. 1986). Distribution. — Known from the western In- dian Ocean, the Bismarck Sea, and the Sala y Gomez Ridge in 540-810 m, but probably ex- tends across the intervening areas on seamounts and oceanic ridges of the Indian Ocean and South Pacific Ocean. Material Examined. — Bismarck Sea: Holotype: AMS 1.15602-002 (85.1 mm HL, 360+ mm TL). Sala y Gomez Ridge (5 spec): ZMMGU 1 7746 (70.3 HL, 340+ TL); 540 m; lchthyandr cr. 5, tr. 53. ZMMGU 17747 (2:66.5-70.7 HL, SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 81 Figure 32. Ventrifossa macrodon new species; paratype, CAS 5 1 796, from Nazca Ridge in 540 m. Drawn by Amy Pertschuk. Scale bar equals 25 mm. 280+-325 +TL); 6 10-620 m; Hercules tr. 70. ZMMGU 17748 (2:72.5-78.5 HL, 361 H — 391 -t- TL); 750-800 m; Prof. Shtok- mancr. 18, sta. 1996. Ventrifossa macrodon new species (Figures 31-33) Ventrifossa sp. 1 : Parin et al. 1981:12 (counts and measure- ments of 24 specimens here described as new). Ventrifossa sp. nova 1 Sazonov and Iwamoto": Parin 1 990: 1 7 (listed from Sala y Gomez Ridge). Parin et al. 1 990:42 (stom- ach contents). Kotlyar and Parin 1990:106, fig. 4b (otolith). Diagnosis.— A species of subgenus Ventrifos- sa with moderately deep body (depth 76-87% HL). Interorbital 20-24% HL, much less than orbit diameter, which goes about 3 in HL; upper jaw 46-50% HL; barbel less than snout length, 23-27% HL; GR-I 17-19 total (inner). No en- larged spinules on body scales. Second spinous ray of first dorsal fin not serrated; ID. 11,9-1 1; IP. il 9— i26; V. 9-10. Leading edge of snout, su- pranarial ridges, and median nasal ridge black- ish; first dorsal fin blackish, without distinct black blotch. Counts and Measurements. — (24 speci- mens; data for holotype in square brackets) 1 D. 11,9-11 [11,9]; IP. il9-i26 (x = i22.07; SD = 2.129) [i23/i24]; V. 9-10 (1 1) [10]; GR-I (outer/ inner) 8-14 [1 1]/(2-3) + (0-1) + (13-16) (16- 19 total) [3 + 14], GR-II (1-3) + (0-1) + (12- 15) (15-18 total) [3 + 14]/(2-3) + (0-1) + (12- 15) (15-18 total) [3 + 14]; scales below midbase ID. 7.5-10 [8]; below 2D. 8.5-1 1 [10]; caeca 50 in 54.5 mm HL paratype; vertebrae 12 (5 spec). Total length 157 +-363+ [350] mm, HL 26.7- 72 [67.7] mm. The following in percent HL: post- rostral 73.7-80 [77.1]; snout 23.2-28.9 [25.6]; preoral 6.5-10.5 [7.8]; orbit 29.2-43.0 [30.7]; in- terorbital 19.6-26.2 [21.7]; suborbital 9.4-13.2 [1 1.4]; postorbital 39.0-46.7 [41.9]; orbit-preop. 38.0-46.6 [44.2]; upper jaw 46.2-54.8 [5 1.7]; gill slit 22.5-29.1 [22.2]; barbel 22.7-33.9 [29.2]; body depth 73-88 [87]; pre-D. 98-1 18 [110]; pre- V. 100-123 [105]; pre-A. 127-152 [137]; V.-A. 26-43 [38]; height ID. 46-77 [-]; ID. base 21.8- 29.1 [27.0]; 1D.-2D. 48.6-76.8 [66.2]; IP. 49- 61 [53.5]; V. 30-46 [32]. Description. — Head and trunk compressed, relatively deep; greatest width of head about equal to width of trunk over pectoral fin bases, slightly more than postorbital length, about three-fifths body depth. Greatest body depth more than 6 in TL, 1.1-1.3 in HL. Snout low, blunt, scarcely protruding beyond the large mouth; distance be- tween supranarial ridges less than (1.1-1.3 into) least interorbital width, 1.4-1.8 into orbit di- ameter. Orbits large, longer than snout, about 3 (2.8-3.4) in HL, but about 2.3-2.7 in juveniles with HL less than 35 mm. Interorbital space flat to slightly convex, width slightly more than that across supranarial ridges. Ridges on head not well developed; rounded or smoothly angular where present. Suborbital region narrow, gently rounded; a ridge line apparent but not demar- cated by stout scales (as in species of Nezumia). Preopercle deep and broadly lobate; posterior border slightly inclined forward and with a shal- low inflection. A relatively deep notch formed along posterior margin of subopercle. Interoper- cle exposed along its posterior and ventral bor- ders. Strongly toothed jaws unrestricted at angles by lip folds; extend to below level of posterior third of orbits. Barbel long, slender, slightly lon- ger than interorbital width. Gill membranes of opposite sides narrowly united under posterior third of orbits, forming a narrow free fold. 82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 25r- 20 10 50 HEAD LENGTH (mm) Figure 33. Comparison of barbel lengths in three species of the Ventrifossa athewdon species complex. Ventrifossa macrodon (circles), V. athewdon (squares), V. sp. (South China Sea) (triangles). Premaxillary dentition consists of moderately wide band of small teeth (4-6 rows wide) flanked by a row of widely spaced, enlarged, conical teeth with arrowhead-shaped tips. Mandibular teeth smaller than enlarged premaxillary teeth and ar- ranged in basically 1 row anteriorly, but 2 irreg- ular rows laterally (small but prominent teeth with arrowhead-shaped tips form a continuous regularly spaced series interspersed laterally and posteriorly with 1 to 3 much smaller teeth that are not exposed above the gum papillae). Thin, finely spinulated scales uniformly cover all of head except over nostril membranes, lips, fins, and gill membranes. Head scales densely covered with short, conical, erect spinules (none enlarged) giving a shagreenlike feel to surface. No scutelike scales on ridges; no enlarged ter- minal scute