Synopsis of the grenadier fishes (Gadiformes; Teleostei) of Taiwan

Abstract

Species of grenadier fishes (Order Gadiformes) in Taiwan are reviewed. The species list of Shao et al. (2008) is revised. A total 71 species in 21 genera and 3 families is recognized, including 5 species that are tentatively identified and 5 species, Coelorinchus hexafasciatus, C. cf. macrorhynchus, C. cf. notatus, Hymenocephalus papyraceus, and Ventrifossa sazonovi, that are first records for Taiwan. Ventrifossa fusca is recognized as a junior synonym of V. misakia. Keys to families, genera and species are provided. Species descriptions are based mainly on Taiwanese specimens but supplemented with specimens from various other sources. Figures of species firstly reported by Shao et al. (2008) are provided.

Full text

Synopsis of the Grenadier Fishes
(Gadiformes; Teleostei) of Taiwan
Tomio Iwamoto1, Naohide Nakayama2, Kwang-Tsao Shao3, and Hsuan-Ching Ho4
1Section of Ichthyology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco,
CA 94118, USA; 2Laboratory of Marine Biology, Faculty of Science, Kochi University,
2–5–1 Akebono-cho, Kochi, 780–8073, Japan; 3Research Center for Biodiversity, Academia Sinica No.
128, Sec. 2, Academia Road, Nankang, Taipei 115, Taiwan; 4National Museum of Marine Biology and
Aquarium and Institute of Marine Biology, National Dong Hwa University, No. 2, Houwan Road,
Checheng, Pingtung, 944 Taiwan; Email: hohc@nmmba.gov.tw
Table of Contents
Abstract.....................................................................34
Introduction..................................................................34
MaterialandMethods..........................................................35
Results......................................................................36
The diversity and current status of grenadier species in Taiwan . . . . . . . . . . . . . . . . . . . . . . . 36
Systematics..................................................................36
KeytothegrenadierfamiliesinTaiwan..........................................36
FamilyBathygadidae.........................................................37
Key to the genera and species of Bathygadidae in Taiwan . . . . . . . . . . . . . . . . . . . . . . . . . 37
Genus Bathygadus Günther,1878...........................................37
B. antrodes (JordanandGilbert,1904)...................................37
B. furvescens Alcock,1894............................................38
B. garretti GilbertandHubbs,1916.....................................39
B. nipponicus (JordanandGilbert,1904).................................40
Bathygadus sp.indet. ................................................41
Genus Gadomus Regan,1903..............................................41
G. colletti JordanandGilbert,1904 .....................................41
G. magnifilis GilbertandHubbs,1920...................................42
G. cf. multifilis [sensu (Günther,1887)]..................................43
FamilyMacrouridae .........................................................44
Key to the genera and some species of Macrouridae in Taiwan . . . . . . . . . . . . . . . . . . . . . 44
Genus Cetonurus Günther,1887............................................46
C. globiceps Vaillant,1884............................................46
Genus Coelorinchus Giorna,1810..........................................47
Key to the species of Coelorinchus inTaiwan...............................48
C. anatirostris JordanandGilbert,1904..................................50
C. asteroides Okamura,1963 ..........................................50
C. brevirostris Okamura,1984 .........................................51
C. cingulatus GilbertandHubbs,1920...................................52
C. divergens OkamuraandYatou,1984..................................53
C. formosanus Okamura,1963.........................................54
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Reprint from Proceedings of the California Academy of Sciences, ser. 4, vol. 62, 15 April 2015. © 2015

C. fuscigulus Iwamoto,HoandShao,2009...............................55
C. hexafasciatus Okamura,1982 .......................................56
C. hubbsi Matsubara,1936............................................57
C. japonicus (TemminckandSchlegel, 1846) .............................57
C. kamoharai Matsubara,1943.........................................58
C. kishinouyei JordanandSnyder,1900..................................59
C. leptorhinus Chiou,ShaoandIwamoto,2004............................59
C. longissimus Matsubara,1943........................................60
C. macrochir (Günther,1877)..........................................61
C. cf. macrorhynchus [sensu Smith and Radcliffe, 1912] . . . . . . . . . . . . . . . . . . . . 62
C. multispinulosus Katayama,1942.....................................63
C. cf. notatus [sensu SmithandRadcliffe,1912]...........................64
C. parallelus (Günther,1877)..........................................65
C. productus GilbertandHubbs,1916...................................66
C. sheni Chiou,Shao,andIwamoto,2004................................67
C. smithi GilbertandHubbs,1920......................................67
C. cf. spinifer [sensu GilbertandHubbs,1920]............................68
Genus Coryphaenoides Gunnerus,1874......................................69
Key to the species of Coryphaenoides inTaiwan.............................69
Coryphaenoides cf. asper [sensu Günther,1877]...........................69
C. microps (SmithandRadcliffe,1912)..................................69
C. nasutus Günther,1877 .............................................71
C. rudis Günther,1878 ...............................................73
Genus Hymenocephalus Giglioli,1882.......................................73
Key to the species of Hymenocephalus inTaiwan............................75
H. lethonemus JordanandGilbert1904..................................75
H. longiceps SmithandRadcliffe,1912..................................75
H. papyraceus JordanandGilbert,1904..................................76
H. striatissimus striatissimus Jordan and Gilbert, 1904 . . . . . . . . . . . . . . . . . . . . . . 77
Genus Hymenogadus GilbertandHubbs,1920 ................................78
H. gracilis GilbertandHubbs,1920.....................................78
Genus Kumba Marshall,1973..............................................79
Key to the species of Kumba inTaiwan....................................79
K. gymnorhynchus IwamotoandSazonov,1994 ...........................79
K. japonica (Matsubara,1943).........................................80
K. punctulata IwamotoandSazonov,1994 ...............................80
Genus Kuronezumia Iwamoto,1974.........................................82
K. dara (GilbertandHubbs,1916)......................................82
Genus Lucigadus GilbertandHubbs,1920 ...................................83
L. nigromarginatus (SmithandRadcliffe,1912) ...........................84
Genus Macrosmia Merrett, Sazonov and Shcherbachev, 1983 . . . . . . . . . . . . . . . . . . . . 84
M. phalacra Merrett, Sazonov and Shcherbachev, 1983 . . . . . . . . . . . . . . . . . . . . . 85
Genus Malacocephalus Günther,1862.......................................86
M. nipponensis GilbertandHubbs,1916.................................86
Genus Mataeocephalus Berg,1898 .........................................87
Key to the species of Mataeocephalus inTaiwan.............................88
M. (Mataeocephalus)cristatus Sazonov, Shcherbachev and Iwamoto, 2003 . . . . . 88
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M. (Hyostomus)hyostomus (Smith and Radcliffe, 1912) . . . . . . . . . . . . . . . . . . . . . 88
Genus Nezumia Jordan,1904..............................................90
Key to the species of Nezumia inTaiwan...................................90
N. condylura JordanandGilbert,1904...................................91
N. evides (GilbertandHubbs,1920).....................................92
N. proxima (SmithandRadcliffe,1912)..................................93
N. spinosa (GilbertandHubbs,1916) ...................................94
N. cf. coheni [sensu IwamotoandMerrett,1997 ...........................95
Genus Pseudocetonurus Sazonov and Shcherbachev, 1982 . . . . . . . . . . . . . . . . . . . . . . . 97
Pseudocetonurus cf. septifer Sazonov and Shcherbachev, 1982 . . . . . . . . . . . . . . . 97
Genus Pseudonezumia Okamura,1970.......................................98
P. pusilla (Sazonov and Shcherbachev, 1982) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Genus Sphagemacrurus Fowler,1925 .......................................99
Key to the species of Sphagemacrurus inTaiwan ...........................100
S. pumiliceps (Alcock,1894) .........................................100
S. richardi (Weber,1913) ............................................101
Genus Spicomacrurus Okamura,1970......................................102
S. kuronumai (Kamohara,1938).......................................103
Genus Trachonurus Günther,1887.........................................103
Key to the species of Trachonurus inTaiwan...............................103
T. sentipellis GilbertandCramer,1897..................................104
T. villosus (Günther,1877) ...........................................104
Genus Ventrifossa GilbertandHubbs,1920..................................106
Key to the species of Ventrifossa inTaiwan................................107
V. divergens GilbertandHubbs,1920...................................107
V. garmani (JordanandGilbert,1904)..................................109
V. longibarbata Okamura,1982 .......................................110
V. macroptera Okamura,1982.........................................111
V. misakia JordanandGilbert,1904....................................112
V. nigrodorsalis GilbertandHubbs,1920................................114
V. rhipidodorsalis Okamura,1984......................................115
V. saikaiensis Okamura,1984.........................................116
V. sazonovi IwamotoandWilliams,1999................................117
FamilyMacrouroididae........................................................118
Genus Squalogadus GilbertandHubbs,1916 ................................118
S. modificatus GilbertandHubbs,1916.................................118
Acknowledgments............................................................119
LiteratureCited..............................................................119
Appendix...................................................................125
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 33

Species of grenadier fishes (Order Gadiformes) in Taiwan are reviewed. The species
list of Shao et al. (2008) is revised. A total 71 species in 21 genera and 3 families is
recognized, including 5 species that are tentatively identified and 5 species, Coelor-
inchus hexafasciatus,C. cf. macrorhynchus,C. cf. notatus,Hymenocephalus
papyraceus, and Ventrifossa sazonovi, that are first records for Taiwan. Ventrifossa
fusca is recognized as a junior synonym of V. misakia. Keys to families, genera and
species are provided. Species descriptions are based mainly on Taiwanese specimens
but supplemented with specimens from various other sources. Figures of species
firstly reported by Shao et al. (2008) are provided.
KEYWORDS: Pisces, taxonomy, Bathygadidae, Macrouridae, Macrouroididae, Taiwan
Grenadiers constitute the largest group of deepwater demersal fishes found at continental-
slope depths between 200 m and 4000 m of all oceans. We here use the common name grenadier
to include members of four distinct families—Bathygadidae, Macrouridae, Macrouroididae, and
Trachyrincidae—of the order Gadiformes, in which the highly valued codfishes of the family Gadi-
dae are a part. Macrouridae is the largest family in the order with more than 350 species classified
into about 30 genera. Grenadiers are almost exclusively benthopelagic in habit, with only a few
species having taken up a bathypelagic existence. Pelagic captures of some normally benthopelag-
ic species suggest that individuals may on occasion forage well off the bottom, and these forays
may be vertical or offshore movements.
Recent deepwater trawl collections from Taiwanese waters have shown an astonishing diver-
sity of benthopelagic fishes, among which the grenadiers are represented by the highest number of
species. Shao et al. (2008) recorded 71 species of grenadiers categorized into 19 genera and three
families; 33 of those species (in 10 genera) were newly recorded from Taiwan and were captured
only since 2001. Our subsequent re-examination and re-evaluation of specimens have changed the
composition of the list slightly, but retained the same number of species (70 if Coelorinchus pro-
ductus is accepted to be a synonym of C. anatirostris) classified in 21 genera (2 genera formerly
treated as subgenera Hymenogadus and Spicomacrurus). Shao et al. (2008) analyzed the species
composition of the group as related to depth and geography in Taiwanese waters and found a dis-
tinct separation at 600 m and 1000 m. The principal factors influencing grenadier species compo-
sition in the catches were depth, geographic region, and net type.
Documenting the diversity of Taiwan’s continental-slope fauna is highly important at this time
because of the recently developed deepwater trawl fishery in the country, which has been exploit-
ing the offshore resources heavily. The primary species targeted in this fishery are the shrimps and
prawns, with the more valuable food fishes in the catches hand-selected out at the dockside mar-
kets. Whatever bycatch remains, and that includes many species of grenadiers, is ground up to be
used as feed stock in the country’s extensive aquaculture and poultry industries. It is doubtful that
such heavy exploitation of these resources can continue without a dramatic depletion of the stocks,
owing to the presumed slow growth rate of most deepwater animals. The classic, well-document-
ed depletion of the deepwater orange roughy Hoplostethus atlanticus stocks off New Zealand and
Australia is mirrored in the North Atlantic by the stocks of roundnose grenadier Coryphaenoides
rupestris and roughhead grenadier Macrourus berglax, which by some estimates can be considered
as threatened in the northwestern part of the Atlantic (Devine et al. 2006; Devine and Haedrich
2008). Merrett and Haedrich (1997) provide detailed case examples as well as an overview of
development, history, and future of deepwater demersal fisheries in their highly informative book
Deep-sea demersal fish and fisheries.
The many species of grenadiers brought into Taiwanese ports and the difficulty in identifying
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Series 4, Volume 62, No. 3

them have prompted us to develop this identification guide, which we hope will allow its users to
accurately identify and document one of the chief components of most deepwater catches. Such
identifications are crucial for obtaining accurate statistical information on Taiwan’s fishery
resources and ultimately to developing regulations that will ensure their sustainablility into the dis-
tant future, as well as maintaining that part of the rich biodiversity heritage of the country.
MATERIAL AND METHODS
Data herein provided are from the specimens we have examined, mostly from Taiwan and
Japan, but sometimes from other areas as indicated in the Materials Examined sections. Specimens
are primarily from the Academia Sinica, Research Center for Biodiversity (formerly Institute of
Zoology) (ASIZ), with supplemental material from the California Academy of Sciences (CAS),
Laboratory of Marine Biology, Faculty of Science, Kochi University, Kochi (BSKU); Hokkaido
University Museum, Hakodate, Hokkaido (HUMZ); National Museum of Marine Biology and
Aquarium (NMMB-P), Kenteng, Taiwan; National Science Museum, Tokyo (NSMT); and United
States National Museum of Natural History, Washington DC (USNM). Methods for making counts
and measurements follow Iwamoto (1970) and Iwamoto and Sazonov (1988), and further elaborat-
ed on by Iwamoto and Williams (1999). The reader is referred to Eschmeyer’s Catalog of fishes
(1998) for complete references to species and generic names. An updated online version can be
accessed at <http//:www.calacademy.org>. Institutional abbreviations are taken from Fricke and
Eschmeyer (2009).
In the MATERIAL EXAMINED sections, the ASIZP specimens are grouped by general locality
(abbreviated as in following paragraph) and listed by institutional catalog number followed in
parentheses by number of specimens, place name or station number, and size range. Collection data
for research vessel stations and fishing ports are provided in Appendix 1 (Table 1). The official and
unofficial Taiwanese names of Taiwan fishing ports, in English and Chinese, and the geographic
coordinates for each locality can be found in Ebert et al. (2013: table 5, fig. 1).
For non-ASIZP specimens more complete data are provided in these sections. The reader
should note that characters used in the keys and diagnoses for genera are often not repeated in the
descriptions of the species of the particular genus. The DISTINGUISHING FEATURES in the species
descriptions are usually only those needed to distinguish Taiwan species from others of the genus
and may not apply if used with collections from other areas. Distributions are given in more detail
for Taiwan records than for other areas. There was no attempt to make the synonymies comprehen-
sive; they are limited to those most applicable to the Taiwanese fauna. Illustrations have been kept
to a minimum because a forthcoming book on the codfishes of Taiwan, geared for a general audi-
ence, will be fully illustrated with photographs or line drawing of all Taiwan species.
ABBREVIATIONS. For measurements and counts: TL, total length; PAL, pre-anal length; HL,
head length; 1D, first dorsal fin; 2D, second dorsal fin; P, pectoral fin; V, pelvic fin; A, anal fin;
GR-I, gill rakers on first arch, GR-II, gill rakers on second arch; pyl. caeca, pyloric caeca. Other:
descr., description; fm, fathom[s]; e., east, eastern; local., locality[ies]; n., north, northern; s., south,
southern; spec., specimen[s]; sta., station; w, west, western. Abbreviations for localities in Taiwan
follow Shao et al. (2008): ET, eastern Taiwan; NET, northeastern Taiwan; SET, southeastern Tai-
wan; SWT, southwestern Taiwan; SCS, South China Sea.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 35