at snout tip. Scales over suborbital shelf small, generally similar to other scales of head. Body scales rather deciduous; those along base of second dorsal fin lack spinules or only partially covered with spinules; those more ven- trad densely covered with minute, fine, conical, reclined spinules, arrayed in roughly quincunx pattern; 50-70 spinules per scale on larger of these scales in specimens 63.3-70.5 mm HL. Spi- nules of head and body scales flecked with me- lanophores. Fins unsealed except at base. Vent region as for most others of genus (see Marshall 1973:655, fig. 49; Okamura 1970a:74, text-fig. 32). Light organ and alimentary canal similar to that illustrated by Okamura (1970a, text-fig. 34). A small, ill-defined posterior dermal window immediately before anus, connected by a narrow isthmus of black skin to a small oval anterior dermal window situated in a shallow fossa between bases of pelvic fins. Pyloric caeca numerous, thin walled, short (length less than pupil diameter), branched at bases. Ovaries of 63.6 mm HL female (CAS 51795) and 63.3 mm HL female (CAS 51796) large, but eggs small, 0.3-0.4 mm diameter. First dorsal fin relatively low, height less than postrostral length of head, about equal to length pectoral fin; first ray of first dorsal fin rudimen- tary, closely appressed to spinous second ray, which is slender and without serrations; second dorsal fin poorly developed over entire length, begins far posterior to first dorsal. Color in alcohol somewhat swarthy overall; SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 83 abdominal region bluish to violet. Ridges of snout along leading edge, over medial nasal ridge, and over supranarial ridges marked with prominent black edges. A diffuse transverse band extends across each parietal ridge at posterior end of su- praoccipital region. Behind level of orbits, pa- rietal region blackish. Suborbital region darker dorsally than ventrally. An irregularly triangular area of black pigmentation extends postero ven- trally from orbit to preopercle. A blackish hor- izontal streak on ventral aspect of eye just below cornea in some but not in others. Lips, gular membrane, and gill membranes blackish; barbel blackish at base, grayish to pale distally. Papillae of gums blackish; oral valve on roof of mouth edged with black; most linings of mouth whitish; pharynx gray. Gill filaments, gill arches, and hy- oid region of gill cavity pale, remainder of gill chamber blackish. Peritoneal lining finely pep- pered with small melanophores ventrally, dusky to blackish dorsally; stomach pale. Fins blackish; anal fin paler posteriorly. Food. — Stomachs of two individuals (CAS 51795 and 51796) contained unidentifiable re- mains of a fish and a shrimplike crustacean. Pa- rin et al. (1990:42) reported the mysid Paralo- phogaster glaber in the diet of 31% of their 13 specimens; a galatheid crab, Munida sp., in 1 5%. Penaeids, copepods {Pleromammama sp.), am- phipods (Eurisidae), and the fish genus Cyclo- thone were also eaten. Size.— To about 40 cm. Distribution.— Known only from the Sala y Gomez Ridge in 345-770 m. Etymology.— The name macrodon (latinized Greek macros, long, and odus, tooth) refers to the presence of enlarged teeth in the outer row of the premaxillary, a not uncommon character in this genus. The name, however, also reflects the close relationship of the new species to V. macroptera Okamura, 1982, and V. atherodon (Gilbert and Cramer, 1897) by the combination of elements forming each name. Comparisons and Remarks.— The new spe- cies is most closely related to V. atherodon, V. macroptera, and an undescribed species from the South China Sea. The four can be distinguished from each other by characters given in the key that follows. In addition, the new species has a somewhat longer barbel than the other three: 23- 34% HL cf. 12-24% in V atherodon, 18-24.5% in V. macroptera, and 1 7-22% in the South Chi- na Sea species (see also Fig. 33). Key to the species in the Ventrifossa atherodon complex la. Scales below second dorsal fin with dis- tinctly enlarged spinules species (South China Sea) 1 b. No enlarged spinules on scales 2 2a. Pectoral fin 1.3-1.5 in HL; orbits 25- 32.5% HL macroptera (Kyushu-Palau Ridge) 2b. Pectoral fin 1 .7-2.0 in HL; orbits 20-43% HL 3 3a. Length barbel 23-34% HL; outer gill slit 22-29% HL (usually 22-26%) macrodon (Sala y Gomez Ridge) 3b. Length barbel 12-24% HL; outer gill slit 26-32% HL (usually 26-28%) atherodon (Hawaiian Islands) The four species constitute a well-defined spe- cies group in the genus Ventrifossa distinguished by a non-serrated spinous dorsal ray, low blunt snout, high gill raker counts, rather large mouth, and 9-10 pelvic fin rays. The subgeneric name Atherodus Gilbert and Hubbs, 1920, is available for this group. Although a count of V. 9 is com- mon among species of Ventrifossa, a count of 1 is normally encountered only in this species group and in V. ctenomelas (usually 9-10), a species endemic to the Hawaiian Islands where it is sym- patric with V. atherodon. Material Examined. —(138 spec.) all from Sala y Gomez Ridge: Holotype: ZMMGU 18132 (67.7 mm HL, 350 mm TL); 580- 564 m; Prof. Shtokman cr. 18, sta. 1964. Paratypes (39 spec): ZMMGU 17751 (2:38-55 HL, 192- 277+ TL); 770 m; Ichthyandrcr. 