RESULTS
The Diversity and Current Status of Grenadier Species in Taiwan
In this study, 71 species of grenadiers in 21 genera and 3 families are recognized. Despite a
relatively limited coastline (500 nautical miles), the species diversity in Taiwan is very high com-
pared to nearby areas. Of this total, 63 species (of 366, or ca. 17%) are in the family Macrouridae,
7 species (of 27, or ca. 30%) in the family Bathygadidae, and 1 species (of 2) in the family
Macrouroididae. The top three largest genera, all in Macrouridae, are Coelorinchus (23 species),
Ventrifossa (9 species) and Nezumia (5 species). Six species were described from Taiwan, and three
of them (Coelorinchus leptorhinus, C. sheni and C. fuscigulus) have not been reported elsewhere.
Shao et al. (2008) published a list of grenadier species from Taiwan with a discussion of their
distribution pattern.
After our detailed examination of specimens from Taiwan and based on new evidence, we
have made the following name changes to Shao et al.’s list: (1) specimens of Gadomus multifilis
are now recognized as Gadomus cf. multifilis; (2) records of Coelorinchus cylindricus in Taiwan
have been replaced by C. fuscigulus Iwamoto, Ho & Shao, 2009; (3) the specimen of Coelorinchus
spinifer is now recognized as C. cf. spinifer; (4) the specimen of Coryphaenoides asper is now
recognized as C. cf. asper; (5) Hymenocephalus gracilis has been reassigned to Hymenogadus and
Hymenocephalus kuronumai has been reassigned to Spicomacrurus; (6) the western Pacific popu-
lation of Malacocephalus laevis is now recognized as M. nipponensis; (7) specimens of Nezumia
coheni are now recognized as N. cf. coheni; (8) the specimen of Nezumia loricata is now reidenti-
fied as N. proxima; (9) a specimen of Pseudocetonurus septifer is now recognized as P. cf. septifer;
(10) the specimen of Paracetonurus cetonuropsis is now reidentified as Pseudonezumia pusilla;
and (11) Ventrifossa fusca is now recognized as a junior synonym of V. misakia.
We also report as first records for Taiwan Coelorinchus hexafasciatus,C. cf. macrorhynchus,
C. cf. notatus, Hymenocephalus papyraceus, and Ventrifossa sazonovi. Specimens of six species
are provided tentative identifications because they showed certain character differences compared
with published descriptions. Further study may result in recognizing them as new species or sub-
species. Moreover, several specimens of Bathygadus could not be confidently identified and were
therefore listed with no names: the genus is in great need of revision.
SYSTEMATICS
Key to the Grenadier Families in Taiwan
1a. One dorsal fin, no part elevated; head notably inflated; eyes tiny, horizontal orbit diameter more
than10timesin HL ............................................Macrouroididae
1b. Two dorsal fins, the first elevated; eyes larger, orbit diameter much less than 10 times in HL
........................................................................2
2a. Second dorsal fin high, much better developed than anal fin; gill rakers long and lathlike; outer
gillslitwhollyunrestricted.........................................Bathygadidae
2b. Second dorsal fin usually much less developed than anal fin; gill rakers short, tubercular; outer
gill slit variously restricted by folds of skin connecting gill arch above and below . . . . . . . .
...............................................................Macrouridae
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Family Bathygadidae
Key to the Genera and Species of Bathygadidae in Taiwan
1a. Chin barbel long, greater than half orbit diameter; elongated ray(s) in 1D, P and V; V 8 (rarely
9)..............................................................2(Gadomus)
1b. Chin barbel usually absent, if present, length much less than half orbit diameter; usually no
greatly elongated rays except in V, and if present, ray extremely fine distally and usually less
thanHL;V8–10(rarely 11) .......................................4(Bathygadus)
2a.GR-Iwithblunttips,4–6+20–23 ...............................................3
2b.GR-Iwithpointedtips,6+22–26 ....................................G. cf. multifilis
3a. Chin barbel about 3 times or less into HL; pyl. caeca 24–29. . . . . . . . . . . . . . . . . G. magnifilis
3b. Chin barbel about 2 times into HL; pyl. caeca 95–134. . . . . . . . . . . . . . . . . . . . . . . . G. colletti
4a. Gular membranes scaled; orbit width 1.1–1.2 times into interorbital . . . . . . . . . . . . . . . . . . . 5
4b. Gular membranes wholly naked; orbit width 1.2–2.1 times into interorbital . . . . . . . . . . . . . 6
5a. V 10 (rarely 11); barbel rarely present; pyl. caeca 12–19. . . . . . . . . . . . . . . . . . . B. nipponicus
5b. V 9 (rarely 10); rudimentary barbel usually present; pyl. caeca 34–54 . . . . . . . . . . . B. garretti
6a.Head bones weak, skin thin and fragile; interorbital width 31–36% HL; pyl. caeca 10–18 . . . .
.................................................................B. antrodes
6b.Head bones relatively strong, skin usually rather stout and thick; interorbital width 26–30% HL;
pyl.caeca20–22..................................................B. furvescens
Genus Bathygadus Günther, 1878
DISTINGUISHING FEATURES.— Chin barbel usually absent; when present barbel tiny and diffi-
cult to see without magnification. In some species, 1D, P, and V with a relatively elongated ray, but
never extremely long and thickened to degree found in Gadomus species.
REMARKS.— Identifying some members of this genus has been difficult and uncertain, partly
owing to the often poor condition of the specimens. Their delicate bones and integument are easi-
ly damaged in trawls, and crucial identification features are often distorted, damaged, or destroyed.
The many species described from the broad area encompassing Japan, Taiwan, the South China
Sea, Philippines, and Indonesia and our limited knowledge of distributional limits of the species
leave the possibility of encountering one or another species wide open. Our study material includ-
ed a number of specimens for which we could not arrive at a satisfactory identification; such spec-
imens are listed under Bathygadus sp. indet. and accompanied by brief remarks.
The genus was reviewed by Howes and Crimmen (1990), but many of their taxonomic con-
clusions are questionable (see Sazonov 1994:101; Iwamoto and Merrett 1997:479).
Bathygadus antrodes (Jordan and Gilbert, 1904)
Melanobranchus antrodes Jordan and Gilbert, 1904:606–607, pl. 4, fig. 7. (holotype: USNM 50932, 265 TL,
Sagami Bay, Japan, 501–749 fm [916–1,370 m], Albatross sta. 3696; other spec. from Sagami and Suruga
bays, 480–677 fm [878–1238 m]).
Bathygadus antrodes: Gilbert and Hubbs, 1916:149–150 (16 spec., 9 local. off Honshu I., Japan; 440–712 fm
[805–1,302 m]).— Okamura, 1970:30–33, pl. XI, text-fig. 16 (descr.; 15 spec.; Japan); Okamura,
1984:197, 356, fig. 138 (descr. in Japanese and English; 4 spec.; s. Japan, Okinawa Trough, 792–1,200
m).— Howes and Crimmen, 1990:191 (USNM spec.; s. Japan). Sazonov 1994:100–101, fig. 1 (5 spec.,
Nintoku Seamount [on Emperor Seamounts chain]; 1120–1160 m).— Chiou et al., 2004b:39. fig. 5 (2
spec., NET; one spec. we currently re-identify as B. garretti).— Shao et al., 2008: table 2 (19 spec., NET,
SWT, SET, SCS).
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 37

MATERIAL EXAMINED (7 spec.).— NET: ASIZP 61225 (1, 635 TL); Da-xi. SCS: ASIZP 65515
(1, 66.8 HL, 350 TL); CD 322, 1098 m; ASIZP 65633 (2, 150–375+ TL); CD 229, 880–1062 m.
SET: ASIZ 67033 (1, 78.1 HL, 367+ TL); CP 350, 1148 m; ASIZP 66100 (1, 62 HL, 310 TL); CD
322, 1098 m; ASIZP 66110 (1, 73.1 HL, 377+ TL); CD 322, 1098 m.
DISTINGUISHING FEATURES.— A species of Bathygadus with no scales on gular membrane;
chin barbel absent; dorsal profile behind head elevated, nape somewhat humpbacked; 1D II7–9; P
i14–i18; V 8–9; GR 5–6+18–20; pyl. caeca 10–18*; interorbital width 31–36% HL; orbit diameter
18–22%; upper jaw length 53–64%; filamentous rays of 1D, P, and V when intact slightly longer
than head.* Head bones and integument notably thin and fragile. Color overall dark; fins, head,
abdomen, mouth, and gill cavity usually black. (* indicates data from Gilbert and Hubbs, 1916).
DISTRIBUTION.— From n. part of Japan (Tohoku region of Honshu Is.) s. to Taiwan (NET,
SET, SCS) and on Emperor Seamounts Chain (at 40°05ʹN, 170°43ʹE), from about 800 to 1370 m.
REMARKS.— Chiou et al. (2004b:39, 42, fig. 5) recorded this species from Taiwan based on
two specimens, but we re-identified one of them (ASIZP 61226) as B. garretti. Bathygadus
antrodes is closely similar to B. bowersi (Gilbert, 1905) from Hawaii and to B. spongiceps Gilbert
and Hubbs, 1920 from the Philippines and Indonesia. Characters used to distinguish the three are
minimal and need to be further supported. A closer comparison with many more specimens from
throughout the western and central Pacific should prove fruitful.
Bathygadus furvescens Alcock, 1894
Figures 1A–B.
Bathygadus furvescens Alcock, 1894:128 (holotype ZSI F13047, “20.5 inches” [ca. 52 cm]; off Maldives,
Investigator sta. 150; 719 fm [1,315 m]).— Gilbert and Hubbs, 1920:388–391 (descr.; 5 spec., 5 loc.,
Philippine and Indonesia; 565–976 fm [1033–1785 m]).— Iwamoto and Merrett, 1997:479 (validated
position of species in Bathygadus).— Iwamoto and Graham, 2001:422–423, fig. 17 (descr., 10 spec., se.
coast Australia, “depths between 1,000 and 1, 240 m”).— Shao et al., 2008: table 2 (2 spec., SET, first
record for Taiwan).
MATERIAL EXAMINED (10 spec).— SET: ASIZP 65510 (1, 91.8 HL, 475+ TL); CP 127, 1263–
1268 m. Others questionably assigned to species, but with black head and abdomen, lacking paler
areas on head: SET: ASIZP 66938 (1, 475 TL); CP 127, 1263–1268 m. SWT: ASIZP 66115 (1,
100 HL, 450 TL); CD 322, 1098 m; ASIZP 70215 (1, 345 TL); ASIZP 63788 (4, 118+-380+ TL);
CD 192, 1305 m; ASIZP 65530 (2, 280–335 TL); CD 134, 736–1040 m.
DISTINGUISHING FEATURES.— A species of Bathygadus with no scales on gular membrane;
chin barbel absent; dorsal profile slightly elevated over nape; 1D II,8–10; P i15–i19; V 8–9; gill-
rakers outer arch (5–6) + (18–20); pyl. caeca 20–22. Interorbital width 26–30% HL; orbit diame-
ter 21–23%; suborbital width 14–15%; postorbital length 51–55%; distance orbit to preopercle
48–50%; length upper jaw 56–59%; length P 61–81%; length V 67–83%; length longest gill-raker
13–15%. Fins well developed; outer V ray elongated and distally filamentous. Flesh and head
bones relatively firm. Fins black to dark dusky. (After Iwamoto and Graham [2001:479] except for
lengths of postorbital and longest gill-raker, the ranges of these were extended by our specimens.)
DISTRIBUTION.— Recorded from Maldives, Indonesia, Philippines, se. Australia, and now off
Taiwan, in depths between approximately 1000 m and 1800 m.
REMARKS.— Howes and Crimmen (1990:195) treated the species as a member of Gadomus
based on a specimen they erroneously considered a syntype, but which not only was non-type
material but also a species different from the holotype (see Iwamoto and Merrett 1997:479). The
large Taiwan specimen was similar in most mensurable and count characters to the specimens we
called B. antrodes. However, its head bones were stouter, integument notably tougher, with scale
38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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pockets well developed, color overall paler, the interorbital width slightly narrower (27–28% HL),
and none of the fin rays longer than the head. The specimen agreed in these respects with
B. furvescens as circumscribed by Iwamoto and Graham (2001:422–423) and represents the first
record of the species from Taiwan waters.
Bathygadus garretti Gilbert and Hubbs, 1916
Bathygadus garretti Gilbert and Hubbs, 1916:151–153, pl.8, fig. 1 (holotype USNM 76863, 513 TL, Suruga
Gulf, Japan, Albatross sta. 5059, 197–297 fm [360–543 m]).
Bathygadus (Melanobranchus)garretti: Gilbert and Hubbs, 1920:380 (in key).
Bathygadus garretti: Okamura, 1970:34–36, pl. 12, text-fig. 17 (descr. based on paratype, USNM 135351);
Okamura, 1984:197, 357, fig. 139 (descr. in Japanese and English; 7 spec., Okinawa Trough, 360–650
m).— Chiou et al., 2004b:42, fig. 6 (1 spec., Da-xi, NET).— Shao et al., 2008: table 2 (6 spec., NET, SWT,
SCS).
Bathygadus nipponicus: Howes and Crimmen, 1990:191 (in part; synonymized B. garretti with B.
nipponicus). Chiou et al., 2004b:42, fig. 7 (1 spec., NET).
MATERIAL EXAMINED (20 spec.).— NET: ASIZP 61226 (1, 86.5 HL, 446 TL); Da-xi; ASIZP
61227 (2, 70–77 HL, 362–380 TL); Da-xi; ASIZP 61228 (1, 96.8 HL, 458 TL); Da-xi, ASIZP
64274 (1, 91.3 HL, 365 TL); CP 235, 764 m; ASIZP 70241 (1, 281 TL); Da-xi; ASIZP 70244 (1,
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 39
FIGURE 1. Bathygadus furvescens Alcock, 1894. A. ASIZP 65510, 475+ mm TL, preserved. B. ASIZP 63788, 1 of 4,
380+ mm TL, preserved.
A
B

153 TL); Da-xi; ASIZP 70246 (3, 127–175 TL); Da-xi; ASIZP 70249 (3, 360–456 TL); Da-xi;
ASIZP 70253 (1, 444 TL); Da-xi. SWT: ASIZP 65512 (1, 58.1 HL, 320 TL); CP 130, 709–728 m;
ASIZP 65593 (1, 70 HL, 338 TL); CD 140, 280–452 m; ASIZP 65599 (1, 65.6 HL, 291+ TL); CD
137, 316–477 m. No data: ASIZP 65580 (1, 71.6 HL, 335+ TL). Other material:Japan: BSKU
98225 (1, 62.5 HL, 305+ TL) and BSKU 98224 (1, 58.9 HL, 320 TL); Suruga Bay, 520–545 m; 23
Nov. 1978.
DISTINGUISHING FEATURES.— A species of Bathygadus with gular membrane scaled; a rudi-
mentary chin barbel usually present (sometimes absent); dorsal profile behind head relatively low,
nape not humpbacked; 1D II8–10; P i15–i18; V 10 (rarely 11); GR-I (5–6) + (17–20); pyl. caeca
34–46 (or more); interorbital width 27–32% HL; orbit diameter 21–26%; upper jaw length
52–58%; none of fins with greatly elongated rays.
DISTRIBUTION.— Southern Japan s. to Taiwan (NET, SWT) in 360–650 m.
REMARKS.— Our identification of specimens of this species was based on a combination of
characters, including the presence of a rudimentary chin barbel. In a few specimens, the barbel was
not present, but other characters that we deemed important appeared to confirm our identification.
Howes and Crimmen (1990:1910) synonymized B. garretti with B. nipponicus, stating that the only
difference between the two was the presence of a rudimentary chin barbel in the former, a sugges-
tion made earlier by Okamura (1984:358). However, our ASIZ specimens suggest that B. garretti
almost always has V 10 (11 on one side in a paratype) vs. mostly 9 in B. nipponicus (the holotype
has 10, fide Okamura 1970 and Howes and Crimmen 1990, not 9 as stated in the original descrip-
tion), a slightly larger orbit (1–26% HL vs. 17–22%), and more pyloric caeca (34–46
[5 spec.] vs. 12–19 [5 spec.]). The count of pyloric caeca in the single paratype of B. garretti was
given as 50 by Gilbert and Hubbs (1916:152), 49 by Okamura (1970:34), and 54 by Howes and
Crimmen (1990:191).
Specimen ASIZP 61227, listed in Chiou et al. (2004b) as the first Taiwan record of B. nippon-
icus, was determined by us to be B. garretti. One of two specimens of B. antrodes (ASIZP 61226)
recorded in the same paper is now re-identified as this species.
Bathygadus nipponicus (Jordan and Gilbert, 1904)
Regania nipponica Jordan and Gilbert, 1904:605–606, fig. (holotype, USNM 50931, 590+ TL; Albatross sta.
3721, Suruga Bay, Japan, 207–250 fm [379–457 m]).
Bathygadus nipponicus: Gilbert and Hubbs, 1916:142 (listed).— Okamura, 1970:33 (compiled).— Howes
and Crimmen, 1990:191–192 (in part; holotype data; see Remarks under B. garretti).— Chiou et al.,
2004b:42, fig. 7 (1 spec., NET, re-identified as B. garretti).— Shao et al., 2008: table 2 4 spec., NEW,
SWT, SCS).
Bathygadus (Melanobranchus)nipponicus: Gilbert and Hubbs, 1920:380 (in key).
MATERIAL EXAMINED (6 spec.).— SWT: ASIZP 64117 (1, 60 HL, 210+ TL); CD 193, 821 m;
ASZIP 65633 (3, 44–60.2 HL, 150–375 TL) and ASZIP 65634 (1, 66 HL, 264+ TL); CD 229,
880–1062 m. SCS: ASIZP 66126 (1, 83.0 HL, 480 TL); CD 320, 731 m.
DISTINGUISHING FEATURES.— A species of Bathygadus with gular membrane scaled; chin bar-
bel absent; dorsal profile behind head relatively low, nape not humpbacked; 1D II,8–10, P i15–i17;
V 9 (rarely 10); GR 6+(17–22) (ASIZP 65634 had 2 rudiments and 6 developed rakers on upper
arm); pyl. caeca 12–19; interorbital width 27–32% HL; orbit diameter 22–26%; upper jaw length
53–56%; none of fins with greatly elongated finray.
DISTRIBUTION.— Southern Japan to Taiwan (SWT, SCS) in 731–l062 m.
REMARKS.— Jordan and Gilbert (1904:605) gave the gill raker count as 5+16 in the holotype
and only specimen in their original description of the species; this count was confirmed by Howes
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and Crimmen (1990:191). That the count is lower than the values in our specimens is somewhat
unsettling, but we hold to our identification until the time more specimens become available and
the holotype can be re-examined by one of us. So far as we can determine, our specimens repre-
sent only the second record of the species and the first from Taiwan. Chiou et al. (2004b:42,
fig. 7) listed the species as a first record from Taiwan based on ASIZP 61227, but we re-examined
that specimen and determined it to be B. garretti.
Bathygadus sp. indet.
ASIZP 64268 (1, 39.2 HL, 180+ TL), CD 191, 821 m. This specimen from 821 m in the SCS had
interorbital 33% HL, orbit 20%, suborbital 14%, postorbital 56%, orbit-preopercle 49%, upper
jaw 62%, and V 10, P 17–18, GR-I 5+19, characters that suggest B. spongiceps.
ASIZP 65634 (1, 264+ TL), CD 229, 880–1062 m. This specimen from SWT is in poor condition
and most characters are undecipherable. Interorbital 23 mm, orbit 12.7 mm, orbit-preopercle 28
mm; V 9, GR-I 6+19. Head integument relatively tough; orbit too wide to be B. garretti or
B.nipponicus.
ASIZP 66738 (1, 21.3 HL, 92+ TL), CD 324, 1293 m. This small specimen from 1293 m in the
SCS had a broad interorbital (38% HL), small orbit (22%), deep suborbital (18%), and GR-I
5+18, characters that suggest B. antrodes.
ASIZP 66793 (1, 405 TL); CD 320, 731 m. This specimen from the SCS is in poor condition; it
was originally identified as Bathygadus entomelas.
Genus Gadomus Regan, 1903
DISTINGUISHING FEATURES.— Chin barbel present, usually thick and long. V rays usually 8,
rarely 9. Second spinous ray of 1D, upper ray of P, and outermost ray of V usually elongated, in
some species extremely long; outer V ray usually rather thick.
REMARKS.— The genus was revised by Howes and Crimmen (1990), who based their work
mostly on the literature and old, previously recorded specimens (none recently collected). With
respect to Gadomus aoteanus, they followed McCann and McKnight (1980), who gave an erro-
neous count of V 9, but examination of the holotype and many other specimens (by TI) of that
species showed that there are consistently eight rays in each fin. Gilbert (1905:658) gave a count
of V 9 for the holotype (and only type specimen) of G. melanopterus; this is the only record of nine
rays in a specimen of Gadomus that we are aware of. Asecond non-type specimen (CAS-SU 8545)
that he reported in the original description has V 8.
Seven of the 12 described species of Gadomus are found in the western Pacific, but there may
be others that are new to science. The species from the Indo-Pacific region have, for the most part,
been inadequately circumscribed owing to the lack of large series from many localities. A thorough
review of the genus using more recently collected material and genetic analyses is badly needed.
Gadomus colletti Jordan and Gilbert, 1904
Gadomus colletti Jordan and Gilbert, 1904:603–604 (holotype USNM 50930, Albatross sta. 3721, Suruga
Bay, Japan; 207–250 fm [379–457 m]).— Gilbert and Hubbs, 1916:154–155 (descr.; 4 spec., 68–302 TL;
Suruga Gulf, 211–293 fm [386–536 m]); Gilbert and Hubbs, 1920:392 (in key).— Okamura, 1970:23–26,
pl. I, fig. a; text-fig. 12A, 13 (descr., 44 spec., 177–322 TL); s. Japan, 360–547 m).— Howes and Crim-
men, 1990:199 (descr. based on holotype and USNM spec.).— Chiou et al., 2004b:43, fig. 8 (2 spec.,
NET).— Shao et al., 2008: table 2 (5 spec., NET, SCS).
Bathygadus colletti: Weber, 1913:172 (listed).
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 41

MATERIAL EXAMINED (6 spec.).— NET: ASIZP 61223 (1, 79 HL, 363 TL); Da-xi; ASIZP
61224 (1, 82.7 HL, 382 TL); Da-xi; ASIZP 65636 (1, 68.1 HL, 352 TL); CD 214, 488–1027 m;
Da-xi. ASIZP 70251 (1, 90 HL, 346 TL); Da-xi. SWT: ASIZP 65513 (1, 200+ TL); CP 130,
709–728 m. Other specimens: Japan: HUMZ 37408 (1, 29.8 HL, 146+ TL).
DISTINGUISHING FEATURES.— A species of Gadomus with chin barbel about 2/3 of HL or
slightly longer; elongated ray in 1D, P and V often longer than HL; 1D II,10–11; P (i16) i18–i21;
V 8; outer gill rakers lathlike but relatively short, bluntly tipped, (4–5)+(8–21) (ASIZP 61224 had
1 rudiment and 5 developed rakers on upper arm, 16 developed and 3 rudiments on lower arm);
pyl. caeca small, numerous, about 95 to 165. Interorbital width 14–18% HL; orbit diameter
22–23%; upper jaw length 51–58%; chin barbel 57–91%; 1D spinous ray less than twice HL. Color
relatively pale, mouth and gill cavity dark but paler on outer margins; lips and barbel whitish; fins
dusky to blackish, dorsal fins lighter basally.
DISTRIBUTION.— Southern Japan to Taiwan (NET, SWT) in 488–1027 m.
REMARKS.— This species was first recorded from Taiwan by Chiou et al. (2004b:43). Our four
specimens appeared to have a somewhat shorter barbel (57–91% of HL) than reported by others.
Okamura (1970:24) recorded their length as 0.9–1.2 into HL in 44 specimens from Japan, “about
as long as, or a little shorter than, length of head” in 10 specimens from the Kyushu-Palau Ridge
and Tosa Bay (Okamura 1982:345), and 75–91% HL in five specimens from the Okinawa Trough
(Okamura 1984:356).
Gadomus magnifilis Gilbert and Hubbs, 1920
Figure 2.
Gadomus magnifilis Gilbert and Hubbs, 1920:398–401, fig. 4 (holotype USNM 78208; n. Mindanao; Alba-
tross sta. 5515 in about 700 fm [1280 m]; 2 paratypes, Philippines; 508–554 fm [929–1014 m]).— Howes
and Crimmen, 1990:197 (descr. from type spec.).— Shao et al., 2008: table 2 (1 spec., SWT, first record
for Taiwan).
MATERIAL EXAMINED.— SWT: ASIZP 65627 (1, 70 HL, 385+ TL); CD 229, 880–1062 m.
DISTINGUISHING FEATURES.— A species of Gadomus with chin barbel about 29–72% of HL;
length elongated P ray 2–3 times HL; prolonged ray in 1D and V greater than 1.5 times HL; 1D
II,9–10; P i17–i18; V 8; outer gill rakers bluntly tipped, lathlike, short, about half orbit diameter,
(5–6)+(21–23); pyl. caeca 24–29; interorbital width 16–17% HL; orbit diameter 21–24%; subor-
bital depth 10–15%; upper jaw length 55–60%; chin barbel 29–72% (compiled from Gilbert and
Hubbs 1920 and ASIZP 65627).
42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 2. Gadomus magnifilis Gilbert and Hubbs, 1920. ASIZP 65627, 385+ mm TL, preserved.