5, tr. 52. CAS 51796 (2:63- 69 HL, 290+-305 TL) and ZMMGU 17752 (55 HL, 307 + TL); 540 m; Ichthyandrcr. 5, tr. 53. ZMMGU 17753 (9:30.5- 56 HL, 169+-307 TL) and CAS 51794 (2:48-60 HL, 265- 290 TL); 535-575 m; Ichthyandrcr. 5, tr. 54. ZMMGU 17754 (9:36.3-69 HL, 201 +-367+ TL) and CAS 51795 (3:38-70.5 HL, 210+-353+ TL); 345-540 m; Ichthyandr cr. 5, tr. 55. ZMMGU 17755 (2:59-72 HL, 325+-350+ TL); 550 m; As- tronomer tr. 104. ZMMGU 18133 (9:54-69 HL, 245+-350 TL); same data as for holotype. Other material: ZMMGU 18071 (3:26.7-50 HL, 157-260 TL); 565-555 m; Prof. Mesiatzev cr. 13, tr. 1. ZMMGU 18072 (4:34.3-53 HL, 177-271 TL); 550-560 m; Prof. Mesiatzev cr. 13, tr. 2. ZMMGU 18073 (2:39.5-46.3 HL, 184+-246 TL); 530 m; Prof. Mesiatzev cr. 1 5, tr. 49. ZMMGU 1 8 1 34 ( 1 1 :32- 76 HL, 168-404 TL); 562-545 m; Prof. Shtokman cr. 18, sta. 1965. ZMMGU 18135 (33.7 HL, 181 TL); 570-580 m; Prof. Shtokman cr. 18, sta. 1971. ZMMGU 18136 (13:29-55 HL, 160-260+ TL); 563-590 m; Prof. Shtokman cr. 18, sta. 1976. ZMMGU 18137 (36:29-61 HL, 132+-288+ TL); 545-600 m; Prof. Shtokman cr. 18, sta. 1977. ZMMGU 18138 (11: 29.5-73 HL, 169-383 TL); 750-600 m; Prof Shtokman cr. 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 34. Ventrifossa obtusirostris new species; holotype, ZMMGU 18130 (49 mm HL), from Sala y Gomez Ridge in 739- 760 m. Prof. Shtokman cr. 18, sta. 1996. Photograph by Sergei Dudarev. 18, sta. 1996. ZMMGU 18139 (17:26.3-73 HL, 131 H — 359 TL); 730-790 m; Prof. Shtokman cr. 18, sta. 2018. Ventrifossa obtusirostris new species (Figure 34, 35) " Ventrifossa sp. nova. 3 Sazonov and Iwamoto' 1 7 (listed from Sala y Gomez Ridge). Parin 1990: Diagnosis.— A species of subgenus Ventrifos- sa with snout blunt and rounded, scarcely pro- duced beyond mouth, its length 26.5-28.3% HL; upper jaw 47.7-52% HL; barbel long, 137-166% of orbit diameter, 48-54% of HL; pectoral fins long, 72-78% HL. Color dark overall, first dorsal blackish distally, paler near base; anterior por- tion of anal fin with black margin. Counts and Measurements.— (from 5 spec- imens; holotype data in square brackets) 1 D. II, 1 1 [11]; IP. i21-i23[i21];V. 9-10 [9/10]; GR-I (out- er/inner) 8-10 [9]/(l-2) + (12-14) [2 + 13] (14- 15 total), GR-II 2 + 12/(1-2) + (11-13) [2 + 11] (11-14 total); scales below ID. 10-13 [11], below midbase ID. 7.5-9.5 [9.5], below 2D. 9- 10 [9]. Total length 249 +-294 mm, HL 44.0-51.5 mm. The following in percent HL: postrostral 74.8-78.6 [78.3]; snout 26.5-28.1 [27.2]; preoral 11.7-17.3; interorbital 25.7-27.4 [26.5]; orbit 33.5-36.3 [33.5]; suborbital 13.5-14.3 [13.5]; postorbital 42.7-46.3 [46.3]; orb.-preop. 40.9- 45.5 [44.6]; upper jaw 47.7-52.0 [51.1]; barbel 47.7-54.3 [54.3]; gill slit 26.3-30.4 [30.4]; pre- 1D. 112-122 [123]; pre-A. 128-133 [133]; pre- vent 111-121 [119]; pre-V. 102-110 [111]; V.- A. 26-31 [27.8]; body depth 80-100 [100]; 1D.- 2D. 39-54 [54]; ID. base 30-36 [32.6]; ID. height 77-89 [80]; IP. 72-78 [78]; V. 39-45 [45]. Description. — Body deep, greatest depth al- Figure 35. Ventrifossa obtusirostris new species; paratype, ZMMGU 17756 (51.5 mm HL), Sala y Gomez Ridge in 739- 760 m; Prof Shtokman cr. 18, sta. 1996. Drawn by Tomio Iwamoto. Scale bar equals 25 mm. SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 85 most equal to HL, tapering abruptly behind ab- domen before leveling off to long tail. Head 5.5- 6.0 into TL, width slightly more than postorbital length of head. Snout blunt, low, scarcely pro- truding beyond mouth, its length about equal to its width across lateral angles, about equal to orbit diameter. Orbits large, about 1.4 into post- orbital length, uppermost rim barely entering dorsal profile of head. Interorbital essentially flat, width less than orbit. Suborbital region almost vertical, gently curved in cross-section, the ridge scarcely discernible in well-preserved specimens, the dorsal shelf uniformly wide. Preopercle mar- gin and ridge broadly rounded, not formed into a lobe; interopercle broad, exposed along pos- teroventral margins. Upper jaw about 2 in HL, maxillary extends to below posterior one-third of orbits; rictus extends to below midorbit. Bar- bel notably long, moderately thick, extending be- yond junction of gill membranes, its length more than postorbital, about 2 in HL. Teeth all small, short, conical, rather bluntly pointed, in a broad band in premaxillary with a slightly enlarged outer series. Mandibular teeth similarly small, barely perceptible among gum papillae, in a long narrow band. Scales small, uniformly covering all of body and head except fins, gill membranes, lips, nasal membranes. Scales thin, covered with short, fine, black, partially reclined spinules, rather sparsely and irregularly arranged in an irregularly quin- cunx pattern. Abdominal area anterior to anal fin short, dis- tance pelvic fin base to anal fin origin much less than distance isthmus to pelvic fin base. Vent close between pelvic fins, slightly behind their insertions; periproct region with anterior exten- sion to small teardrop-shaped dermal window situated anterior to pelvic fin origin. First dorsal fin moderate in height, slightly more than postrostral length of head, its second spinous ray finely serrated along leading