DISTRIBUTION.— Taiwan (SWT) to n. Mindanao, Philippines, in 929–1280 m.
REMARKS.— Our identification of the single Taiwan specimen is tentative. Its barbel length is
somewhat shorter (29% HL cf. 61–72%) and suborbital somewhat deeper (15% HL cf. 10%) than
in the type specimens of G. magnifilis. The Taiwan specimen appears similar to G. introniger in its
gill-raker count, barbel length, orbit diameter, and bluntly tipped gill rakers, but differs in having
a slightly narrower interorbital (17% cf. 20–23%), slightly deeper suborbital (15% cf. 10–13%),
and longer orbit-to-preopercle distance (51% cf. 44–49%). The pyloric caeca could not be count-
ed in our specimen because they had been previously extracted and were unavailable; they may
have offered additional clues to the correct identity. Gilbert and Hubbs (1920:396) considered
G. denticulatus as closely related to G. magnifilis, but compared to our specimen, G. denticulatus
has a much narrower interorbital space (1.5–1.8 into orbit diameter cf. 1.3), slightly larger orbit
(23–24% HL cf. 21% in our spec.), and slightly fewer gill rakers (4–6)+(17–22) (23–27 total), cf.
6+23 (29 total).
Gadomus cf. multifilis [sensu (Günther, 1887)]
Figures 3A–B.
Gadomus multifilis (Günther, 1887): Shao et al., 2008: table 2 (4 spec., SWT, first record for Taiwan).
MATERIAL EXAMINED (9 spec.).— SWT: ASIZP 64092 (4, 30–50.6 HL, 130+-260+ TL); CD
193, 821 m. SCS: ASIZP 66334 (1, 38 HL, 240 TL) and ASIZP 66240 (1, 37.1 HL, 223+ TL), CD
307, 1591 m; ASIZP 66189 (1, 35.1 HL, 170+ TL) and ASIZP 66810 (1, 35.5 HL, 185+ TL); CD
322, 1098 m. Other specimens: ASIZP 68056 (1, 34.7 HL, 223 TL); CC2702 Aurora, 944–1004
m, 27 May 2007.
DISTINGUISHING FEATURES.— A species of Gadomus with chin barbel about 50–75% of HL;
length elongated P ray 2–3 times HL, prolonged ray in 1D and V much greater than HL; 1D II,8–9,
P i16–i19; V 8; outer gill rakers sharply tipped, lathlike (6–7) + (23–26); pyl. caeca 24–29. Interor-
bital width 16–17% HL, orbit diameter 21–24%; suborbital depth 10–15%; upper jaw length
55–60%; chin barbel 29–72% (compiled from Gilbert and Hubbs, 1920).
DISTRIBUTION.— Taiwan (SWT, SCS) and broadly across Indo-West Pacific, if these speci-
mens are the same as G. multifilis.
REMARKS.— Our identification of nine Taiwan specimens has been problematical, although
they for the most part fit the original description and subsequent descriptions of G. multifilis by
Gilbert and Hubbs (1920) and Howes and Crimmen (1990). However, four of the ASIZ specimens
(ASIZP 64092) have a slightly lower count of GR-I (6 + 23) and more pyloric caeca (30–47 in 4
spec.); ASIZP 66334 had more rakers, 7 + 26 rakers. Gilbert and Hubbs (1920:406–408) gave the
gill-raker count as 6 + (26–27) and pyloric caeca as 12 and 16. Howes and Crimmen
(1990:195–197) counted 6 + 25 gill-rakers for the holotype and eight other specimens, and counts
of 25 and 15 pyloric caeca in two of those specimens. The low gill-raker counts and high pyloric
caeca counts of ASIZP 64092 agree rather well with those Gilbert and Hubbs (1920:403) gave for
G. introniger (gill-rakers (5–6) + (20–24) and “pyloric caeca [in several specimens, 35 to 52”]).
However, the orbit diameter and interorbital width are less in our specimens and the gill-rakers are
pointed, not bluntly tipped, as in G. introniger. Similarly, the counts of gill-rakers and pyloric
caeca, plus the interorbital and orbit dimensions, agree closely with G. magnifilis, but the gill-rak-
ers are also bluntly tipped in that species. We see no resolution to our dilemma without a thorough
revision of the genus using extensive material from throughout the Indo-West Pacific, and espe-
cially from that critical region around the Philippine Islands and the Malay Archipelago.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 43

Family Macrouridae
Key to Genera and Some Species of Macrouridae in Taiwan
(number of species in Taiwan in parentheses after genus)
1a.Sixbranchiostegalrays.......................................................2
1b.Sevenbranchiostegalrays.....................................................4
2a. Spinous 1D ray smooth; a stout continuous suborbital ridge terminating posteriorly in a sharp
point;V7................................................Coelorinchus (23 spp.)
2b. Spinous 1D ray serrated (sometimes weakly) along leading edge; suborbital ridge not continu-
ousandnotendinginasharppoint;V7–14......................................3
3a. Gill-rakers absent on lateral side of first gill arch; V7 or 8; anus in middle third of distance
betweenVandA......................................Mataeocephalus hyostomus
3b. Gill-rakers present on lateral side of first gill arch; V 7–14; anus usually immediately anterior
toA...................................................Coryphaenoides (4 spp.)
4a. Abdomen and isthmus with patches of fine black striations with silvery underlayment (especial-
ly when fresh); small lens-like light organ on chest connected by black medioventral line to a
44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 3. Gadomus cf. multifilis [sensu (Ganther, 1887)] A. ASIZP 64092, 260+ mm TL, fresh. B. ASIZP 66334, 240
mm TL, preserved.
A
B

secondlensimmediatelybeforeanus ...........................................5
4b. Abdomen and isthmus lacking areas of fine black striations; light organ, if present, not as above
.........................................................................7
5a. Nasal bones forming three flat platelike horizontal processes; gular membrane with net- or
mesh-like pattern; prolonged distal portion of outer pelvic fin ray with narrow membranous
flange................................................Spicomacrurus kuronumai
5b. Nasal bones not forming horizontal platelike processes; gular membrane with thin black trans-
verse lines at right angle to median line; outer pelvic fin ray tapers evenly to distal tip . . . . 6
6a. First dorsal fin with weakly denticulate spinous ray; gill rakers on inner side of first arch with
12–16 rakers on lower limb; V 7–9, usually 8. . . . . . . . . . . . . . . . . . . . Hymenogadus gracilis
6b. First dorsal fin spinous ray entirely smooth along leading edge; gill rakers on inner side of first
arch with 15–22 rakers on lower limb; V 7–12. . . . . . . . . . . . . . . . . Hymenocephalus (4 spp.)
7a. Anus and urogenital pore within broad black naked area, the periproct, the posterior border of
which abuts origin of A and spans most of space between V and A. . . . . . . . . . . . . . . . . . . . 8
7b. Periproct smaller, not occupying most of space between V and A; anus (and urogenital pore)
either abuts A origin, or separated from A by several scales rows, sometimes closer to inser-
tionofVthantooriginofA .................................................10
8a. Origin of V behind P origin; 2nd spinous ray of 1D smooth . . . . . . . . . . Trachonurus (2 spp.)
8b. Origin of V below or anterior to P origin; spinous 1D ray serrated along leading edge . . . . . 9
9a. Head greatly inflated, broad and deep; snout fully scaled; a series of enlarged scales along ante-
riorsectionof2D ...........................................Cetonurus globiceps
9b. Head not inflated, moderately compressed; underside of snout naked; no series of enlarged
scalesalong2D .........................................Sphagemacrurus (2 spp.)
10a. Olfactory organ massive, its diameter almost equal to that of orbit . . . . Macrosmia phalacra
10b. Olfactory organ small to moderate, much less than orbit diameter . . . . . . . . . . . . . . . . . . . 11
11a. Most of dorsal surface and entire ventral surface of snout naked . . . . . . . . . . Kumba (3 spp.)
11b. Dorsal surface of snout fully scaled; ventral surface of snout variously naked . . . . . . . . . . 12
12a. Low jaws bearing enlarged, widely space, fang-like teeth in 1 or 2 rows; spinous 1D ray
smooth.............................................Malacocephalus nipponensis
12b. Lower jaw teeth normal in size, closely spaced, none fang-like, in 2 or more rows to broad
band; spinous 1D ray smooth or serrated along leading edge. . . . . . . . . . . . . . . . . . . . . . . . 13
13a. V with 6 rays, placed anterior to P base; anus closer to A origin than to V insertion . . . . . . .
........................................................Pseudonezumia pusilla
13b. V with 7 or more rays, position variable from below 1D to under preopercle; anus removed
from A origin, usually closer to V insertion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a. Head broadly inflated, interorbital space about 40% of HL; orbit diameter less than 25% HL;
suborbitaldeep,about20%HL ..........................Pseudocetonurus cf. septifer
14b. Head not especially inflated, interorbital space less than 35% HL; orbit diameter usually more
than25%HL,suborbital lessthan 20%HL .....................................15
15a. Terminal and lateral snout scutes present, the terminal scute paired; suborbital ridge marked
bytworowsofcoarselymodifiedscales........................................16
15b. Terminal and lateral snout scutes generally not well developed, the terminal scute single, if
present, not large; no coarsely modified scales along suborbital ridge . . . . . . . . . . . . . . . . . 17
16a. Outer series of rakers on GR-I usually absent (sometimes with a few rudimentary spicules);
V7or8................................................Mataeocephalus (2 spp.)
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 45

16b. Outer series of rakers on GR-I always present, although sometimes rudimentary; V 7 to 17.
.............................................................Nezumia (5 spp.)
17a. Upper jaw extending to below posterior one-third or more of orbit; premaxillary tooth band
extends posteriorly beyond maxillary process; inner series of gill rakers on GR-I 14–20 total;
no scales on gular or branchiostegal membranes . . . . . . . . . . . . . . . . . . . . Ventrifossa (9 spp.)
17b. Upper jaw extending posteriorly to below anterior half of orbit; premaxillary teeth band does
not extend beyond maxillary process; gill rakers on inner series of first arch usually less than
14 total; some species with scales on gular or branchiostegal membranes. . . . . . . . . . . . . . 18
18a. Color brown to blackish, no silvery pigmentation; fins uniformly blackish; teeth in broad
bands.......................................................Kuronezumia dara
18b. Color pale to greyish; silvery pigmentation ventrally in fresh specimens; fins often with black
blotches or streaks; teeth in relatively narrow bands. . . . . . . . . . . Lucigadus nigromarginatus
Genus Cetonurus Güünther, 1887
DISTINGUISHING FEATURES.— A distinctive genus owing to its large, soft, globose head, fully
scaled including branchiostegal rays (and gular membrane in some specimens); enlarged scales
along dorsal-fin interspace and anterior portion of 2D; interrupted lateral line; BR 7; large periproct
region spanning most of the short space between V and A; and serrated spinous 1D ray.
REMARKS.— Two species recognized, one in our area; genus reviewed by Sazonov and
Shcherbachev (1985).
Cetonurus globiceps Vaillant, 1884
Figures 4A–B.
Macrurus globiceps Vaillant in Filhol 1884:183, fig. 2 (name and figure, Spanish Sahara; lectotype: MNHN
1886–0092; 7 paralectotypes).
Hymenocephalus crassiceps (non Günther, 1878): Vaillant, 1888:214–218, pl. 20, figs. 1, 1a-e (descr.; Atlantic
off France, North Africa, and Azores)
Cetonurus robustus Gilbert and Hubbs, 1916:207–210, pl. 11, fig. 2 (holotype USNM 76870, Albatross sta.
4971, off central Hondo [Honshu], Japan, 33°23ʹ30ʺN, 135°34ʹ00ʺE; 649 fm [1187 m]; 4 paratypes, sta.
4973, 600 fm [1097 m])
Cetonurus globiceps: Sazonov and Shcherbachev, 1985 (descr., distr.). Iwamoto and Williams, 1999:169 (7
spec., w. and se. Australia, 792–1030 m).— Iwamoto and Graham, 2001:458 (17 spec., 940–1200 m).—
Shao et al., 2008: table 2 (3 spec., SET, SCS, 998–1290 m; first Taiwan record)
MATERIAL EXAMINED (5 spec.).— SCS: ASIZP 65559 (1, 170+ TL); CD 136, 998–1211 m;
ASIZP 66095 (1, 270+ TL); CD 324, 1293 m. SWT: ASIZP 65620 (1, 331 TL); CD 228,
1262–1290 m. SET: ASIZP 67020 (1, 310+ TL, 61 HL); CP 353, 1205 m. Other material: ASIZP
68059 (1, 390 TL); Aurora, the Philippines.
DISTINGUISHING FEATURES.— A species of Cetonurus with large orbit (24–28% HL); small
scales (18–19 rows below 1D and 14–16 rows below 2D); and 3–4 rows of closely spaced teeth on
premaxilla. Size to 510 mm TL.
DISTRIBUTION.— This species is widespread in tropical and subtropical waters of the Indo-
West Pacific and Atlantic but is absent in the central and e. Pacific. In the w. Pacific it is found off
Japan, Taiwan, New Zealand, and Australia. Four specimens were collected in the South China Sea
and the s. coast of Taiwan at depths between 998 and 1293 m.
REMARKS.— The only congener, C. crassiceps (Günther, 1878), is a closely similar species
that has been recorded from the Kermadecs, Hawaiian Islands, Vanuatu, Loyalty Ridge, Norfolk
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Ridge, Lord Howe Rise, and in the central and South Atlantic. Characters used by Sazonov and
Shcherbachev (1985) to distinguish the only two members of the genus are weak and need to be
further tested. Three of the specimens here listed were the basis for Shao et al.’s (2008:fig. 2) first
record of the species from Taiwan.
Genus Coelorinchus Giorna, 1810
DISTINGUISHING FEATURES.— BR 6; suborbital ridge formed of stout, coarsely spined, modi-
fied scales extending continuously from tip of snout to posterior angle of preopercle, ending in a
sharp point; second spinous ray of first dorsal fin smooth along leading edge (some juveniles and
exceptional adults may have a few spinules near distal tip); ventral light organ variously developed,
from small gland anterior to anus not externally visible, to large prominent organ with one or two
dermal fossae along ventral midline; no rakers along outer (lateral) side of first gill arch. V almost
always 7, rarely 6.
REMARKS.— The most diverse genus of family Macrouridae with more than 118 recognized
species, categorized by some into as many as seven subgenera; 23 species known from Taiwan.
Relationships closest to Macrourus, the only significant differences between the two genera the
presence of denticulations on spinous second ray of 1D and generally more pelvic fin rays (8 or 9,
rarely 7) in Macrourus. Preliminary cladistic analysis (e.g., Roa-Varón and Ortí 2009:700) suggests
that Macrourus is deeply nested within Coelorinchus, a situation that could prove to be a nomen-
clatural nightmare.Greatest diversity found in tropical regions of the Indo-West Pacific. Species
most common in upper- to middle-slope waters between 200 and 1000 m, although some species
range into shallow continental-shelf depths and others into depths greater than 1000 m. Many
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 47
FIGURE 4. Cetonurus globiceps Valliant, 1884. A. AISZP 67020, 310+ mm TL, fresh. B. ASIZP 65559, 170+ mm TL,
preserved.
A
B