edge, rather flexible and weak, its tip scarcely pro- longed; second dorsal fin rudimentary for almost entire length; anal fin well developed. Pectoral fin extends to end of abdomen; pelvic fin rather small, outer ray thin, short, extending past base of third or fourth anal fin ray. Origin of pelvics far forward, below hind margin of operculum; pectoral fin origin slightly behind that; first dorsal origin about on same vertical as pectoral or slightly behind, anal fin origin below or forward of hind edge of first dorsal. Color in alcohol dark brownish dorsally, paler on body below midlateral line; bluish to dark gray or blackish over abdomen and head. Lead- ing edge of snout blackish, but inconspicuous because of dark ground color. Pectoral and pelvic fins black; first dorsal overall blackish, but some- what paler along base; anal fin pale except an- terior rays blackish distally. Barbel starkly con- trasting in white. Oral cavity completely pale or white except for thin dark edge of dorsal oral valve and gums of lower jaws; lips black. Gill cavity black along inner wall and peripherally on outer wall; pale in remaining areas; gill arches, rakers, and filaments pale. Size.— To at least 30 cm. Distribution. — Known only from a single collection on the Sala y Gomez Ridge in 750- 800 m. Etymology. — From the Latin, obtusus, blunt, and rostrum, snout, in reference to the short blunt snout of the species as compared with others of the genus. The name is to be treated as a feminine adjective. Comparisons and Remarks.— It is somewhat perplexing that this species was collected only once in the NSG region, despite relatively ex- tensive trawling at appropriate depths. Whether this scarcity of collections can be attributed to unavailability of populations to the sampling gear, insufficient collecting within their primary range, or a general scarcity of individuals is not known. Relationships of V. obtusirostris appear to be closest to three western tropical Pacific species, all of which have a relatively long barbel (about one-half HL): V. saikaiensis Okamura, 1984, V. longebarbata Okamura, 1982, and V. rhipido- dorsalis Okamura, 1984. The first species differs from V. obtusirostris in the following: somewhat fewer pelvic rays (8-9), a broad area of spinule- less scales behind the first dorsal, a slightly short- er pectoral fin (6 1-72% HL), and a slightly short- er barbel (115-133% of orbit). Members of the last two species can be distinguished by their shorter pectoral fin (less than 70% HL). In ad- dition, V. longebarbata has a somewhat shorter postorbital length (39^41%) and narrower inter- orbital (23-24% HL). The long pectoral fin char- acteristic of V. obtusirostris is matched only in V. macroptera, but that species differs in its members having a shorter barbel (20-25% HL, less than orbit diameter), more gill rakers (GR-I 16-18 total inner), a smaller orbit (18-31% HL), and broader interorbital (30-32% HL). 86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Figure 36. Ventrifossa teres new species; holotype, ZMMGU 18131 (52.2 mm HL), from Sala y Gomez Ridge in 730-790 m, Prof. Shtokman cr. 18, sta. 2018, 7 May 1987. Photograph by Sergei Dudarev. Material Examined. — (all from Sala y Gomez Ridge) Ho- lotype: ZMMGU 18130 (46.0 mm HL, 275+ mm TL); Sala y Gomez Ridge, 750-800 m; Prof. Shtokman cr. 1 8, sta. 1 996, trawl 29. Paratypes (same locality as holotype): ZMMGU 17756 (3: 44.5-5 1 .5 HL, 258 +-294 TL) and CAS uncat. (44.0 HL, 249 + TL). Ventrifossa teres new species (Figures 36, 37) Ventrifossa sp. 2: Parin et al. 1981:12 (counts and measure- ments of 4 specimens from Sala y Gomez Ridge). "Ventrifossa sp. nova 2 Sazonov et Iwamoto": Parin 1990:17 (listed from Sala y Gomez Ridge). Kotlyar and Parin 1990: 107, fig. 4b (otolith). Diagnosis.— A species of subgenus Ventrifos- sa with slender body (greatest depth 57-75% HL). Interorbital width 1 8-24% HL, much less than orbit diameter, which goes about 3 in HL; upper jaw 40^5.5% HL; barbel less than snout length, 19-27% HL. First dorsal fin spine weakly ser- rated; ID. 11,9-10; V. 8-9; GR-I 13-15 total (in- ner). Leading edge of snout blackish; no black median nasal streak; first dorsal blackish, with- out distinct black blotch. Counts and Measurements.— (from 38 spec; holotype data in square brackets) ID. 11,(8) 9- 10 [11,10]; IP. i20-i22 (rarely il9 or i23) [i23]; V. 8-9 [9]; GR-I (outer/inner) 7-12 [10]/(l-2) + (0-1) + (10-13) [1 + 1 + 12] (12-15 total), GR- 11 (1-2) + (0-1) + (9-12) [2+11] (1 1-14 total)/ (1-2) + (0-1) + (9-12) [1 + 1 + 12] (11-14 total); caeca 34-36 (4 spec); abdominal verte- brae 1 1 . Total length 1 3 1-239 mm, HL 23.8-52.2. The following in percent HL: postrostral 72-78 [75.7]; snout 26.4-3 1 .8 [26.8]; preoral 1 1 .2-1 7.6 [ 1 3.6]; orbit 31.6-37.3 [31.4]; interorbital 18.5-23.9 [18.1]; suborbital 7.2-12.3 [9.2]; postorbital 37.0- 41.4 [38.3]; orbit-preop. 32.9-38.5 [36.8]; upper jaw 38.7-44.7 [42.1]; gill slit 18.8-25.9 [21.1]; barbel 19.4-26.7 [27.4]; body depth 57-75 [71]; pre-D. 103-1 12 [1 1 1]; pre-V. 99-122 [107]; pre- A. 126-159 [144]; V.-A. 27^19 [42]; ID. height 47-68 [49]; ID. base 20.5-30.2 [27.8]; 1D.-2D. 48-80 [63]; IP. 49-61 [54]; V. 28-37 [34]. Description.