species of shallower waters are highly restricted in their geographic distribution. Maximum size
attained range from about 20 cm TL to more than 87 cm.
In statements concerning the ventral light organ, Groups I through IV refer to categories
assigned by Iwamoto (1990) based on the external development of the dermal window of the light
organ.
Key to the Species of Coelorinchus in Taiwan
1a. Light organ with two widely separated, usually blackish fossae, one immediately behind isth-
mus, the second immediately before anus, both connected by a medioventral line; anus/uro-
genital opening immediately before A origin; ventral snout surface with a series of overlapping
scales at anterolateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. Light organ variously developed, from scarcely visible to prominent with a single large fossa
between or slightly anterior to V bases; anus/urogenital opening immediately before or sepa-
rated by several scale rows from A origin; no overlapping series of scales ventrally on antero-
lateral snout margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2a. Underside of head behind mouth mostly scaled . . . . . . . . . . . . . . . . . . . . . . . . . C. formosanus
2b. Underside of head essentially wholly naked (tiny isolated scales sometimes present above pos-
terior angle of mouth and on preopercle) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Anterior rays of 2D about equal in height to those of A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3b. Anterior rays of 2D decidedly shorter than those of A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
4a. Body with prominent dark longitudinal stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hubbsi
4b. Body with prominent saddles or completely lacking body markings . . . . . . . . . . . . . . . . . . . 5
5a. No body markings; snout rather short, broad, depressed, 39–44% HL. . . . . . . . C. brevirostris
5b. Prominent saddle marks on body; snout 36–50% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. 1D with elongated spinous ray; BR membrane dusky to pale. . . . . . . . . . . . . . . . C. cingulatus
6b. No elongated spinous ray in 1D; BR membrane dark to black . . . . . . . . . . . . . . . C. fuscigulus
7a. Body covered with bold vermiculations and blotches, more or less aligned in 2 or 3 longitudi-
nal rows; origin of 2D slightly before vertical through A origin; 1D-2D interspace about equal
to length base of 1D . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. multispinulosus
7b. Body immaculate or with irregular, sometimes faint, markings; origin of 2D about on same ver-
tical as, or somewhat behind, A origin; 1D-2D interspace greater than length base of 1D . . 8
8a. Gular and branchiostegal membranes heavily peppered with black pigmentation, chest region
relatively dark, underside of head covered with tiny black sensory papillae; spinules on body
scales broadly triangular, in irregular, somewhat quincunx pattern . . . . . . . . . . C. kamoharai
8b. Underside of head faintly dusky to immaculate, chest region light dusky, sensory papillae on
underside of head scattered and inconspicuous; spinules on body scales weak, small, in 6–14
somewhat parallel to divergent rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. longissimus
9a. Snout relatively blunt, not acutely pointed; a distinct curve in anterior portion of suborbital
shelf; anus about midway between A origin and base of outer V rays, preceded by large black
fossa of light organ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. macrochir
9b. Snout acutely pointed in lateral view; suborbital shelf lacking distinct curve anteriorly; anus
closer to A origin than to base of outer V rays, fossa of light organ present or absent . . . . 10
10a. A blackish round blotch above P; anus removed from A by several scale rows; large fossa of
light organ extending forward between V bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
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10b. No blackish blotch above P; anus immediately before or removed from A origin; light organ
situated posterior to V bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11a. Black blotch above P large, spanning 5–7 diagonal rows of scales; gular and branchiostegal
membranes black; height 1D 45–54% of pre-vent length . . . . . . . . . . . . . . . . . C. kishinouyei
11b. Black blotch above P small, spanning 3 or 4 diagonal scale rows; gular and branchiostegal
membranes pale or whitish; height 1D 67% of pre-vent length. . . . . . . . . . . . . . C. cf. notatus
12a. Underside of head entirely or almost entirely naked (underside of head completely naked
except for ventral portion of preopercle in C. hexafasciatus) . . . . . . . . . . . . . . . . . . . . . . . 13
12b. Underside of head scaled (underside of snout naked in C. leptorhinus, but posteriorly head
scaly) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
13a. Dark saddles on trunk and tail . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hexafasciatus
13b. No saddle marks on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a. Spinules on body scales below 2D with 3–5 divergent rows of strong, triangular spinules, mid-
dle row higher than lateral rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. productus
14b. Spinules on body scales below 2D with 4–11 somewhat parallel to divergent, sharply crest-
like rows of narrow, triangular spinules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15a. Scales of median nasal ridge 9–12, spinule rows directed laterally and posteriorly only . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. anatirostris
15b. Scales of median nasal ridge 6–8, with spinules radiating in all directions . . . . C. asteroides
16a. Prominent saddle markings on body; snout tip lacking sharp terminal scute . . . . . . . C. sheni
16b. No saddle markings on body; a sharp terminal snout scute present. . . . . . . . . . . . . . . . . . . 17
17a. Underside of snout naked (head scaly from above mouth posteriorly) . . . . . . . C. leptorhinus
17b. Underside of snout and head fully scaled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a. Anterolateral margin of snout incompletely supported by bone. . . . . . . . . . . . . . . . . . . . . . 19
18b. Anterolateral margin of snout completely supported by bone . . . . . . . . . . . . . . . . . . . . . . . 20
19a. Body scales with 4–6 parallel rows of strong, high, usually broad-based (i.e., large lateral but-
tresses) spinules; scales between occipital ridges and on underside of head mostly with one
spinule row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. parallelus
19b. Body scales with 3–8 somewhat parallel to broadly divergent rows of spinules; scales between
occipital ridges with 1–5 spinule rows, on underside of head with 1–3 rows . . . C. divergens
20a. Light organ visible only as crescent-shaped area of perianal ring, no separate fossa; most
scales between occipital ridges with a single row of spinules . . . . . . . . . . . . . . . C. japonicus
20b. Light organ with short fossa extending anteriorly almost to or between V fins; scales between
occipital ridges with 2–5 rows of spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21a. Nasal fossa naked; large strong spinules on body scales in 3 divergent rows; scales between
occipital ridges with a single row of spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cf. spinifer
21b. Nasal fossa usually scaled anteroventrally, seldom naked; spinules on body scales in 4–6 (3–7)
divergent rows; scales between occipital ridges with multiple rows (usually 2–4) of spinules
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22a. Length 1D base 1.2 times into 1D-2D interspace; nasal fossa densely scaled anteroventrally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cf. macrorhynchus
22b. Length 1D base 1.3–2.0 times into 1D-2D interspace; nasal fossa rather sparsely scaled on
anteroventral surfaces, sometimes naked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. smithi
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 49

Coelorinchus anatirostris Jordan and Gilbert, 1904
Coelorhynchus anatirostris Jordan and Gilbert, 1904:619 (holotype: USNM 51471 [ex CAS-SU 8550], 40 cm
long; Misaki, Japan).— Okamura, 1970:186–189, pl. VIII, text-fig. 80 (72 spec.; s. Japan, 300–540 m).—
Okamura in Okamura et al., 1982:171, 352 (2 spec.; s. Japan).— Okamura in Masuda et al., 1984:98, pl.
83E.— Yatou in Okamura and Kitajima, 1984:233. 368 (4 spec; Okinawa Trough, 300–550 m).
Coelorinchus anatirostris: Iwamoto, 1990:141 (compiled).— Shen et al., 1993:168.— Shao et al., 2008: table
2 (5 spec., NET, SWT, 200–441 m).
Caelorinchus anatirostris: Iwamoto and Merrett, 1997:486, fig. 6c (7 spec., New Caledonia, Chesterfield and
Ballona Plateau, 600–855 m).— Merrett and Iwamoto, 2000:743 (7 spec., Vanuatu, New Caledonia,
450–1160 m).— Chiou et al., 2004b:36, 47 (in key, Taiwan, listed).
MATERIAL EXAMINED (20 spec.).— NET: ASIZP 61324 (1, 346 TL), Da-xi; ASIZP 65572 (1,
432 TL), Da-xi; ASIZP 70217 (3, 157–337 TL), Da-xi; ASIZP 70219 (3, 330–358 TL), Da-xi;
ASIZP 70698 (1, 79 HL, 272+ TL); Da-xi; CAS 214596 (2); Da-xi; CAS 214728 (4, 215+-325+
TL), Su-ao. SWT: ASIZP 65540 (4, 160+-255 TL), CD 138, 441 m. SCS: CAS 224495 (ex ASIZP
65594) (1, 63.1 HL, 227 TL), CD 141, 985–1110 m.
DISTINGUISHING FEATURES.— 1D II 8–10; P i16–i18; GR-I (inner) 5–7 (total). Scales below
midbase 1D 3.0–4.5, below 2D 4–6; pyl. caeca 21–30. Snout pointed, length moderate, 38–45%
HL, 1.3–1.6 times orbit; preoral length 36–40% HL; anterolateral margin of snout completely sup-
ported by bone; orbit 26–31% HL; upper jaw 21–27% HL. Nasal fossa scaled ventrally; underside
of head naked except for two small scale patches under orbit and preopercle angle; light organ
group II of Iwamoto (1990), fossa narrow and short, not extending to V bases; median rostral ridge
scales 9–12, with spinules radiating laterally and posteriorly; body scales covered with narrowly
triangular spinules in 4–10 (usually 6–8) slightly divergent, ridgelike rows, median row strongest.
Dorsally grayish, ventrally heavily peppered over ivory ground, bluish over abdomen, pale over
chest; mouth and gill cavity grayish to blackish; leading edge of 1D black, other fins dusky to pale.
Attains about 430 mm TL.
DISTRIBUTION.— Widely distributed in the w. Pacific from s. Japan and East China Sea, Tai-
wan, New Caledonia, Chesterfield and Bellona Plateau, Vanuatu, and off ne. Australia. Depth range
300–1160 m.
REMARKS.— An apparent distributional gap in the Philippines and East Indies is somewhat
disconcerting; specimens from the sw. Pacific should be carefully compared with those from Tai-
wan and points north. Specimen CAS 224495 from the SCS has scale spinules that are short and
uniform, unlike those in others in the materials examined, and the snout seemed too slender for the
species. It is tentatively included here, but should be further compared with other closely related
species. Among Taiwan species, C. anatirostris is most closely similar to C. productus and
C. asteroides, but the former has stronger and fewer (3–5) divergent rows of spinules on body
scales and the latter has fewer scales (6–8) on the median rostral ridge. Fukui et al. (2009) syn-
onymized C. anatirostris and C. productus, but we have reservations about accepting their conclu-
sion (see discussion in description of C. productus).
Coelorinchus asteroides Okamura, 1963
Coelorhynchus asteroides Okamura, 1963:21, figs. 1–4 (holotype, FAKU 23801, 7 paratypes; off Owase, Mie
Pref., Japan).— Okamura, 1970:189–192, pl. XLI, text-figs. 81, 82 (8 spec., s. Japan, 320–360 m).— Oka-
mura in Masuda et al., 1984:98, pl. 83F.— Yatou in Okamura and Kitajima, 1984:235, 369 (5 spec., Oki-
nawa Trough).
Caelorinchus asteroides: Chiou et al., 2004b:43, fig. 9 (2 spec., NE, SW Taiwan).
Coelorinchus asteroides: Shao et al., 2008: table 2 (3 spec., NET, SWT, 100–452 m).
50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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MATERIAL EXAMINED (8 spec.).— NET: ASIZP 61339 (1, 302 TL); Da-xi; ASIZP 65658 (1,
250+ TL); Da-xi. SWT: ASIZP 61340 (1, 310 TL); Dong-gang; ASIZP 62194 (2, 224–224 TL);
Dong-gang; ASIZP 65560 (1, 230+ TL); CD 140, 280–452 m; ASIZP 70617 (1, 159 TL); Dong-
gang. Other specimens: ASIZP 68016 (1, 375 TL); Aurora, 909–922 m.
DISTINGUISHING FEATURES.— 1D II, 7–9; P i15–i19; GR-I (inner) 5–7 (total); scales below
midbase 1D. 3.0–4.5, below 2D. 4.0–5.5; pyl.caeca 40–50. Snout fairly broad, tipped with a slen-
der sharp scute, length 39–46% HL, 1.3–1.5 times orbit, preoral length 32–35% HL; anterolateral
margin of snout completely supported by bone; orbit 25–31% HL; upper jaw 22–29% HL. Nasal
fossa naked ventrally; underside of head naked; median rostral series of scales 6–8, with spinules
radiating in all directions; body scales with short, sharp lanceolate to triangular spinules in 4–11
slightly divergent rows, the median row strongest. Light organ group II, fossa narrow and short.
Ground color grayish, silvery ventrally, abdomen bluish, paler anteriorly on chest; mouth dark, gill
cavity blackish. Attains more than 390 mm TL.
DISTRIBUTION.— Geographically very confined; East China Sea from s. Japan to SCS off Tai-
wan, in 100–600 m.
REMARKS.— First recorded from Taiwan by Chiou et al. (2004b). Among the Taiwan mem-
bers of the genus, C. asteroides is most likely to be confused with C. anatirostris, but it has fewer
platelike scales on the median rostral ridge, and the spinule rows on these scales are arranged radi-
ally in all directions, compared to the lateral and posterior orientation of rows in C. anatirostris.
Coelorinchus brevirostris Okamura, 1984
Coelorhynchus brevirostris Okamura in Okamura and Kitajima, 1984:225 (holotype, BSKU 29562; Okinawa
Trough, 25°47.4ʹN, 124°23.4ʹE, 600 m.).
Caelorinchus brevirostris: Chiou et al., 2004b:301 (redescription, 11 spec., Taiwan).
Coelorinchus brevirostris: Shao et al., 2008: table 2 (6 spec., NET, ET, SCS, 445–1185 m).
MATERIAL EXAMINED (21 spec.).— NET: ASIZP 61350 (1, 175+ TL), Da-xi; ASIZP 61351 (1,
213+ TL), Da-xi; ASIZP 61352 (1, 144 TL), Da-xi; ASIZP 65613 (1, 175+ TL), CP 120, 520–640
m; ASIZP 65518 (1, 183+ TL), 24.66°N, 122.18°E, CD 209, 508–522 m; ASIZP 66806 (1, 44.6
HL, 163+ TL,); CP 315, 509 m; ASIZP 66814 (4, 110–200 TL); CD 311, 516 m; CAS 215542 (2,
51.9 HL, 215+TL); Da-xi; CAS 228339 (1, 215+ TL); Nan-fang-ao. ET: ASIZP 65519 (1, 215 TL)
and ASIZP 65615 (1, 134+ TL), CD 210, 445–1185 m. SCS: ASIZP 65675 (1, 140+ TL), CD 311,
516 m; ASZIP 66170 (1, 196+ TL) and ASIZP 66191 (1, 170+ TL), OCP 313, 513 m; ASIZP 66186
(1, 194+ TL) and ASIZP 66874 (1, 92+ TL), 21.67°N, 117.72°E, CP 314, 506 m; CAS 224494 (ex
ASIZP 66182) (1, 185 TL), 21.67°N, 117.72°E; coll. P-F Lee, 17 Aug. 2005.
DISTINGUISHING FEATURES.— 1D II 7–8; P i14–17; GR-I (inner) 6–8 (total); scales below mid-
base 1D 4.5–5.5, below 2D 4.5–5.5; pyl.caeca 13–16. Snout rather short, broad, and depressed,
39–44% HL; terminal scute small; anterolateral snout margin incompletely supported by bone;
orbit 25–28% HL; 1.5–1.7 in snout length; upper jaw 21–24% HL; body terete. Underside of snout
and head naked; nasal fossa scaled; body scales large, with 5–8 parallel spinule rows. Second spin-
ous ray of 1D prolonged, 1.3 times HL; rays of 2D well developed, about as long as opposite rays
in A. Light organ group IV (of Iwamoto in Cohen et al. 1990), extends from anus to just behind
isthmus. Body without prominent markings. Attains about 220 mm TL.
DISTRIBUTION.— East China Sea (Okinawa Trough) n. of Ishigaki Is. to sw. Taiwan in South
China Sea at depths of 400–1185 m.
REMARKS.— The species was originally described from a single specimen taken in the East
China Sea in 600 m, but subsequently redescribed by Chiou et al. (2004b) from 11 specimens col-
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 51

lected from ne. Taiwan. More specimens were recently collected from the South China Sea at
depths between 445 and 1185 m (see Shao et al. 2008: table 2). Our specimens agree well with the
original description except that they had fewer P rays (14–17 versus 19) and a shorter barbel
(16–20% of orbit versus 24.7%). However, one of us (NN) re-examined the holotype and found
P i16–i17 and barbel length 15% of orbit diameter, which is in agreement with our Taiwan speci-
mens.
Among the Taiwan species of Coelorinchus, C. brevirostris is likely to be confused only with
the recently described C. fuscigulus and C. cingulatus, but the lack of body markings distinguish-
es it from other members of the genus having an elevated 2D and long light organ with the anteri-
or fossa just posterior to isthmus.
Coelorinchus cingulatus Gilbert and Hubbs, 1920
Coelorhynchus cingulatus Gilbert and Hubbs, 1920:480, fig. 15 (holotype, USNM 78221, South China Sea,
near Taiwan, 421 m; paratype, USNM 78223, off n. Luzon, 410 m).
Coelorinchus cingulatus: Okamura in Okamura and Kitajima, 1984:229, 366 (1 spec., Okinawa Trough, 250
m). Shen 1984:146.— Shao et al., 2008: table 2 (17 spec., NET, SWT, SCS, 236–1211 m).
Caelorinchus cingulatus: Iwamoto and Merrett, 1997:493–495, fig. 8 (17 spec., New Caledonia region, e.
coast Australia; 480–580 m).— Merrett and Iwamoto, 2000:744 (5 spec., New Caledonia, Vanuatu,
460–525 m).— Chiou et al., 2004b:37, 47 (in key, listed from Taiwan).
MATERIAL EXAMINED (29 spec.).— NET: ASIZP 63249 (1, 233 TL), Da-xi; ASIZP 64545 (2,
152–155 TL), Nan-fang-ao; ASZIP 65662 (1, 232 TL), Da-xi; ASIZP 70211 (5, 263–304 TL), Da-
xi; ASIZP 70245 (1, 205 TL), Da-xi. SWT: ASIZP 65520 (6, 120–173 TL), CD 137, 316–477 m;
ASIZP 65582 (2, 195–200 TL), CD 141, 985–1110 m; ASIZP 65583 (5, 175–200 TL), CD 136,
998–1211 m; ASIZP 65605 (2, 160–170 TL), CD 138, 441 m; ASIPZ 66183 (1, 240 TL), OCP 313,
513 m. Other specimens: ASIZP 67859 (1, 234 TL), Aurora, 422–431 m; ASIZP 67962 (1, 154
TL), Aurora, 431–493 m; ASIZP 68017 (1, 226 TL), Aurora, 357–367 m.
DISTINGUISHING FEATURES.— 1D II 8–9; P i16–19; GR-I (inner) 6–8 (total); scales below mid-
base 1D 3.5–5.5, below 2D 4.5–6.5; pyl.caeca 10–15. Snout sharply pointed, 43–50% HL; termi-
nal scute slender, sharp; anterlateral snout margin incompletely support by bone; orbit 22–26% HL,
1.6–2.2 in snout length; upper jaw 19–24% HL; body somewhat cylindrical, greatest width slight-
ly less than greatest depth. Underside of snout and head naked; nasal fossa naked or sparsely
scaled; body scales large, densely covered with 7–15 parallel spinule rows. Second spinous ray of
1D slightly prolonged; rays of 2D well developed, almost as long as opposites in anal fin. Light
organ group IV, extends from anus to just behind isthmus. A series of dark saddles on body, ante-
rior two directed obliquely downwards and forward. Attains about 300 mm TL.
DISTRIBUTION.— Originally described from the South China Sea near Taiwan and n. Luzon,
but subsequently recorded from Japan (East China Sea), the e. coast of Australia, and the regions
around New Caledonia, Vanuatu, and Loyalty Island. It was collected in Taiwan (NET, SWT) at
depths between 236 and 1211 m.
REMARKS.— Specimens from the sw. Pacific show slight differences from those from the n.
hemisphere, but we have not been able to tease out any specific characters that would differentiate
the populations. The species is most likely to be confused with C. fuscigulus, but that species lacks
an elongated 1D spinous ray and has differences in body markings.
52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Coelorinchus divergens Okamura and Yatou, 1984
Figures 5A–C.
Coelorincus divergens Okamura and Yatou in Okamura and Kitajima, 1984:236–239 (holotype, BSKU 26865,
Okinawa Trough [East China Sea] sw. of Kyushu, Japan, 1000 m; paratype, BSKU 33464, Okinawa
Trough, 780–810 m).— Shao et al., 2008: table 2 (15 spec., NET, SCS, SWT, 646–1110 m; first record
from Taiwan).
MATERIAL EXAMINED (17 spec.).— NET: ASIZP 64145 (1, 280+ TL), CP 235, 764 m; ASIZP
65575 (1, 286 TL), Da-xi; ASIZP 65577 (1, 245 TL), CP 196, 646–787 m. SWT: ASIZP 63799 (1,
338 TL), CD 193, 821 m; ASIZP 65538 (1, 330 TL), CD 141, 985–1110 m. SCS: ASIZP 66084 (1,
74 HL, 212+ TL), ASIZP 66788 (2, 260+-295+ TL), and ASIZP 66838 (1, 166+ TL), CD 321, 954
m; ASIZP 66752 (5, 225+-350+ TL) and CAS 224888, ex ASIZP 66752 (2, 323+-385+ TL), CD
320, 731 m. Other specimens: ASIZP 68052 (1, 398 TL), Aurora, 944–1004 m.
DISTINGUISHING FEATURES.— 1D II, 7–8; P i17–i18; GR-I (inner) 8(total); scales below mid-
base 1D 4–5, below 2D 4.5–5.5; lateral line scales over distance equal to pre-1D length 30–31;
pyl.caeca 9–11. Snout broadly spade-shaped (viewed dorsally), sharply pointed, length 38–45%
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 53
A
B
C
FIGURE 5. Coelorinchus divergens Okamura and Yatou, 1984. A. ASIZP 64145, 280+ mm TL, fresh. B-C. CAS 224888,
385+ mm TL, preserved, dorsal view (B) and ventral view (C) of head.

HL, terminal scute relatively small but pointed, anterolateral margin snout incompletely supported
by bone; orbit 24–28(32)% HL, 1.4–1.8 in snout; interorbital 23–27%, postorbital 32–39%, orbit-
preopercle 34–37%, upper jaw 26–31%, barbel 8–14%; preanal length 161–163%; height 1D
48–49%; length P 41–42%; isthmus to A 38–40%. Nasal fossa sparsely scaled anteroventrally;
scales atop head with 2–5 crestlike rows of spinules in parallel to slightly divergent rows; scales on
underside of head with broad, high spinules aligned in one (rarely 2) crestlike row; body scales
large, thick, adherent, spinules in about 5 divergent ridgelike rows, middle row enlarged, spinule
rows flanking middle row much lower, incomplete, not reaching scale margin. Light organ short,
narrow, Group II. Ground color in alcohol brownish or grayish; fins dark or dusky; mouth, gill cav-
ities, branchiostegal membranes, and jaws blackish. Attains at least 40 cm TL.
DISTRIBUTION.— Known from East China Sea, Taiwan, and South China Sea in 646–1110 m.
REMARKS.— Our specimens (as reported by Shao et al. 2008) represent the first record of the
species from Taiwan and the South China Sea. According to Okamura and Yatou (in Okamura and
Kitajima 1984) this species is very similar to C. parallelus Günther and C. kermadecus Jordan and
Gilbert, but can be distinguished from the former in having 2–5 spinule rows (vs. single row) on
scales atop head, slightly shorter snout (38–45% HL vs. 42–48% HL), and longer upper jaw
(26–31% vs. 22–26%). Coelorinchus kermadecus differs from C. divergens in having 3–5 spinule
rows (vs. 1 row) on scales on the underside of head, a longer snout (44–48% HL), and shorter upper
jaw (23–26%).
Coelorinchus formosanus Okamura, 1963
Coelorhynchus formosanus Okamura, 1963(Mar.):37 (holotype FAKU 35856, Da-xi, Taiwan; 4 paratypes,
FAKU 35857–60).— Okamura, 1970:161–165, pl. XXXIV, text-figs. 65, 66 (descr. from type specimens).
Coelorhynchus intermedius Chu and Lo in Chu, Chan and Chen, 1963 (Aug.):173, fig. 139 (East China
Sea).— Xiong, Zhan, and Deng 1988:187–188, fig. 148 (4 spec., 164–238 mm TL; East China Sea off
Ryukyu Is., 28°48ʹN, 127°00ʹE, 213–285 m).
Coelorhynchus abbreviatus Chu and Lo in Chu, Chan and Chen, 1963(Aug.):174 (East China Sea).
Coelorinchus formosanus: Okamura in Masuda et al., 1984:97, pl. 82–J.— Shao et al., 2008: table 2 (9 spec.,
NET, SCS, SWT, 600–1110 m).
Caelorinchus formosanus: Iwamoto, 1990:158 (descr.). Shen et al., 1993 (descr.).— Chiou et al., 2004b:37,
47 (in key, list).
MATERIAL EXAMINED (38 spec.).— NET: FAKU 35836 (holotype), Da-xi; FAKU 35857
(paratype), Da-xi; ASIZP 57976 (1, 184 TL), Da-xi; ASIZP 61048 (1, 220 TL), Da-xi; ASIZP
61342 (1, 272 TL), Da-xi; ASIZP 65584 (3, juveniles), CP 119, 123–140 m; ASIZP 65650 (1, 251
TL), Da-xi; ASIZP 65661 (1, 260+ TL), Da-xi; ASIZP 65664 (1, 310 TL), Da-xi; ASIZP 61342 (1,
272 TL), Da-xi; ASIZP 70693 (5, 224–333 TL), Da-xi; ASIZP 70714 (1, 154 TL), Da-xi; CAS
214457 (2, 278–297 TL), Su-ao; CAS 224169 (1, 242 TL), Da-xi. SWT: ASIZP 65595 (4, 215–305
TL), CD 141, 985–1110 m; ASIZP 65822 (1, 218 TL), Dong-gang; ASIZP 58024 (1, 173 TL),
Dong-gang; ASIZP 62195 (1, 168 TL), Dong-gang; ASZIP 62196 (2, 156–160 TL), Kaoshiung;
ASIZP 62197 (2, 260–297 TL), Dong-gang; ASIZP 70661 (3, 84–165 TL), Dong-gang. NT: ASIZP
61009 (1, 215 TL), Jin-shan; ASIZP 65568 (1, 326 TL), Da-xi. No data: ASIZP 65578 (1, 322 TL).
DISTINGUISHING FEATURES.— 1D II, 8–10; P i15–i18; GR-I (inner) 7–9 (total); scales below
midbase 1D 3.5–4.5, below 2D 3.5–4.5; pyl.caeca 9–12. Snout sharply pointed, length 42–45%
HL, anterolateral margin snout incompletely supported by bone; orbit 22–26% HL, 1.5–2.0 in
snout; upper jaw 28–35% HL. Nasal fossa and broad areas anteriorly and laterally atop snout
almost entirely naked; underside of snout naked except along anterolateral margins where scales
broadly overlap edge; underside of head behind mouth and mandibular rami scaly; short triangular
54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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spinules on body scales in irregularly quincunx to widely divergent rows. Light organ long, extends
forward to near isthmus, Group IV. Ground color brownish to grayish, irregular darker blotches
over dorsal portions of body, silvery ventrally; fins dark or dusky; mouth whitish, gill cavities,
gular and branchiostegal membranes, and jaws dark dusky; 1D black along leading edge, dark on
basal half or so, V and A blackish distally. Attains at least 326 mm TL. (Adapted from Okamura
1970, with additions from current material.)
DISTRIBUTION.— Known from East China Sea e. of Amami Is. (Ryukyu Islands) and Taiwan
(NET, SWT, and SCS) in depths between about 100 and 600 m. Most abundant in 100–400 m.
REMARKS.— Coelorinchus formosanus is the most abundant grenadier species in Taiwan in
depths less than 600 m (Wu 2002). Among the Group IV species of Coelorinchus of Taiwan, it is
alone in having a scaly posterior underside of head. Okamura (1970:151) synonymized without
comment C. intermedius Chu and Lo, 1963 and C. abbreviatus Chu and Lo, 1963 with C. for-
mosanus. Xiong et al. (1988:148), however, considered C. intermedius as distinct, based on lesser
amounts of scaly areas under the head and below the nasal fossa in the latter species. We have not
been able to verify this with comparative material. They also provided no comment on the status
of C. abbreviatus, suggesting agreement with Okamura’s assessment of the nominal species.
Coelorinchus fuscigulus Iwamoto, Ho, and Shao, 2009
Coelorinchus fuscigulus Iwamoto, Ho, and Shao, 2009:40–45, figs. 1–2 (holotype ASIZP 70169, 74.8 mm
HL, 322 mm TL; NET, Da-xi, 24.94°N, 121.9°E; 9 paratypes NET and East China Sea).
Coelorinchus cylindricus (non Iwamoto and Merrett, 1997): Shao et al., 2008: table 2 (listed, 1 spec. NET,
400–600 m).
MATERIAL EXAMINED.— All type specimens as listed in original description (Iwamoto et al.,
2009). ASIZP 70169 (holotype, 74.8 HL, 322 TL), Da-xi,); ASIZP 63249 (1, paratype, 56.1 HL,
233 TL), Da-xi; ASIZP 66922 (1, paratype, 66.2 HL, 286 TL), CP 248, 536 m; ASIZP 66973 (1,
paratype, 72.7 HL, 293 TL), Nan-fang-ao; ASIZP 70168 (1, paratype, 77.1 HL, 301+ TL), Da-xi;
CAS 228337 ( 2 paratypes, ex. ASIZP 70168, 66.9–74.1 HL, 302+-285+ TL), Da-xi; CAS 228338
(1, paratype, 66.9 HL, 266 TL), Nan-fang-ao; East China Sea: ASIZP 63193 (1, paratype, 52.2
HL, 228 TL) and CAS 224492 (1, paratype, ex. ASIZP 63193, 45.4 HL, 190 TL), Diao-yu-tai
Archipelago, Yilan, Taiwan.
DISTINGUISHING FEATURES.— 1D II, 8–9; P i16–i18; GR-I (inner) 7–9 total; scales below mid-
base 1D 5.5–6.5, below 2D 6.5–7.5; pyl.caeca 19–24. Snout produced and sharply pointed, length
36–41% HL, 1.4–1.7 times orbit; anterolateral margin of snout incompletely supported by bone;
orbit 24–25% HL, upper jaw 26–30% HL. Nasal fossa scaled ventrally; underside of head naked
except for overlapping scales on anterolateral snout margin; body scales with small, sharp spinules
arranged in 10–13 parallel rows. Light organ Group IV of Iwamoto (1990); anterior fossa near isth-
mus, posterior before anus, no black stripe connecting the two. Rays of 2D well developed, about
equal to opposite rays of anal fin. Overall color medium brown to grayish, but in fresh specimens
silvery on ventral sides of head and trunk (bluish in preserved specimens); 8–11 prominent saddles
on body; fins dusky to blackish; lips and jaws pale; branchiostegal region blackish. Attains at least
322 mm TL.
DISTRIBUTIONs.— Known only from NET and East China Sea in Diaoyutai Archipelago, Tai-
wan, in approximately 600 m.
REMARKS.— Coelorinchus fuscigulus was previously confused with C. cylindricus, a species
known only from the holotype taken off New Caledonia, but that species has a complete bony sup-
port of the anterorlateral snout margin. Among the Taiwan members of the genus, C. fuscigulus is
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 55

most similar to C. brevirostris, C. cingulatus, and C. hubbsi. The body markings easily distinguish
the species from C. brevirostris and C. hubbsi; the blackish branchiostegals that contrast strongly
with the pale gular membrane, the fully blackish 1D, and lack of an elongated 1D spinous ray dis-
tinguish it from C. cingulatus.
Coelorhinchus hexafasciatus Okamura, 1982
Figure 6.
Coelorhynchus hexafasciatus Okamura in Okamura, Amaoka and Mitani, 1982:173, pl. 104 (Holotype: BSKU
30443; Kyushu-Palau Ridge, 26°45.0ʹN, 135°19.0ʹE, 336 m; 31 paratypes from Kyushu-Palau Ridge).
MATERIAL EXAMINED.— ASIZP 71202 (1, 133.3 mm HL, 494+ mm TL), Da-xi.
DISTINGUISHING FEATURES.— 1D II,9; P i17–18; V 7; GR-I (inner) 8; GR-II (outer/inner) 7/9;
scale rows below 1D origin 7.5, below midbase 1D 5.5, below 2D origin 6. Snout conical, 40%
HL; terminal scute rather blunt; anterolateral margin of snout incompletely support by bone; orbit
23% HL, 1.8 in snout length; mouth large, upper jaw 31% HL; posterior end of rictus scarcely
restricted by a lip fold. Premaxillary teeth arranged in wide tapered band; outer series distinctly
enlarged. Light organ restricted as a short naked streak before anus. Underside of head complete-
ly naked except for lower preopercle; nasal fossa scaled ventrally. Body scales large, adhered, cov-
ered with keel-like spinules in 7 widely divergent rows. Second spinous ray of 1D not elongated;
2D poorly developed throughout its length. About 6 dark saddles dorsally on trunk and tail; gill
membrane blackish posteriorly; fins generally dark, but outer V ray somewhat paler. Attains at least
70 cm TL.
DISTRIBUTIONs.— Known from the Kyushu-Palau Ridge, s. Japan, and the East China Sea off
Taiwan, in about 340–1320 m.
REMARKS.— This species may be confused with C. sheni (known only from Taiwan), but the
two are readily distinguished by squamation on the head (underside of head mostly naked in
C. hexafasciatus vs. completely scaled in C. sheni). The specimen herein reported represents the
first record of the species from Taiwanese waters. The species was originally described from the
Kyushu-Palau Ridge, but no additional specimens had been reported after the original description.
56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 6. Coelorinchus hexafasciatus Okamura, 1982. ASIZP 71202, 494+ TL, preserved. Photograph by N. Nakaya-
ma.

Coelorinchus hubbsi Matsubara, 1936
Coelorhynchus hubbsi Matsubara, 1936:358 (holotype, FAKU. Owase, Mie-ken, Japan, 100–200 fm [about
183–366 m]).— Okamura in Okamura et al., 1982:166 (photograph of fresh specimen), 169 (comparison
with C. matsubarai).
Coelorinchus hubbsi: Okamura in Masuda et al., 1984:96, pl. 82–L (photograph).— Shen et al., 1993:169.—
Shao et al., 2008: table 2 (2 spec., NET, 100–400 m).
Caelorinchus hubbsi: Chiou et al., 2004b:37, 47 (in key, list).
MATERIAL EXAMINED (2 spec.).— NET: ASIZP 63189 (1, 260 TL), Diao-yu-tai; ASIZP 64536
(1, 162 TL), Diao-yu-tai.
DISTINGUISHING FEATURES.— 1D II, 8–10; P i15–i18; GR-I (inner) 8–10 total; scales below
midbase 1D 4.0–4.5, below 2D 5.5–6.5; pyl.caeca 20–28. Snout produced and sharply pointed,
length 38–45% HL, 1.26–1.65 times orbit; anterolateral margin of snout incompletely supported by
bone; orbit 26–32% HL, upper jaw 22–27% HL. Nasal fossa scaled ventrally; underside of head
naked except for overlapping scales on anterolateral snout margin; body scales with slender, sharp
spinules, arranged in slightly divergent to subparallel rows. Light organ long, extending forward
nearly to isthmus, Group IV of Iwamoto (1990). Rays of 2D well developed, about equal to oppo-
sites of anal fin. Ground color grayish, silvery on sides of head and trunk; three dark longitudinal
streaks on side of body; underside of head and abdomen densely peppered; fins blackish to light
dusky; mouth and gill cavities blackish. Attains at least 260 mm TL.
DISTRIBUTIONs.— Known from s. Japan to ne. Taiwan; restricted in Taiwan to off a small
island called “Fishing Island [Uotsuri-shima of the Senkaku Shoto],” at depths less than 400 m.
REMARKS.— This species has an apparently restricted distribution; it was not captured during
the Japanese expeditions off the Kyushu-Palau Ridge (Okamura et al. 1982) and the Okinawa
Trough (Okamura and Kitajima 1984). The prominent body stripes on this species distinguish it
from all other Taiwan Coelorinchus.
Coelorinchus japonicus (Temminck and Schlegel, 1846)
Macrurus japonicus Temminck and Schlegel, 1846:256, pl. 112, fig. 2, 2A–B (bays in provinces Oomura and
Shimabara, Japan. Lectotype: RMNH 3476. Paralectotypes: RMNH D1405–1406 [2 stuffed]).
Coelorhynchus japonicus: Gilbert and Hubbs, 1916:178–179 (5 spec., 158–307 mm TL; Japan).— Okamura,
1970:183–186, pl. XL, text-figs. 78, 79 (93 spec., 138–656 mm TL; range given as “tropic and subtrop-
ic regions of Indo-western Pacific including southern part of Japan”).— Yatou in Okamura et al.,
1982:181, photo 106, 353 (8 spec., 438–750 m TL, 300–700 m).
Coelorinchus japonicus: Okamura in Masuda et al., 1984:98, pl. 83–D.— Yatou in Okamura and Kitajima,
1984:231, photo 163, 367 (10 spec., 315–680 mm TL, 560–1000 m).— Xiong, et al., 1988:190–192, fig.
152 (4 spec., 275–373 mm TL; East China Sea, 490–716 m).— Iwamoto, 1990:162–163, fig. 378.—
Shen et al., 1993:169 (descr.).— Kim et al,. 2005:172 (Korea).— Shao et al., 2008: table 2 (3 spec., NET,
100–650 m).
Caelorinchus japonicus: Chiou et al., 2004b:36, 47 (in key, list).
MATERIAL EXAMINED (7 spec.).— NET: ASIZP 56349 (1, 292 TL), Da-xi; ASIZP 65567 (1,
370+ TL), Da-xi; ASIZP 63280 (2, 370–600 TL), Da-xi; ASIZP 70216 (2, 363–450 TL), Da-xi;
CAS 214622 (1, 300 TL), Da-xi.
DISTINGUISHING FEATURES.— 1D II, 8–10; P i16–i20; GR-I (inner) 7–9 total; scales below
midbase 1D 4.5–6.0, below 2D 5.5–7.5; pyl.caeca 41–60. Snout produced and sharply pointed,
length 40–48% HL; anterolateral margin of snout completely supported by bone; orbit 22–30%
HL, upper jaw 23–29% HL. Nasal fossa scaled ventrally; underside of head, including mandibular
rami scaly; most head scales with single crestlike spinule row; body scales with broadly triangular
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 57