— A slender-bodied species of Ventrifossa, width across pectoral fin bases about three-fourths body depth; greatest head width three-fourths its depth; greatest body depth about 1.3-1.8 into HL, 6.4-8.7 into TL. Nape and first dorsal fin base low. Snout low, conical in lateral profile; relatively blunt and narrow in dorsal view, width across anterolateral angles slightly greater than interorbital width; length about 1.2 into or- bit. Orbits large, about 3 in head, about 1.3 into postorbital length. Interorbital space probably relatively flat in live specimens, but slightly con- cave in examined preserved specimens. Subor- bital region traversed by a low, rounded ridge, the upper and lower surfaces of region meeting at an obtuse angle (in cross-section); upper shelf area not markedly narrowed anteriorly. Pre- opercle broadly lobate; interopercle broadly ex- posed posteriorly beyond preopercle. Upper jaw long, about 2.2-2.6 into HL; maxillary extends to below posterior one-third of orbits; rictus (mouth cleft) not restricted posteriorly. Barbel slender, moderately long, about equal to inter- orbital width. Dentition of premaxillary in a moderately wide band; inner teeth very small, bordered by an out- er series of slightly enlarged, widely spaced, con- ical, recurved teeth. Mandibular teeth all small, in 2 irregular series. Tips of outer premaxillary teeth and all mandibular teeth shaped like ar- rowheads. Scales uniformly cover all of head except for nostril membranes, lips, gill membranes, and a narrow median triangular area above upper lip. Head scales thin, densely covered with short, SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 87 Figure 37. Venthfossa teres new species; paratype, ZMMGU 17759 (39.5 mm HL), Sala y Gomez Ridge in 354-540 m, Ichthyandr cr. 5, trawl 55. Drawn by Amy Pertschuk. Scale bar under head equals 10 mm, that under scale equals 1.0 mm. conical, erect spinules, none enlarged. No scute- like scales on ridges; no enlarged terminal scute at snout tip; scales over suborbital shelf not coarse or scutelike. Most body scales missing on type specimens; those remaining dorsally below sec- ond dorsal fin covered with fine, conical, reclined spinules, arrayed in roughly quincunx pattern; 24-26 spinules per scale on larger of these scales. Spinules of head and body scales flecked with melanophores. Fins unsealed except at base. The 43.5 mm HL specimen from CAS 51797 had large ovaries filled with eggs of different sizes, some measuring as much as 1 .4 mm in diameter. Pyloric caeca relatively few for a Venthfossa (ac- curate number could not be determined), length slightly less than interorbital diameter. Vent- periproct region much like that of other members of genus; anterior dermal window of light organ small, circular, slightly anterior to pelvic bases. First dorsal fin relatively short, length less than postrostral length of head; first spinous ray close- ly appressed to second spinous ray and not pro- truding through integument. Second spinous ray weakly denticulate, the short denticles confined to distal half. Paired fins moderate in size, neither pelvics nor pectorals much longer than postor- bital length of head. Color in alcohol overall pale with widely spaced dots on ventral half of trunk and tail; abdomen bluish. Branchiostegal and gular membranes blackish on exposed surfaces. Lips edged with thin black margin. Diffuse blackish blotches over posterior part of posttemporal and ventrally on preopercle at junction with infraorbitals. Barbel completely white to base. Edges of mandibular ramus black. Leading edge of snout posteriorly to anterior end of suborbital ridge blackish; su- pranarial ridges blackish, this coloration extend- ing onto tops of orbital rim. Outer edge of gill cover with a diffuse blackish margin; upper edge of opercle with a thin black margin; a diffuse horizontal black streak slightly below upper edge of opercle. Oral cavity completely pallid; bran- chial cavity generally pallid with scattered punc- 88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 tations and blackish outer margins. Gill fila- ments pale; arches and rakers dusky. Paired fins black; first dorsal blackish with no pronounced blotches, but base pale; anal and second dorsal pale. Size.— A small species, attains about 24 cm. Distribution. — Known only from the Sala y Gomez Ridge, in 345-610 m. Etymology.— From Latin, teres, terete, in ref- erence to the slender, cylindrical body. Comparisons and Remarks. — Ventrifossa teres resembles V. mucocephalus Marshall, 1973, from the tropical western Atlantic in its low head profile, relatively terete body, and small chin bar- bel. The two species differ in the following: (1) V. teres usually has V. 9, rarely 8, cf. V. 8, seldom 7 or 9 in V. mucocephalus; (2) ID. 11,9 or 10 in V. teres cf. ID. 11,1 1-12 (rarely 11,10); (3) inter- orbital 18.5-23.9% of HL in V. teres cf. 23-28% HL; (4) orbit to preopercle distance 32.9-38.5% HL cf. 40.5-46.0%; (5) black margin along su- pranarial ridges in V. teres, but unmarked in V. mucocephalus; and (6) V. teres is a smaller spe- cies attaining about 24 cm TL, cf. more than 4 1 cm in V. mucocephalus. Ventrifossa teres also resembles V. nasuta (Smith, 1964) from southern Africa in its low head profile and relatively terete shape, but the two differ markedly in the latter having a stout terminal snout scute and lacking distinctive head markings. Several morphomet- ric and meristic features also distinguish the two. Material Examined.— (39 spec, all from Sala y Gomez Ridge) Holotype: ZMMGU 18131 (mature 2, 52.2 mm HL, 237 mm TL); 730-790 m; Prof. Shtokman cr. 28, sta. 2018. Paratypes: CAS 51797 (2:39.0-43.5 HL, 174-184 TL); 770 m; IchthyandrcT. 5, tr. 52. ZMMGU 17758 (38 HL, 188 TL); 535-575 m; Ichthyandr cr. 5, tr. 54. ZMMGU 17759 (39.5 HL, 196 TL); 345-540 m; Ichthyandr cr. 5, tr. 55. ZMMGU 17760 (2:41-48.5 HL, 196+-226+ TL); 610 m; Hercules tr. 68. ZMMGU 17761 (13:36.5-47. 5 HL, 157+-214+ TL); 750- 800 m; Prof. Shtokman cr. 18, sta. 1996. ZMMGU 18067 (4: 23.8^19.2 HL, 131-239 TL); 562-545 m; Prof. Shtokman cr. 18, sta. 1965. CAS uncat. (5:41-45 HL, 184+-202+ TL) and ZMMGU 18068 (8:38^16.5 HL, 172.5+-207+ TL); same data as for holotype. ZMMGU 18069 (2:31.1-32.7 HL, 143+-143 + TL); 545-600 m; Prof. Shtokman cr. 18, sta. 1977. Biogeographical Considerations Table 5 lists the 25 species of macrourids known from NSG and compares their distribu- tions in various parts of the world's oceans. Only two species are known from the continental slope of South America: Caelorinchus immaculatus and Nezumia convergens, and the identification of the latter from a single specimen is as yet tentative. Caelorinchus immaculatus is a representative of a southern temperate fauna, as its relationships appear closest to C. karrerae of the southeastern Atlantic and southern part of the Indian Ocean, from Africa to Australia. None of the NSG spe- cies are common to the tropical eastern Pacific fauna, although the widespread Malacocephalus laevis is also known from the northeastern Pacific in slope waters of southern California and on seamounts off California and Mexico. Eight of the 25 species are widespread tropical or subtropical taxa: Macrouroides inflaticeps, Squalogadus modificatus, Cetonurus crassiceps, Coryphaenoides paradoxus, Hymenocephalus gracilis, Malacocephalus laevis, Nezumia propin- qua, and Trachonurus villosus (but unresolved taxonomic problems exist with Trachonurus). Eight species are endemic to NSG and four (possibly five) others (Caelorinchus spilonotus, Hymenocephalus striatulus, Mataeocephalus acipenserinus, Pseudocetonurus septifer, and pos- sibly Gadomus sp. cf. melanopterus) are known only from NSG and the Hawaiian Islands. The ties to the Hawaiian fauna are strong, with 1 1 species (three are questionably included here) common to the two areas, the most for any re- gion. The western tropical Pacific and Indian Ocean also show a high affinity, with eight and nine species in common, although all but Ven- trifossa johnboborum are widespread species. Those areas are followed closely by the Atlantic Ocean, with seven species in common with the NSG. Thus, despite the proximity of the Nazca Ridge to the mainland coast of Peru, the relationship of the fauna of the Nazca and Sala y Gomez ridges is clearly to the west, in the tropical central and western Pacific. This is in line with the dis- tributions of other deep-sea fish and invertebrate taxa known from this area (see Parin 1 990; Parin et al. 1981; Nesis 1 990) and also the shore fauna of many eastern Pacific islands (Rosenblatt et al. 1972; Springer 1982). Wilson et al. (1985) dis- cuss this relationship in the fauna of seamounts of the central North Pacific. The 32% endemism of the grenadier fauna (8 of 25 spp.) is somewhat lower than the 46% endemism (65 of 141 spp.) recorded by Parin (1990) for the entire NSG fish fauna. That so many of the grenadiers repre- sented are circumglobal species accounts for the lower percentage of endemic species. It can be expected that more drags made below 1,500 m SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 89 Table 5. List of the grenadiers of the Nazca and Sala y Gomez ridges, comparing their distributions in different areas. Species denoted with an asterisk (*) may have still-unresolved taxonomic problems and may involve more than one species. A question mark (?) following a plus sign ( + ) indicates the possible occurrence of the species in the region. Abbreviations: A— Atlantic Ocean; E— endemic to Nazca and Sala y Gomez ridges; ESP— continental eastern South Pacific; HI— Hawaiian Island region; 10— Indian Ocean; Nazca— Nazca Ridge; SyG = Sala y Gomez Ridge; WP— western tropical Pacific; WSP— Australian- New Zealand region (western South Pacific). will result in the capture of other species that will further reduce the percentage of endemics, be- cause abyssal grenadiers are more broadly dis- tributed than their shallow-water counterparts, as a general rule. The relative richness of the two ridges is clearly skewed towards the Sala y Gomez Ridge, with 2 1 species of grenadiers represented, compared with only eight on the Nazca Ridge (see Table 5). Nezumia sp. cf. convergens was taken only in the region between the two ridges; Macrouroides inflaticeps was taken in the intermediate area and on the Nazca Ridge; Cetonurus crassiceps and Hymenocephalus sp. cf. aterrimus were taken on both ridges and in the intervening area. Parin ( 1 990) considered the Nazca fauna a