spinules lacking strong transverse buttresses, arranged in 3–7 slightly divergent, sharply crestlike
rows. Head ridges stout, heavily spinulated. Light organ short, Group I of Iwamoto (1990). Ground
color brownish to grayish, paler below, darker over abdomen and gill covers; gular and BR mem-
branes pale, lips white, mouth and gill cavities blackish; fins blackish to light dusky. Attains at least
750 mm TL. (Description mostly after Okamura, 1970.)
DISTRIBUTION.— From s. Japan (including Japan Sea), East China Sea (Okinawa Trough), to
ne. Taiwan; 100–1000 m, but most abundant off Japan between 300 and 600 m.
REMARKS.— A large species with a substantial vertical distribution. One of the few grenadiers
recorded as occurring in the Japan Sea, although that record is questionable and must be verified
(see Kim et al. 2009:113). Among the Taiwan Coelorinchus with short light organ immediately
anterior to A origin, C. japonicus is distinguished by the combination of anterolateral snout margin
completely supported by bone, scales on underside and top of head with spinules in one crestlike
row, and body scales with broad spinules in 3–7 sharp crestlike rows.
Coelorinchus kamoharai Matsubara, 1943
Coelorhynchus kamoharai Matsubara, 1943:136, fig. 4 (holotype, FAKU 2498 [apparently lost], Kumano-
Nada, Japan; paratypes, FAKU 1593–1596 [4], 2496 [1], and 4 other unnumbered lots of 1, 3, 3 and 2 spec-
imens, respectively [apparently lost]).— Okamura, 1970:159–161, text-fig. 64, pl. I, fig. b (111 spec.,
115–284 TL; s. Japan)
Coelorinchus kamoharai: Okamura in Masuda et al., 1984:97.— Iwamoto, 1990:164–165, fig. 382 (com-
piled).— Yatou in Okamura and Kitajima, 1984:221 (fig. 157 on p. 220), 365 (10 spec.; East China Sea
[Okinawa Trough], s. Japan, Taiwan; 220–400 m).— Shen et al., 1993:169 (descr.).— Shao et al., 2008:
table 2 (6 spec., NET, 100–650 m).
Caelorinchus kamoharai: Chiou et al., 2004b:47 (list).
MATERIAL EXAMINED (9 spec.).— NET: ASIZP 61336 (4, 158–211 TL), Da-xi; ASIZP 65796
(1, 176 TL), Da-xi; ASIZP 66336 (1, 221 TL), Nan-Fang-Ao; CAS 56041 (1, 152 TL), Dong-gang;
NMMST-P (1, 172 TL), Hsiao-liu-chiu. Other specimens: ASIZP 67947 (1, 190 TL), Aurora,
269–277 m.
DISTINGUISHING FEATURES.— 1D II, 8–10; P i15–i19; GR-I (inner) 9–12 total; scale below
midbase 1D 3.5–4.0, below 2D 4.5–5.0; pyl.caeca 7–13. Snout sharply pointed, length 31–37%
HL, anterolateral margin snout incompletely supported by bone; orbit 24–29% HL; upper jaw
28–34% HL. Nasal fossa and broad areas anteriorly and laterally atop snout naked; underside of
head naked except along anterolateral margin of snout where scales broadly overlap edge; under-
side of head covered with unpaired black papillae; spinules on body scales short, broad based, in
irregularly quincunx order. Light organ long, extends forward to near isthmus, Group IV. Dorsum
brownish gray with irregular darker blotches, remainder of body silvery; chest and area around
periproct and anal-fin origin blackish or dark; fins dusky; median nasal process black; mouth and
gill cavities white; gular and branchiostegal membranes heavily peppered, but underside of snout
covered with hair-like black papillae. Attains at least 284 mm TL.
DISTRIBUTION.— Southern Japan, East China Sea, ne. Taiwan, and Philippines, in depths of
100–650 m. The Philippine specimen represents the southernmost record of the species.
REMARKS.— Coelorinchus kamoharai is closely similar to C. multispinulosus, C. formosanus,
and C. longissimus, but differs from them in its body markings; in addition, C. multispinulosus has
slender, dense scale spinules on body scales and sparse, paired papillae under the head, C. for-
mosanus has scales posteriorly on underside of head; and C. longissimus has scale spinules
arranged in definite rows, small scales on nasal fossa, and underside of snout almost immaculate.
Five paratypes (FAKU 1593–1596 and 2496) were rediscovered by one or us (NN) from the FAKU
collection.
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Coelorinchus kishinouyei Jordan and Snyder, 1900
Coelorhynchus kishinouyei Jordan and Snyder, 1900:376–377, pl. XX (holotype [unique], USNM 49395;
Misaki, Japan.).— Gilbert and Hubbs, 1916:170–172 (7 spec., 192–295 TL, Suruga Gulf, Japan).— Oka-
mura, 1970:172–175, pl. XXXVI, text-fig. 71 (92 spec., 123–362 TL; s. Japan, 250–450 m).
Coelorinchus kishinouyei: Okamura in Masuda et al., 1984:97.— Iwamoto, 1990:167 (compiled.).— Shen et
al., 1993:169 (descr.).— Shao et al., 2008: table 2 (15 spec., NET, SCS, SWT, 227–1211 m).
Caelorinchus (Paramacrurus)kishinouyei: Chiou et al., 2004b:47, table 1 (listed).
MATERIAL EXAMINED (55 spec.).— NET: ASZIP 56351 (4, 109–267 TL), Da-xi; ASIZP
58020 (8, 109–232 TL), Da-xi; ASIZP 58266 (4, 139–158 TL), Da-xi; ASIZP 58633 (2, 105–110
TL), Da-xi; ASIZP 58647 (2, 186–199 TL), Da-xi; ASIZP 60245 (2, 168–248 TL), Da-xi; ASZIP
61049 (1, 181 TL), Da-xi; ASZIP 63148 (1, 160 TL), Da-xi; ASIZP 65082 (1, 181 TL), Da-xi;
ASIZP 65645 (1, 256 TL), Da-xi; ASIZP 65654 (1, 309 TL), Da-xi; ASIZP 65655 (1, 222 TL), Da-
xi; ASIZP 65656 (1, 222+ TL), Da-xi; ASZIP 65659 (1, 220 TL), Da-xi; ASIZP 65660 (1, 185+
TL), Da-xi; ASIZP 65991 (1, 152 TL), Da-xi; ASIZP 70662 (5, 103–160 TL), Da-xi; ASIZP 70696
(6, 150–212 TL), Da-xi; ASIZP 70732 (1, 210 TL), Da-xi; CAS 214461 (1, 240+ TL), Su-ao; CAS
214612 (2, 192+-213+ TL), Da-xi; CAS 224170 (2, 205–202+ TL), Da-xi. SWT: ASIZ 62158 (1,
216 TL), Dong-gang. SWT: ASIZP 65555 (1, 188 TL), CD 142, 227–335 m; ASIZP 65585 (1, 205
TL), CD 136, 998–1211 m; ASIZP 65596 (1, 205 TL), CD 141, 985–1110 m; ASIZP 70616 (1, 146
TL), Dong-gang; ASIZP 70660 (1, 210 TL), Dong-gang.
DISTINGUISHING FEATURES.— 1D II, 8–10; P i15–i19; GR-I (inner) 9–10 total; scales below
midbase 1D 3.5–4.0, below 2D.5–4.5; pyl.caeca 19–26. Snout sharply pointed, sides gently con-
vex, tipped with a blunt scute, length 33–45% HL, anterolateral margin snout completely support-
ed by bone; orbit 31–38% HL; upper jaw 23–29% HL. Nasal fossa and underside of head naked;
spinules on body scales short, conical, in 8–17 parallel to slightly divergent rows. Light organ
short, extends forward only to base of pelvic fins, Group II. Dorsum brownish, belly blackish, rest
of body silvery, a large (2⁄3orbit diameter) ocellated dark blotch above pectoral fin; underside of
head densely peppered; mandibular rami and gill membranes blackish; fins generally dusky, sec-
ond 1D spine and tips of rays black; mouth and gill cavities whitish. Attains at least 362 mm TL.
DISTRIBUTION.— Off s. Japan, South China Sea, and Taiwan (NET, SWT) in 200 m to more
than 600 m
REMARKS.— Coelorinchus kishinouyei was strangely not recorded by Okamura and Kitajima
(1984) from the Okinawa Trough, where it would be expected owing to its presence in s. Japan and
Taiwan. In this regard, the distribution is similar to that of C. hubbsi. It is relatively abundant in
landings at Da-xi in ne. Taiwan. This highly distinctive species is closely similar to C. jordani,
which has a much smaller pectoral blotch and a longer light organ that extends well forward of the
pelvic bases and onto the chest. The Taiwan specimens of C. kishinouyei appeared to have a slight-
ly longer snout, smaller orbit, darker overall coloration with little or no silvery, with anal fin black
overall, and blotch above pectoral fin more obscure and lacking pale outer ring (i.e., not ocellated).
Coelorinchus leptorhinus Chiou, Shao and Iwamoto, 2004
Caelorinchus leptorhinus Chiou, Shao and Iwamoto, 2004a:299, figs. 1–3 (holotype, ASIZP 061344 and 23
paratypes, Da-xi, ne. Taiwan, 24°54.63ʹN, 122°03.49ʹE, 400–600 m).
Coelorinchus leptorhinus: Shao et al., 2008: table 2 (23 spec., NET, SWT, 100–650 m).
MATERIAL EXAMINED (54 spec.).— NET: ASZIP 58636 (1, 146 TL), Da-xi; ASIZP 58648 (2,
285–311 TL), Da-xi; ASIZP 60246 (1, 192 TL), Da-xi; ASIZP 60247 (1, 310 TL), Da-xi; ASIZP
61344 (1, holotype, 800 TL), Da-xi; ASIZP 61345 (1, paratype, 380 TL), Da-xi; ASIZP 61346 (10,
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 59

paratype, 142–176 TL), Da-xi; ASIZP 61347 (2, paratypes, 480–640 TL), Da-xi; ASIZP 64288 (1,
400 m, 146 TL), Da-xi; ASIZP 65569 (1, 567 TL), Da-xi; ASZIP 65571 (1, 407 TL), Da-xi; ASIZP
65642 (1, 370 TL), KSD sta.4; ASZIP 65646 (1, 230+ TL), Da-xi; ASZIP 65647 (1, 308 TL), KSD
sta.4; ASZIP 65649 (1, 261 TL), KSD sta.; ASIZP 65781 (1, 346 TL), Da-xi; ASIZP 66253 (1, 352
TL), Da-xi; ASIZP 66917 (2, 212–224 TL), CP 248, 526 m; ASIZP 66939 (1, 133 TL),Da-xi;
ASIZP 70214 (2, 334–394 TL ), Da-xi ; ASIZP 70252 (1, 147 TL), Da-xi; ASIZP 70692 (18,
200–375 TL), Da-xi. SWT: ASIZP 66405 (1, 240 TL), Dong-gang; ASIZP 65628 (1, 546 TL), CD
233, 448–526 m.
DISTINGUISHING FEATURES.— 1D II, 8–9; P i17–i18; GR-I (inner) 6–8 total; scales below mid-
base 1D 5.5–7.5, below 2D 5.5–7.5; pyl.caeca 42–48. Snout long, narrow, pointed in lateral view,
tip rounded, length 39–46% HL, anterolateral margin of snout completely supported by bone; orbit
16–19% HL; upper jaw 26–29% HL. Nasal fossa scaly, underside of snout naked, but finely scaled
posteriorly on head; spinules on body scales in 5–8 sharp, slightly divergent ridge rows. Light
organ short, Group II of Iwamoto (in Cohen et al. 1990); anus removed from A origin. Ground color
brown, bluish on belly and halfway onto chest; underside of head, mouth, and gill cavities dark;
fins generally dusky to blackish. Attains more than 850 mm TL.
DISTRIBUTION.— Apparently endemic to Taiwan (NET, SWT, SCS). The species is most abun-
dant in 300–400 m off s. Taiwan and 400–800 m in ne. Taiwan.
REMARKS.— Shen et al.’s (1993) record of C. tokensis was a misidentification of C. leptorhi-
nus. The species is abundant in the bycatch of the deepwater trawl fisheries off Da-xi (NET). It is
most closely similar to the Philippines species C. macrorhynchus Smith and Radcliffe, 1912 in hav-
ing a notably long snout, long jaws, relatively long barbel, and similar light organ. The main dif-
ferences between the two species lie in C. leptorhinus having the underside of snout naked and hav-
ing a slightly shorter snout (preoral length 35–40% HL cf 40–45%).
Coelorinchus longissimus Matsubara, 1943
Coelorhynchus longissimus Matsubara, 1943:140, fig. 5 (holotype, FAKU 1592 [apparently lost]; Kumano-
Nada, Japan).— Okamura, 1970:165–168, pl. XXXV, text-fig. 67 (48 spec., 200–357 mm TL; s. Japan,
280–400 m).
Coelorinchus longissimus: Okamura in Masuda et al., 1984:97, pl. 82–K (compiled).— Yatou in Okamura and
Kitajima, 1984:223, 366, fig. 159 (5 spec., 175–325 mm TL; Okinawa Trough).— Shao et al., 2008: table
2 (3 spec., NET, SWT, 100–650 m).— Kim et al., 2005:172 (Korea; compiled).
Caelorinchus longissimus: Nakabo, 2003:430 (compiled).— Chiou et al., 2004b:43–44, fig. 10 (2 spec., NET,
SWT).
MATERIAL EXAMINED (9 spec.).— NET: ASIZP 61337 (1, 202 TL), Da-xi; ASIZP 70695 (2,
212–235 TL), Da-xi. SWT: ASIZP 57602 (1, 182 TL), Dong-gang; ASIZP 61338 (1, 262 TL),
Dong-gang); ASIZP 70614 (1, 165 TL), Dong-gang. Other material: Philippines: ASIZP 68091
(1, 84 TL), Aurora, 184–200; ASIZP 68423 (2, 100–103 TL), Aurora, 184–200 m.
DISTINGUISHING FEATURES.— 1D II, 8–10; P i16–i20; GR-I (inner) 6–9 total; scales below
midbase 1D 3.0–4.5, below 2D 3.5–4.5; pyl.caeca 16–23. Snout sharply pointed, length 42–45%
HL, anterolateral margin snout incompletely supported by bone; orbit 24–27% HL; upper jaw
23–28% HL. Most of nasal fossa and broad areas atop snout naked; underside of head naked except
along anterolateral margin of snout where scales broadly overlap edge; spinules on body scales
short, weak, slender in 6–14 parallel to slightly divergent rows. Light organ long, extends forward
to near isthmus, Group IV of Iwamoto (in Cohen et al. 1990). Dorsum gray with faint irregular
blotches, remainder of body silvery; fins dusky, 1D with blackish membrane behind long spinous
ray; median nasal process black; mouth white, gill cavities blackish; gular and branchiostegal
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membranes heavily peppered; underside of snout clear, mostly lacking melanophores except along
outer margins. Attains at least 357 mm TL.
DISTRIBUTION.— Pacific coast of s. Japan from Suruga Bay s. into the East China Sea (Oki-
nawa Trough), ne. Taiwan, the South China Sea off Taiwan, and in the Philippines, in 280–400 m.
REMARKS.— This species is closely similar to C. kamoharai, differing only in the slender
scale spinules arrayed in many divergent rows (as compared with short broad spinules in quincunx
pattern) and paler, slightly different pattern of blotches on the body.
Coelorinchus macrochir (Günther, 1877)
Figure 7.
Macrurus macrochir Günther, 1877:438 (holotype, BMNH 1887.12.7.123; off “Inoshima” [= Enoshima],
Japan, Challenger sta. 232, 345 fm [631 m]).
Abyssicola macrochir: Goode and Bean, 1896:417 (compiled, type-species for new genus Abyssicola).—
Gilbert and Hubbs, 1916:183–184–186 (36 spec., 91–634 mm TL; Pacific coast of Japan, 129–437 fm
[236–799 m]).— Okamura, 1970:145–148, pl. XXXII, text-fig. 58 (136 spec., 173–680 mm TL; Japan:
Hokkaido to East China Sea off Kagoshima Prefecture).— Okamura in Masuda et al., 1984:96, pl. 82–G
(compiled).— Okamura in Okamura and Kitajima, 1984:221, 364 (1 spec., 590 mm TL; Okinawa
Trough, 820 m).— Nakabo, 2002:429 (compiled).
Coelorhynchus (Abyssicola)macrochir: Gilbert and Hubbs, 1920:425 (in key).
Coelorinchus macrochir: Iwamoto, 1990:171 (compiled).— Shao et al., 2008: table 2 (2 spec., NET, 100–650
m; first record from Taiwan).— Honma et al., 2008:65–74 (Japan Sea).— Kim et al., 2009:110–112, fig.
5, table 2 (East Sea [Japan Sea], Korea).
MATERIAL EXAMINED (2 spec.).— NET: ASIZP 65574 (1, 424 TL), Da-xi; ASIZP 66972 (1,
570 TL), Da-xi.
DISTINGUISHING FEATURES.— 1D II, 9–11; P i16–i20; GR-I (inner) 10–12 total; scales below
midbase 1D 4.5–6.0, below 2D 5.5–7.0; pyl.caeca 42–48. Snout rather bluntly conical to rounded
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 61
FIGURE 7. Coelorinchus macrochir (Gunther, 1877). ASIZP 55972, 570 mm TL, preserved.