depauperate one containing a few peculiar elements that make it distinctive from that of the Sala y Gomez. Nesis (1990), using cephalopod data, confirmed the existence of a zoogeographical boundary on the southwestern side of the Nazca Ridge at about 84°-85°W. Comparison of the grenadiers of the NSG and Hawaiian Islands shows some interesting par- allels and contrasts (Table 6). The Hawaiian Is- land region has a slightly greater number of re- corded species (29) than NSG (25), and the number of endemics is notably higher (15, or 52%, compared with 9 and 36%). This high en- demism of the Hawaiian fauna is reflected in the few species shared in common with the western tropical Pacific— the number of NSG species in this category is twice that of the Hawaiian Islands (8 vs. 4). Strangely, however, the number of In- dian Ocean and Altantic Ocean species shared in common with NSG on the one hand, and the Hawaiian Islands on the other hand, are com- parable. If we exclude five wide-ranging (and problematic) species {Cetonurus crassiceps, Cor- yphaenoides paradoxus, Hymenocephalus ater- rimus, Malacocephalus laevis, and Trachonurus villosus), the contrasts become even more inter- esting. In the Hawaiian fauna, Kuronezumia bu- bonis and Mesobius berryi remain as the only 90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 Table 6. List of the grenadiers of the Hawaiian Islands region, comparing their distributions in different areas. Species denoted with an asterisk (*) may have still-unresolved taxonomic problems and may involve more than one species. A question mark (?) following a plus sign ( + ) indicates the possible occurrence of the species in the region. Abbreviations: A— Atlantic Ocean; E— endemic to Hawaiian Islands region; ESP— continental eastern South Pacific; IO— Indian Ocean; Nazca— Nazca Ridge; SyG— Sala y Gomez Ridge; WP— western tropical Pacific; WSP— Australian-New Zealand region (western South Pacific). two species also found in the Indian and Atlantic oceans. Of the NSG fauna, only Hymenocepha- lus gracilis and Squalogadus modificatus remain. That the two faunas are closely related is shown in the number of species they share exclusively: Gadomus melanopterus, Caelorinchus spilono- tus, Hymenocephalus striatulus, Mataeocephalus acipenserinus, and Pseudocetonurus septifer. It thus appears that the two faunas share a host of wide-ranging species as well as several unique ones, but each has been isolated long enough to develop a large percentage of endemic elements. Okamura and Yatou (in Okamura et al. 1982) reported on the grenadiers from the Kyushu-Pa- lau Ridge and Tosa Bay, collected by the Fish- eries Agency of the Government of Japan (Table 7). The Kyushu-Palau Ridge extends in a south- easterly direction from the island of Kyushu to the Palau Islands at a latitude of approximately 6°N. The ridge area investigated was confined to approximately 25°-30°N. Of the 25 species re- corded, 15 were confined to the Kyushu-Palau Ridge, and 10 others were taken on the conti- nental slope off Tosa Bay (Shikoku Island, Japan) and other continental margins. Although the coverage area and collection depths were much more limited in the Japanese surveys than in the Soviet surveys of the NSG, some interesting comparisons can be made. There were three species that were possibly the same in the two areas: Malacocephalus nipponensis (?=M laevis), Nezumia propinqua, and Tracho- nurus villosus (?). Six of the 1 5 species (40%) were endemic to the Kyushu-Palau Ridge, and these were equally divided among Caelorinchus and Ventrifossa. The two genera were represented in SAZONOV AND IWAMOTO: GRENADIERS OF THE NAZCA AND SALA Y GOMEZ RIDGES 91 the NSG fauna by four and three endemic spe- cies. The Kyushu-Palau Ridge, although being relatively closely adjacent to an exceeding rich faunal region (Japan and the Philippines), none- theless has a distinctive fauna, which appears to be less diverse than the faunas of the Hawaiian Archipelago and the Nazca and Sala y Gomez ridges. The propensity for the genera Caelorin- chus and Ventrifossa to have the most species in these ridge areas is notable, although the thought is tempered with the knowledge that Caelorin- chus is by far the most diverse genus of grena- diers, with more than 100 species. Acknowledgments We thank the many people who helped in var- ious ways in the preparation of this paper. Fore- most, we thank Nikolai V. Parin, head of the Laboratory of Oceanic Ichthyofauna, for initi- ating this cooperative study, for his encourage- ment and advice, and for his help with materials, facilities, personnel, and other things too nu- merous to mention. Yuri N. Shcherbachev (IOAN) contributed enormously with his advice, knowledge of the group and collections, and as- sistance with manuscript review, references, data gathering, and facilities arrangements (the last especially for TI). Many people were extremely hospitable and helpful to TI during his visits to their institutions. These include, but are not lim- ited to: E. Karmovskya, Y. N. Shcherbachev, V. Tchuvasov, and others of IOAN; I. A. Verighina and others of ZMMGU; A. Andriashev, V. V. Barsukov (deceased), A. V. Neyelov, and others of