in lateral profile, length 29–36% HL, anterolateral margin of snout incompletely supported by
bone; a distinct curve in anterior portion of suborbital ridge; orbit 27–33% HL; mouth relatively
large, upper jaw 34–42% HL. Nasal fossa and underside of head scaly; spinules on body scales
short, broad, in 4–9 widely divergent rows. Vent about midway between pelvic-fin base and anal-
fin origin; naked fossa of light organ close before vent (Group II of Iwamoto, 1990). Ground color
brown, bluish ventrally on trunk; mouth and gill cavities blackish; fins dusky to blackish. Attains
at least 680 mm TL.
DISTRIBUTION.— Pacific coast of Japan (Hokkaido to Kyushu off Kagoshima), s. Okhotsk
Sea, East China Sea (including Okinawa Trough), Japan Sea off Niigata, ne. Taiwan, and the
Philippines, in 235–830 m.
REMARKS.— Shao et al. (2008: table 2) first recorded the species from Taiwan based on the
two specimens listed above; they represent the southernmost record (lat. 24.8°N) of the species.
Most previous workers have included Coelorinchus macrochir in its own genus, Abyssicola Gün-
ther, based primarily on its dentition: short conical teeth in three series in premaxilla, in two series
in dentary. Okamura (1970:143–144) provided a lengthy diagnosis that readily differentiated the
species from the 14 then-known Japanese species of Coelorinchus. However, when viewed on a
worldwide basis of more than 100 species in the genus, the diagnostic characters enumerated by
Okamura are shared by one or more species of Coelorinchus sensu lato and thus fail to differenti-
ate Abyssicola. Fukui et al. (2010) conducted a detailed study of the eggs and larvae of C. kishi-
nouyei, and using DNA nucleotide sequences, provided a (op. cit. fig. 3) phylogenetic tree which
showed C. macrochir deeply nested within other Japanese Coelorinchus, with its sister group being
C. anatirostris/productus (which they considered as synonyms: see Remarks under C. productus).
Coelorinchus cf. macrorhynchus [sensu Smith and Radcliffe, 1912]
Figure 8.
MATERIAL EXAMINED.— SWT: ASIZP 65531 (1, 73.0 mm HL, 222 mm TL), CD 138, 441 m.
Distinguishing feature.— 1D II 9; P i18–i19; GR-I (inner) 2+6, GR-II (outer/inner) 0+6 /
1+7; scales below midbase 1D 4.5, below 2D 6.5, lateral-line scales over distance equal to pre-D
length 47. Snout sharply pointed, slender and spear-shaped in dorsal view, length 51% HL; antero-
lateral margin of snout completely supported by bone; orbit 23% HL, 2.2 in snout length, upper
jaw 23% HL; nasal fossa densely scaled anteriorly and ventrally; underside of head almost fully
scaled (scales with 1–3 short rows of spinules). Light organ group II of Iwamoto (1990), fossa nar-
row and short, not extending to pelvic-fin bases. Anus separated from A origin by 2 or 3 scale rows.
Spinules on body scales blade-like in 3–6 sharp, divergent, ridgelike rows, closely overlapping,
with height increasing abruptly with posteriormost spinule much larger than anterior ones. Ground
color brownish (somewhat faded), bluish over abdomen but not on chest; mouth and gill cavity
dark; most fins dark dusky to blackish. Attains more than 44 mm TL.
DISTRIBUTION.— Broadly distributed from Philippines to e. and w. coasts of Australia, in
485–1107 m. In Taiwan, the single specimen was collected in 441 m.
REMARKS.— This single juvenile was captured in the same haul as two specimens (ASIZP
65523, 250–265 mm TL) that we identified as C. smithi. It differed from those specimens in hav-
ing a more slender and longer snout (orbit 2.2 into snout, cf. 1.3–2.1 in C. smithi), slightly narrow-
er internasal and interorbital widths (16% HL and 18% HL, respectively, cf. 17–19% and 19–23%
in C. smithi) and the nasal fossa was almost fully covered with small scales antero-ventrally (naked
to sparsely covered in C. smithi). The differences are so slight that we are uncertain whether they
simply represent individual variation; thus, our identification must be considered as tentative. The
presence of this species in Taiwan waters should not be surprising, however, as it was originally
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described from captures in the Philippines in the Verde Island Passage off sw. Luzon. If confirmed,
this specimen represents the first record of the species from Taiwan.
Coelorinchus multispinulosus Katayama, 1942
Coelorhynchus multispinulosus Katayama, 1942:332 (holotype, NSMT-P 18224; Tsuiyama Market, Yogo
Pref., Japan [Japan Sea]).— Okamura, 1970:156–158, pl. XXXIII, text-fig. 63 (89 spec., 154–378 mm TL;
s. Japan, Japan Sea, East China Sea, 150–300 m).
Coelorinchus multispinulosus: Okamura in Masuda et al., 1984:97, pl. 82–H (compiled).— Yatou in Okamu-
ra and Kitajima, 1984:223, fig. 158 (p. 222), 365 (7 spec., 230–325 mm TL; Okinawa Trough in 146–200
m).— Iwamoto, 1990:174–175, fig. 398 (compiled).— Shen et al., 1993:170 (descr.).— Kim et al.,
2005:173 (Korea; compiled).— Shao et al., 2008: table 2 (11 spec., NET, 100–650 m).
Caelorinchus multispinulosus: Nakabo, 2002:430 (compiled).— Chiou et al., 2004b:37, 47 (in key, list).
MATERIAL EXAMINED (46 spec.).— NET: ASIZP 57306 (1, 202 TL), Da-xi; ASIZP 58021 (2,
179–274 TL), Da-xi; ASIZP 58635 (4, 158–255 TL), Da-xi; ASIZP 58699 (2, 179–214 TL), Da-
xi; ASZIP 61084 (6, 210–245 TL), Da-xi, 100 m; ASIZP 61104 (1, 135 TL), Da-xi, 50 m; ASIZP
61333 (1, 236 TL), Da-xi; ASIZP 65174 (1, 227 TL), Da-xi; ASZIP 65651 (1, 180 TL), KSD sta.
4; ASZIP 65653 (1, 189 TL), KSD st. 4; ASIZP 66924 (2, 220–255 TL), CP 250, 220 m; ASIZP
61047 (3, 195–230 TL), Da-xi; ASIZP 70663 (13, 195–245 TL), Da-xi; NMMSTP 0183 (2,
208–220 TL), Da-xi; CAS 214597 (6), Da-xi. NT: ASZIP 55477 (1, 203 TL), Ye-liou; ASIZP
61215 (1, 215 TL), Jim-shan, 100 m.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 63
A
B
C
FIGURE 8. Coelorhynchus cf. macrorhynchus [sensu Smith and Radcliffe, 1912]. ASIZP65531, 222 mm TL. A. lateral
view. B. dorsal view of head. C. ventral view of head.

DISTINGUISHING FEATURES.— 1D II, 8–10; P i13–i17; GR-I (inner) 7–10 total; scales below
midbase 1D 3.5–4.5, below 2D 3.5–4.5; pyl.caeca 11–20. Snout sharply pointed, length 40–45%
HL, scattered paired papillae on underside, anterolateral margins incompletely supported by bone;
orbit 22–27% HL; upper jaw 27–31% HL. Most of nasal fossa and broad areas atop snout naked;
underside of head naked except along anterolateral margin of snout where scales broadly overlap
edge; spinules on body scales weak, slender, in quincunx order. Light organ long, extends forward
to near isthmus, Group IV of Iwamoto (1990). Dorsally on body and nape grayish with prominent
vermiculations and blotches that extend below lateral midline, laterally mostly silvery, blackish
ventrally on trunk, thorax, jaws, and gill membranes; fins dusky to blackish, 1D with blackish
membrane behind long spinous ray; mouth white, gill cavities blackish; underside of snout with
scattered paired papillae. Attains at least 378 mm TL.
DISTRIBUTION.— Pacific coast of s. Japan to ne. Taiwan, Japan Sea, and East China Sea (Oki-
nawa Trough), in depth less than 400 m. Most abundant in depths less than 200 m.
REMARKS.— Coelorinchus multispinulosus is one of the most abundant grenadier in s. Japan
and one of only a few grenadier species known to occur in the Japan Sea (Kim, I.S. et al. 2005;
Kim, S.Y. et al. 2009). Its normal depth distribution is also one of the shallowest of all grenadiers
from the general region. Its body marking is unique among the Taiwan members of the genus.
Coelorinchus cf. notatus [sensu Smith and Radcliffe, 1912]
Figure 9.
MATERIAL EXAMINED.— NMMB-P11958 (90++ mm TL, large portion of tail missing, 35.5
mm HL); Taiwan; Nan-fang-ao; ca. 280 m.
DISTINGUISHING FEATURES.— 1D II, 9; P i16; GR-I (inner) 1+1+6, 8 total; scales below mid-
base 1D 3.5, below 2D 5.5; pyl.caeca 25. Snout sharply pointed in lateral view, sides gently con-
vex in dorsal view, tipped with a short, broad scute, length 43% HL, anterolateral margin snout
completely supported by bone; orbit 28% HL; upper jaw 23% HL. Nasal fossa and underside of
head naked; spinules on body scales short, conical, in 5 or 6 almost parallel rows. Light organ short,
extends forward slightly anterior to base of pelvic fins, Group II. Belly bluish, ventral aspects of
body tail pale; a small (pupil width) ocellated black blotch above pectoral fin; underside of head
almost entirely pale to whitish; mandibular rami and gill membranes pale; fins generally dusky,
second 1D spine and tips of rays blackish; mouth and gill cavities blackish, but lips pale.
DISTRIBUTION.— Taiwan off Nan-fang-ao (NET) in about 280 m.
REMARKS.— This single damaged specimen (tail mostly broken off, scales almost entirely
missing on body and head) belongs amongst those Coelorinchus species that Gilbert and Hubbs
(1920:446) included under the subgenus Paramacrurus Bleeker. Most of the species in this group
have a dark blotch above the base of the pectoral fin, an acute snout slightly longer than diameter
of orbit, lateral margins of snout fully supported by bone, snout with convex lateral profile (viewed
dorsally), anus removed by a short distance from anal-fin origin, and light organ moderately devel-
oped with dermal window extending forward between pelvic-fin bases (and beyond in some
species). The subgenus includes C. kishinouyei and C. jordani from Japan, and at least nine others
from the Philippines-East Indies regions, including C. notatus, C. maculatus, C. velifer, C. sexra-
diatus, C. triocellatus, C. dorsalis, C. macrolepis, C. argus, and C. thurla. The Taiwan specimen
follows very closely the original description and illustration of C. notatus Smith and Radcliffe,
1913, as well as the more-extensive description provided by Gilbert and Hubbs (1920:462–465),
who in a footnote (op. cit. p. 464) stated that none of Radcliffe’s “smaller specimens are true nota-
tus.“ Thus, the species is known only from the holotype. NMMB-P11958 differs from C. notatus
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in having a smaller pectoral blotch, which covers only four diagonal scale rows (cf. six in C. nota-
tus) and a snout with a straight dorsal profile (cf. slightly concave). Gilbert and Hubbs (1920:465)
stated that “the belly is blackish only in advance of the ventrals,” but Radcliffe (1913:137) stated,
“belly with blackish shades showing through scales.” In NMMB-P11958, the entire underside of
the trunk is dark bluish, from the gill membranes to beyond the anal-fin origin. The Taiwan speci-
men appears to have more saddle markings than does C. notatus. A broad dark saddle extends from
before the mid-base of 1D to slightly beyond the origin of 2D. This saddle extends one or two scale
rows below the lateral line, about to the mid-lateral line of the body; it is not as dark as the pec-
toral blotch or the short (5 or 6 rows wide) more-posteriorly situated saddle from which it is sepa-
rated by a pale gap occupying three scale rows wide. A third, longer saddle that does not extend to
the lateral line follows a pale gap of three scale rows; this shallow saddle is about 12 scales long
and 2.5 to 3 scales deep. A fourth short saddle appears to be developed, although the specimen is
so damaged that only parts of the integument remain and the precise limits of the marking could
not be confidently acertained. The second and fourth saddles are about in the positions shown for
the two saddle marks in the original illustration of the C. notatus (Radcliffe, 1913: pl. 30, fig. 3),
which does now show the intervening saddles seen in NMMB-P11958. The relatively poor condi-
tion of the Taiwan specimen does not allow us to confidently assign the specimen to a species. A
judgement awaits the collection and study of additional, better-preserved specimens.
Coelorinchus parallelus (Günther, 1877)
Macrurus parallelus Günther, 1877:439 (syntypes, BMNH 1887.12.7.65–68[4], 1887.12.7.69 [1]; MNHN
1890–0125 [ex BMNH] [1]; NMI block reg. no. 114.1899 [1]; off Inoshima, Japan, Challenger sta. 232,
345 fm [631 m]. Specimens [4] from New Zealand and Kermadec Islands subsequently renamed C. ker-
madecus Jordan and Gilbert, 1904).
Coelorhynchus parallelus: Okamura, 1970:198–200, pl. XLIII, text-fig. 86 (2 spec., 400–450 mm TL; Japan).
Coelorinchus parallelus: Kyushin et al., 1977:42 Okamura in Masuda et al., 1984:98–99, pl. 83–3 (com-
piled).— Yatou in Okamura and Kitajima, 1984:245, fig. 171 (p. 244), 370 (9 spec., 275–480 mm TL; Oki-
nawa Trough, in 650–990 m).— Iwamoto, 1990:178–179, fig. 404 (compiled).— Iwamoto and Merrett,
1997:500–502, fig.12 (3 spec., New Caledonia region, 412–970 m).— Iwamoto et al., 1999:52.— Merrett
and Iwamoto, 2000:746–747, fig. 12 (5 spec., Vanuatu, Norfolk Ridge, Lord Howe Rise, in 764–1124 m).
— Shen et al., 1993:170 (descr.).— Shao et al., 2008: table 2 (3 spec., SWT, 100–650 m).
Caelorinchus parallelus: Chiou et al., 2004b:36, 47 (in key, list).
Coelorinchus commutabilis: Shao et al., 2008: table 2 (1 spec., SCS, 731 m; first record from Taiwan).
MATERIAL EXAMINED (6 spec.).— NET: ASIZP 65570 (1, 635+ TL), KSD sta.3; ASIZP 70690
(1, 380 TL), Da-xi. SWT: ASIZP 65542 (1, 405 TL), CD 138, 441 m; ASIZP 65630 (1, 146 HL,
404 TL) and ASIZP 70271 (1, 430 TL), CD 230, 810–850 m; ASIZP 66108 (1, 350+ TL), OCP 301,
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 65
FIGURE 9. Coelorinchus cf. notatus [sensu Smith & Radcliffe, 1912], NMMB-P11958, 90+ mm TL.

687 m. SCS: ASIZP 66785 (1, 62 HL, 210 TL) and CAS 224496 (ex ASIZP 66785) (1, 61 HL, 220
TL), CD 320, 731 m.
DISTINGUISHING FEATURES.— 1D II, 8–9; P i17–i18; GR-I (inner) 7–8 total; scales below mid-
base 1D 3.5–4.0, below 2D 4.5–5.5; pyl. caeca about 9. Snout long, sides converge in gentle curve
towards sharp spinous scute at tip, length 42–48% HL, anterolateral margin snout incompletely
supported by bone; orbit 23–27% HL; upper jaw 22–26% HL. Nasal fossa and underside of snout
scaly; body scales large, spinules stout, broad-based, in 1–6 parallel rows; scales of head mostly
with spinules arranged in single parallel ridge-row. Light organ short, scarcely visible before vent;
Group I of Iwamoto (1990). Color brown, to grayish; mouth and gill cavities dark; fins dusky to
blackish. Attains about 450 mm TL.
DISTRIBUTION.— Pacific coast of s. Japan to Taiwan, East China Sea, South China Sea, and
sw. Pacific. In Taiwan, specimens were collected from NET and SWT in depths of 441–850 m.
REMARKS.— Coelorinchus parallelus has a reportedly disjunct distribution in the western
Pacific. Its apparent absence from the Philippines and Indo-Australian Archipelago and its report-
ed presence on oceanic elevations in the sw. Pacific is suspicious. Specimens from the latter area
should be examined in greater detail to confirm their conspecificity with the type specimens.
Among the Taiwan Coelorinchus species with a simple (Group I) light organ, C. parallelus can be
distinguished by the combination of broad spinules on body scales arranged in 4–6 parallel rows
and its incompletely supported anterolateral snout margin. The two Taiwan specimens previously
reported as C. commutabilis by Shao et al. (2009) from the SCS are questionably recorded here as
this species. In almost all features, they fall in well with C. parallelus, but the spinules on their
body and head scales are more slender and lack the broadly transverse buttresses characteristic of
C. parallelus. Also, the nasal fossa is sparsely scaled anteriorly and the snout viewed dorsally is
somewhat broader and more convexly curved. That these SCS specimens are relatively small
(210+- 220+ mm TL) may account for the differences. A good size series is necessary to confirm
these differences.
Coelorinchus productus Gilbert and Hubbs, 1916
Coelorhynchus productus Gilbert and Hubbs, 1916:175–177, pl. 9, fig. 1 (holotype, USNM 76865; Suruga
Gulf, Japan, 197–297 fm [360–543 m]; paratypes, CAS-SU 22977, USNM 76872, 76873).
Coelorinchus productus: Yatou in Okamura and Kitajima, 1984:233, fig. 166 (p. 232), 369 (3 spec., 210–270
mm TL; Okinawa Trough, 410–600 m).— Iwamoto, 1990:179–180, fig. 406 (descr.).— Shao et al.,
2008: table 2 (3 spec., NET, SWT, 100–650 m).
Caelorinchus productus: Chiou et al., 2004b:44, fig.11 (2 spec., NET).
MATERIAL EXAMINED (5 spec.).— NET: ASIZP 61326 (1, 296+ TL), Da-xi; ASIZP 61327 (1,
221 TL), Da-xi; ASZIP 65573 (1, 333 TL), Da-xi; ASIZP 65644 (1, 239 TL), Da-xi; ASZIP 65648
(1, 241+ TL), Da-xi.
DISTINGUISHING FEATURES.— 1D II 8–10; P i16–i18; GR-I (inner) 6–8 total. Scales below
midbase 1D 3.5–4.5, below 2D 4–6; pyl.caeca about 27. Snout sharply pointed, length 39–42% HL,
1.5–1.7 times orbit; anterolateral margin of snout completely supported by bone; orbit 26–29% HL,
upper jaw 22–23% HL; nasal fossa scaled ventrally; underside of head naked except for occasion-
al small patch below preopercle angle. Light organ group II of Iwamoto (1990), fossa narrow and
short, not extending to pelvic-fin bases. Spinules on body scales daggerlike in 3–5 slightly diver-
gent ridgelike rows, middle row strongest. Ground color grayish-brown, silvery ventrally, bluish
over abdomen becoming blackish ventrally; mouth and gill cavity blackish; fins dusky to blackish.
Attains about 310 mm TL.
DISTRIBUTION.— Pacific coast of s. Japan to ne. Taiwan in 271–651 m.
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REMARKS.— Coelorinchus productus has been confused with the closely similar C. anati-
rostris, into which Okamura (1970:186) synonymized the species. However, Yatou (in Okamura
and Kitajima, 1984:233) treated it as distinct, and Iwamoto (1990:130) and Nakabo (2002:434)
provided characters by which the two species could be distinguished. Chiou et al. (2004b) used the
specimens here listed to document the first record of the species in Taiwanese waters. In a recent
paper, Fukui et al. (2010) re-synonymized C. productus with C. anatirostris based on examination
of two specimens of the former and one of the latter, each collected in Suruga Bay. Their selected
measurements and analysis of nucleotide sequences (16S rRNA) showed the two to be the same.
However, their measurements do not wholly agree with those we took from our specimens of the
two species. Aside from proportional measurements, which can sometimes be misleading because
of individual variation, they mention no other character used to distinguish their specimens of each
species. This raises the question of whether or not they actually had specimens of C. productus (or
C. anatirostris). Obviously, this problem needs further investigation; in the meantime, we will con-
tinue recognition of both species based on characters here given.
Coelorinchus sheni Chiou, Shao, and Iwamoto, 2004
Caelorinchus sheni Chiou, Shao and Iwamoto, 2004a:37–39, figs. 1–4 (holotype, ASIZP 061348; off Da-xi,
ne. Taiwan, 24°54ʹ63ʺN, 120°03ʹ49ʺE, 400–650 m; paratypes: ASIZP 061232 [1], CAS 215541[1]).
Coelorinchus sheni: Shao et al., 2008: table 2 (3 spec., NET, SWT, 100–650 m).
MATERIAL EXAMINED (13 spec.).— NET: ASIZP 61348 (1, holotype, 420 TL), Da-xi; CAS
215541 (1, paratype, 427 TL), Da-xi; ASIZP 70292 (1, 535 TL), Da-xi; ASIZP 70210 (8, 135–377
TL), Da-xi. BSKU 116417 (1, 578 TL), Da-xi. SET: ASIZP 61232 (1, paratype, 937 TL), Lyu-dao.
DISTINGUISHING FEATURES.— 1D II 8–9; P i18–i19; GR-I (inner) 9–10 total. Scales below
midbase 1D 4.5–5.0, below 2D 5–6; pyl.caeca about 27. Snout long, smoothly conical in lateral
profile, narrow in dorsal view, its length 41–56% HL, about 2.0 times orbit; anterolateral margin
of snout incompletely supported by bone; orbit 20–21% HL, its dorsal margin well below dorsal
profile; upper jaw 30–37% HL; nasal fossa scaled anteriorly and ventrally; underside of head fully
scaled. Light organ group II of Iwamoto (1990), fossa narrow and short, not extending to pelvic-
fin bases. Spinules on body scales sharp, blade-like, in 5–7 slightly divergent, crestlike rows, mid-
dle row strongest. Ground color light brown, bluish over abdomen; about 5 prominent saddle mark-
ing interspersed with pale narrow bars from trunk to tip of tail; mouth and gill cavity blackish; fins
dusky, but pelvic fins blackish except for white outer prolonged ray; margin of A blackish. Attains
more than 937 mm TL.
DISTRIBUTION.— Known only from ne. and se. Taiwan in depths of about 100–650 m.
REMARKS.— Coelorinchus sheni appears to be confined to rough-bottom slopes of e. Taiwan
where bottom trawls are ineffective; specimens were all captured by longline.
Coelorinchus smithi Gilbert and Hubbs, 1920
Coelorhynchus smithi Gilbert and Hubbs, 1920:493–498, fig. 20 (holotype, USNM 78212; Indonesia ne. of
Celebes, 298 fm [545 m]; 8 paratypes, Philippines and Indonesia).— Okamura, 1970:179–183, pl.
XXXIX, text-figs. 75–77 (63 spec., 144–322 mm TL; s. Japan in 300–610 m).— Okamura in Okamura et
al., 1982:171, 353, fig. 103 (p.170) (8 spec., 242.5–295 mm TL; Kyushu-Palau Ridge and Tosa Bay, in
300–610 m).
Coelorinchus smithi: Okamura in Masuda et al., 1984:98, fig. 83C (compiled).— Yatou in Okamura and Kita-
jima, 1984:231, 368, fig. 164 (p. 230) (5 spec., 235–385 mm TL; Okinawa Trough [East China Sea], in
400–600 m).— Shao et al., 2008: table 2 (8 spec., NET, SCS, SWT, 441–1110 m).
Caelorinchus smithi: Iwamoto and Williams, 1999:161–164, fig. 22 (10 spec., Java, n. and ne. Australia,
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 67