ZIN; S. Jewett, L. R. Parenti, V. G. Springer, R. P. Vari, (USNM); D. M. Cohen, R. Feeney, R. J. Lavenberg, J. Seigel, and C. C. Swift (LACM); J. E. Randall and A. Sugimoto (BPBM); K. Hartell (MCZ); O. Crimmen, G. Howes, A. Wheeler, and (BMNH); O. Okamura (BSKU). Technical assistance at CAS was provided by D. G. Catania, J. Fong, and P. M. Sonoda; T. Moritz and B. Edgar provided references; S. Tatro trans- lated the abstract into Spanish. Photographs of specimens were provided by S. K. Middleton (CAS) and S. Dudarev (IOAN). Many individ- uals were involved in collecting specimens, but we especially acknowledge the efforts of G. A. Golovan', N. P. Pakhorukov (both from the Se- vastopol Laboratory), and A. N. Kotlyar (IOAN, formerly with VINRO). Although we began this study many years ago, its completion was aided Table 7. Grenadiers of the Kyushu-Palau Ridge and Tosa Bay (after Okamura et al. 1982). Species found on the Kyushu-Palau Ridge (species endemic to ridge marked with an asterisk*): 1. Gadomus colletti Jordan and Gilbert, 1904 2. Caelorinchus gilberti Jordan and Hubbs, 1925 3. C. hexafasciatus Okamura, 1982* 4. C. longicephalus Okamura, 1982* 5. C. matsubarai Okamura, 1982* 6. Hymenocephalus longiceps Smith and Radcliffe, 1912 7. H. striatissimus Jordan and Gilbert, 1904 8. H. lethonemus Jordan and Gilbert, 1904 9. Malacocephalus nipponensis Gilbert and Hubbs, 1916 10. Nezumia propinqua (Gilbert and Cramer, 1897) 11. Trachonurus villosus (Giinther, 1877) 12. V.fusca Okamura, 1982* 13. V. japonica (Matsubara, 1943) 14. V. longibarbata Okamura, 1982* 15. V. macroptera Okamura, 1982* Tosa Bay species not found on Kyushu-Palau Ridge: 1. Caelorinchus anatirostris Jordan and Gilbert, 1904 2. C. smithi Gilbert and Hubbs, 1916 3. C. japonicus (Temminck and Schlegel, 1884) 4. Coryphaenoides nasutus Giinther, 1877 5. C. marginatus Steindachner and Doderlein, 1884 6. Kuronezumia dam (Gilbert and Hubbs, 1916) 7. Nezumia condylura Jordan and Gilbert, 1904 8. N. proxima (Smith and Radcliffe, 1912) 9. Ventrifossa garmani (Jordan and Gilbert, 1904) 10. V. nigrodorsalis Gilbert and Hubbs, 1916 by a 1988 visit to the USSR by TI, sponsored by the National Academy of Sciences/National Research Council through its Soviet and East European Program. Director G. E. Schweitzer and Program Associate C. M. Turzak are thanked for their assistance. The CAS In-House Research Fund provided support to TI for museum visits. Resumen Hay registrado veinticinco especies de grana- deros desde las Cordilleras submarinas de Nazca y Sala y Gomez en el sur del Pacifico Oriente. Las especies nuevas incluyen: Caelorinchus im- maculatus, C. multifasciatus, C. nazcaensis, C. spilonotus, Hymenocephalus neglect issimus, H. semipellucidus, Kuronezumia pallida, Ventrifos- sa macrodon, V. teres, y V. obtusirostris. Cae- lorinchus immaculatus es muy semejante a C. karrerae desde el sur el Atlantico Oriente y el mar de las Indias. El complejo de H. striatissimus esta examinado usando informaciones nuevas. Hymenocephalus semipellucidus y H. neglectis- simus parecen pertenecer a este complejo. El sub- 92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 48, No. 2 especie de H. s. hachijoensis de Japon esta ele- vado a la categoria del especie pleno. Kuronezumia, considerado antes como subge- nera de Nezumia, esta redefinado y elevado a la categoria de genero incluyendo K. pallida, K. bu- bonis, K. leonis, K. macronema, K. dara, y dos especies no descritos. A despecho de la prox- imidad de las Cordilleras submarinas a la cuesta del Peru, los afines de la fauna asociada estan hacia del oeste, particularmente al Pacifico Oc- cidental y las Islas Hawaii. Ocho de los veinti- cinco especies de estas Cordilleras submarinas eran definitivamente conocidos de la proximi- dad de las Islas Hawaii: Caelorinchus spilonotus, Cetonurus crassiceps* Coryphaenoides paradox- us* H. striatulus, Malacocephalus laevis* Ma- taeocephalus acipenserinus, Nezumia propin- qua* y Pseudocetonurus septifer. Tres otros especies cuyos identificaciones no son determi- nados pueden ser parte o tener afines en la fauna de las Islas Hawaii: Gadomus sp. cf. melanop- terus, Hymenocephalus sp. cf. aterrimus, and Trachonurus villosus*! Los cuatro especies mar- cados con asterisco estan distribuidos amplia- mente por los oceanos Pacifico, Indio y Atlan- tico. Malacocephalus laevis se conoce de los taludes continentales en el sur de California y por las montafias submarinas cerca de Baja Cal- ifornia, pero nunca se los encuentra por las cues- tas de America Central y America del Sur. Cae- lorinchus immaculatus tambien se nota de America del Sur; Nezumia convergens esta re- presentada discutiblemente en un especimen un- ico de la cordillera submarina de Sala y Gomez. Literature Cited Amaoka, K., K. Nakaya, H. Araya, and T. Yasui, eds. 1983. Fishes from the North-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fish. Resource Conser. Assoc, Tokyo. 371 pp. Arai, T. and P. J. McMillan. 1982. A new macrourid fish, Coelorinchus biclinozonalis from New Zealand, and rede- scription of C. australis from Australia. Japan. J. Ichthyol. 29(2): 115-126. Barnard, K. H. 1925. Descriptions of new species of marine fishes from S. Africa. Ann. Mag. Nat. 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