402–731 m).— Iwamoto and Graham, 2001:455, fig. 71 (1 spec., se. Australia, 740 m).— Nakabo,
2003:433 (compiled).— Chiou et al., 2004b:36, 47 (in key, list).
MATERIAL EXAMINED (7spec.).— NET: ASIZP 61330 (1, 316 TL), Da-xi; ASIZP 61331 (1,
291 TL), Da-xi. SWT: ASIZP 65523 (2, 250–265 TL), CD 138, 441 m; ASIZP 65594 (2, 230–255
TL), CD 141, 985–1110 m; ASIZP 58062 (1, 230+ TL), Tong-Sha Islands, SCS.
DISTINGUISHING FEATURES.—1D II 7–10; P i15–i19; GR-I (inner) 7–8 total; scales below mid-
base 1D 3.5–4.5, below 2D 4.5–6.0; pyl.caeca 19–37. Snout sharply pointed, length 39–52% HL;
anterolateral margin of snout completely supported by bone; orbit 24–30% HL, upper jaw 20–29%
HL; nasal fossa usually sparsely scaled ventrally; underside of head almost fully scaled (scales with
1–3 short rows of spinules). Light organ group II of Iwamoto (1990), fossa narrow and short, not
extending to pelvic-fin bases. Spinules on body scales blade-like in 3–7 sharp, divergent, ridgelike
rows. Ground color grayish-brown, paler to silvery ventrally, bluish over abdomen but not on chest;
mouth and gill cavity dark; most fins dark dusky to blackish. Attains more than 32 mm TL.
DISTRIBUTION.— Broadly distributed from s. Japan to Indonesia and Australia, in 300–750 m.
In Taiwan, specimens have been collected between 441 m and 1110 m.
REMARKS.— Among the Coelorinchus of Taiwan, C. smithi is most similar to C. leptorhinus,
C. spinifer, C. macrorhynchus, and C. japonicus in having the anterolateral snout margin complete-
ly supported by bone.It is distinguishable from C. leptorhinus inhaving underside of snout fully
scaled (cf. mostly naked) and from C. spinifer in having notably stronger, larger scale spinules on
body scales, arrayed in three divergent rows with the middle row notably larger, and by having the
length of the 1D base entering 1.2 times into the 1D-2D interspace. It differs from C. japonicus in
having a distinct, though short, naked fossa before the vent (lacking in C. japonicus), somewhat
fewer scale rows below the midbase of 1D (3.5–4.5 vs. 4.5–6.0) and below 2D origin (4.5–6.0 vs.
5.5–7.5), fewer pyloric caeca (21–37 vs. 41–60), and narrower scale spinules (broad buttresses in
C. japonicus). Differences from C. macrorhynchus are discussed in the description of that species.
Coelorinchus cf. spinifer [sensu Gilbert and Hubbs, 1920]
Coelorhynchus spinifer Gilbert and Hubbs, 1920:516–519, fig. 30 (holoype: USNM 78226, Gulf of Tomini,
Sulawesi [Celebes], 762 fm [1440 m]).
Coelorinchus spinifer: Shao et al., 2008: table 2 (1 spec., SCS, 1098 m; first record from Taiwan).— Iwamo-
to et al., 2009:47–48, fig. 4A–B (descr. of Taiwan spec.)
MATERIAL EXAMINED (1 spec.).— SCS: ASIZP 66748 (1, 52 HL, 180 TL), CD 322, 1098 m.
DISTINGUISHING FEATURES.— 1D II, 11; P i16; GR-I 2+7 (inner); GR-II 1+7 (outer); scales
below midbase 1D about 7, below 2D 6; lateral line scales over distance equal to predorsal length
ca. 43. Snout sharply pointed, about twice orbit diameter, 45–52% HL; anterolateral margin of
snout completely supported by bone; orbit 22–23% HL, internasal width 19%, interorbital 25%,
orbit to preopercle 30%, postorbital 30%, upper jaw 22%; length 1D base 1.2 times into 1D-2D
interspace; underside of head almost fully covered with tiny deciduous scales. Light organ group
II of Iwamoto (1990), fossa narrow and short, not extending to pelvic-fin bases. Scales strong and
prickly, spinules on body scales in 3 slightly divergent, ridgelike rows, middle row larger. Ground
color brownish, blackish over abdomen, opercles, jaws, and gular and branchiostegal membranes;
mouth and gill cavity black; most fins dusky, but P and V blackish, outer V ray pale. Attains at least
185 mm TL.
DISTRIBUTION.— Indonesia off Sulawesi and South China Sea off Taiwan in depths of 1440 m
and 1098 m, respectively.
REMARKS.— Coelorinchus spinifer is among the deepest-living members of the genus; only a
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handful of Coelorinchus species occur at depths greater than 1000 m. The Taiwan specimen is only
the second known of the species (Iwamoto et al. 2009). The notably long and strong spinules on
the scales, especially on the head ridges, in the holotype and Taiwan specimen, may reflect a juve-
nile condition. Until a larger size series is obtained, we are uncertain of its status within the genus.
It is otherwise similar in most characters to C. smithi.
Genus Coryphaenoides Gunnerus, 1874
DISTINGUISHING FEATURES.— BR 6. Spinous second ray of 1D serrated along leading edge.
Anus usually immediately before anal fin. No light organ.
REMARKS.— Coryphaenoides is the second largest genus of Macrouridae, with more than 60
species currently recognized, only four of which are so far known from Taiwan. Six or seven sub-
genera continue to be recognized, some as full genera, but the circumscription of each of these has
not been based on adequate phylogenetic analyses, although some attempts have been made using
very limited numbers of species (e.g., Wilson et al. 1991; Wilson 1994), Wilson and Attia (2003)
using DNA sequencing, peptide mapping of lactate dehydrogenase, and protein electrophoresis;
and Rao-Varón and Ortí (2009) using nuclear and mitochondrial DNA sequences. Most of the
species appear to have their primary depth range in mid-continental-slope depths, but many occur
at lower-slope depths; a few are primarily found on the continental rise at depths of 2000–4000 m,
and the deepest-living grenadier, C. yaquinae Iwamoto and Stein, 1974, has been captured in
abyssal depths below 6000 m. That species and C. armatus (Hector, 1874) can be expected in Tai-
wan waters deeper than about 4000 m, and other Coryphaenoides species are likely to be found
when depths greater than 2000 m are more thoroughly sampled. As might be expected from fishes
of great depths, many of the species are known from widely separated areas throughout the world
oceans. A few of the larger members of this genus are of some commercial importance. The Round-
nose grenadier (C. rupestris) of the North Atlantic has long been targeted by commercial fishermen
and some stocks have become severely depleted. The Pacific grenadier (Coryphaenoides acrolepis)
is the target of a very limited fishery off northern California.
Key to the Species of Coryphaenoides in Taiwan
1a. V rays 7 or 8; a greatly elongated spinous 1D ray, usually more than 1.5 times HL C. microps
1b. V rays 9–12; spinous 1D ray much less than 1.5 times HL . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Mouth large, upper jaw extends to below posterior margin of orbit or beyond; barbel 10–23%
of HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rudis
2b. Mouth moderate to small, upper jaw not extending beyond posterior 1⁄3of orbit; barbel 6–14%
of HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Interorbital space much smaller than orbit (about 1.3 into); GR-I (inner series) 9–10 total; snout
acutely pointed; preopercle ridge forming acute, lobelike angle posteroventrally. . C. nasutus
3b. Interorbital space much larger than orbit (about 0.7 into); GR-I (inner series) about 7 total;
snout rather bluntly pointed; preopercle ridge forming shallow rounded lobe posteroventrally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cf. asper
Coryphaenoides cf. asper [sensu Günther, 1877]
Coryphaenoides asper Günther, 1877:440 (holotype, BMNH 1887.12.7.88; se. of Cape Nojima [se. of Boso
Peninsula], Japan, 34°37ʹN, 140°32ʹE, 1875 fm [3429 m]). Shao et al., 2008: table 2 (1 spec., Taiwan
[SCS], 1982 m; first record from Taiwan).— Iwamoto et al., 2009:45–47, fig. 3 (descr.; Taiwan [SCS]
spec.).
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 69

MATERIAL EXAMINED (1 spec.).— SCS: ASIZP 66107 (1, 92.9 HL, 435+ TL), CD 325,
1982 m.
DISTINGUISHING FEATURES (figures in square brackets are from original description).— 1D II
[9] 11, P i19–i21 [25], V 10 [11], GR-I (inner) 2+5; GR-II outer 1+6/1+8; scales below 1D origin
7.5 [6], below midbase 1D 5, below 2D origin 5.5, over distance equal to pre-1D 37. Snout 29%
HL, broadly pointed in lateral view, preoral length short, 16%, orbit small, 19%, about 1.5 into
snout length, interorbital width 28%, orbit to preopercle 48%, upper jaw 33%, barbel 14% (about
1.3 times in orbit diameter), greatest body depth 81%, 1D-2D interspace 21%, height 1D 91%,
length 1D base 26%, length V 83%. Second spine of 1D serrated, produced beyond segmented
rays, outer V ray prolonged. Upper jaw extends posteriorly to below middle of orbit. Head fully
scaled except over gular and branchiostegal membranes; modified, thickened scales above subor-
bital ridge; body scales covered with slender spinules in about 5 divergent rows.
DISTRIBUTION.— Known only from two specimens, the holotype taken off Japan in 3429 m
and the Taiwan specimen taken in the South China Sea in 1982 m.
REMARKS.— The ASIZP specimen was first recorded by Shao et al. (2008) and subsequently
described in more detail by Iwamoto et al. (2009). If correctly identified, it represents only the sec-
ond known specimen of the species. The absence of previous captures since the H.M.S. Challenger
made its historic voyage in 1873–1876 is probably owed to the paucity of trawl hauls made at
depths exceeding 2000 m in the area. A few character differences that we found between this Tai-
wan specimen and the holotype have led to some uncertainty as to it identity; it is for this reason
that we consider the identification as tentative.
Coryphaenoides microps (Smith and Radcliffe, 1912)
Macrourus microps Smith and Radcliffe in Radcliffe, 1912:116–117, pl. 25, fig. 3 (holotype USNM 72934,
42.5 cm long; Philippines, Lagonoy Gulf, se. Luzon, 13°37ʹ30ʺN, 123°41ʹ09ʺE; Albatross sta. 5511, 560
fm [1024 m]; 4 paratypes, sta. 5325 off n. Luzon, [CAS-SU 2544])
Coryphaenoides microps: Gilbert and Hubbs, 1920:418–419 (descr.; holotype and 4 paratypes, CAS-SU 2246,
n. Luzon, Albatross sta. 5325, 224 fm [410 m]).— Shcherbachev and Iwamoto, 1995:300–301.— Chiou et
al., 2004a: table 1 (listed from Taiwan).— Shao et al., 2008: table 2 (1 spec., SWT, 687 m).— Kim et al.,
2009:108–110, fig. 3 (descr.; type spec., plus others from Philippines, Taiwan [NET], and Korea [East Sea,
115 m; first record])
MATERIAL EXAMINED (53 spec.).— NET: ASIZP 58265 (1, 250+ TL, 50.8 HL), Da-xi; ASZIP
60252 (1, 137 TL), Da-xi; ASZIP 61128 (4, 267+-330 TL), Da-xi; ASIZP 63838 (2, 306–413 TL),
CP 195, 570 m; ASIZP 65566 (1, 93+ TL), CP 124, 1129–1165 m; ASZIP 65657 (1, 233 TL), Da-
xi; ASIZP 66900 (1, 140 TL), CP 247, 480 m. SWT: ASIZP 63767 (1, 232+ TL), CD 194, 507 m;
ASIZP 63792 (1, 365+ TL) and ASIZP 64144 (1, 268+ TL), CD 193, 821 m; ASIZP 65516 (2,
160–190 TL) and CAS 224886 (1, 175 TL, ex. ASIZP 65516), CD 142, 227–235 m; ASIZP 65526
(2, 150–200 TL), CD 138, 441 m; ASIZP 65535 (4, 130–210 TL), CD 137, 316–477 m; ASIZP
65581 (1, 295 TL), CD 229, 880–1062 m; ASIZP 65591 (8, 160–200 TL), CD 136, 998–1211 m;
ASIZP 65603 (1, 359 TL), CD 133, 690–748 m; ASIZP 65617 (1, 174 TL), CD 137, 316–477 m;
ASIZP 65624 (1, 395 TL), CD 229, 880–1062 m; ASIZP 65639 (1, 385+ TL), CD 229, 880–1062
m; ASIZP 66099 (1, 295 TL), OCP 301, 687 m; ASIZP 66109 (1, 320+ TL), OCP 301, 687 m;
ASIZP 66111 (1, 463+ TL), OCP 301, 687 m; ASIZP 66413 (1, 220+ TL), Dong-gang; ASIZP
66425 (1, 214+ TL), Dong-gang; ASIZP 66427 (1, 166 TL), Dong-gang;ASIZP 66786 (1, 310 TL),
OCP 301, 687 m; ASIZP 66799 (1, 188+ TL), OCP 302, 695 m. No data: ASIZP 65579 (1, 421
TL). Other material (Univ. Philippines, Marine Science Institute, Manila [UPMSI]): UPMSI
uncat. (1, 380 TL) (no data, probably MUSORSTOM II); UPMSI uncat. (5) MUSORSTOM II sta.
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79, off Luzon, 13°31.6ʹN, 120°33.7ʹE, 326–240 m; 1 Dec 1980, 210+–252 TL). Other specimens:
USNM 72933 (holotype, 82.0 HL, 413+ TL), e. coast Luzon, Philippines, 1024 m; CAS-SU 25446
(2 paratypes, 23.7–39.9 HL, 124+–187+ TL), n. Luzon, 18°34’15ʺN, 121°51’15ʺE, 410 m.
DISTINGUISHING FEATURES.— 1D II 9–10; P i19–i21; V 7–8; GR-I (inner) 9–11 total; GR-II
outer/inner 8–10/7–9; scales below midbase 1D 4.5–7.5, below 2D origin 6.0–9.5, over distance
equal to snout tip-to-1D 29–35. Snout 28–33% HL, extending less than pupil diameter beyond
mouth; preoral steep, short, length 11–16%; internasal narrow, width 15–21%; interorbital
18–26%; orbit 17–25%; postorbital length 47–53%; distance orbit to preopercle 40–47%; upper
jaw 30–33%; barbel short, thick 8–14%; greatest body depth 61–96%; 1D-2D interspace 16–38%;
second spinous ray of 1D greatly prolonged, 178–388% HL, serrations sparse, reduced or lost in
large adults; length V 43–87% HL, outer ray moderately prolonged. Upper jaw extends posterior-
ly to below anterior 1⁄3of orbit, mouth opening restricted laterally. Free margin of preopercle broad-
ly rounded. Head scaly except narrow naked margin under snout and suborbital and all of gular and
branchiostegal membranes; a tubercular scale at snout tip; body scales covered with slender
reclined spinules in numerous subparallel rows. Color dark brown in adults, paler in young; fins
generally pale near base, blackish distally.
DISTRIBUTION.— Off Luzon, Philippine Islands, Japan Sea off Korea, and Taiwan (SCS,
NET), in 240–1024 m, but one exceptional capture at 115 m off Korea.
REMARKS.— Chiou et al. (2004b:47, table 1) listed C. microps from Taiwan, indirectly sug-
gesting that the C. marginatus recorded by Shen et al. (1993) was an incorrect identification. Kim
et al. (2009) recorded the species from the East (Japan) Sea off Korea and provided additional
information on the species, comparing it with C. marginatus. Coryphaenoides microps is closely
similar to C. marginatus, and the two species have many features in common. They can be sepa-
rated by a combination of characters including orbit diameter (25–33% HL in C. marginatus cf.
17–25% in C. microps), postorbital length (42–45% HL cf. 47–56%), distance orbit to preopercle
(33–40% cf. 40–47%), and length spinous 1D ray (1.5–2.5 times HL cf 1.8–3.0). The two species
belong to what Gilbert and Hubbs (1920:413) describe as “a rather well-marked group of species
which agree in possessing a produced dorsal spine, a deep and sharply compressed body, and a dor-
sal contour horizontal behind the first dorsal fin.” Other species of this group include C. semiscaber
Gilbert and Hubbs, 1920, C. macrolophus (Alcock, 1889), and C. tydemani Gilbert and Hubbs
1920.
Coryphaenoides nasutus Günther, 1877
Figures 10A–B.
Coryphaenoides nasutus Günther, 1877:440 (2 syntypes, BMNH 1887.12.7.78–79; 1 syntype?, MNHN
1890–0123; “south of Yedo” [Tokyo], Japan, 34°07ʹN, 138°00ʹE, Challenger sta. 235, 565 fm [1033
m]).— Gilbert and Hubbs, 1916:168 (45 spec., Japan: Hokkaido to e. coast Kyushu; 13 Albatross sta.,
250–614 fm [457–1123 m]).— Okamura, 1970:140–143, pl. XXXI, text-fig. 56 (38 spec., 155–470 mm
TL; Hokkaido to East China Sea, 625–1180 m).— Okamura in Okamura et al., 1982:165, 351, fig. 99 (8
spec., 136–234 mm TL; Kyushu-Palau Ridge).— Sawada in Amaoka et al. 1983:113, 195, fig. 64 (5 spec.,
199–391 mm TL, 815–1100 m).— Okamura in Masuda et al., 1984:96, pl. 82F (compiled).— Yatou in
Okamura and Kitajima, 1984:219, 364, fig. 155 (5 spec., 265–470 TL; Kyushu-Palau Ridge, 815–1000
m).— Iwamoto, 1990:216 (descr.).— Shao et al., 2008: table 2 (5 spec., NET, SET, 979–1268 m; first
record from Taiwan).
Macrurus nasutus: Günther, 1887:132, pl. 30, fig. B (descr.; type illustrated).
MATERIAL EXAMINED (9 spec.).— NET: ASIZP 64113 (2, 240+-338+ TL), CP 242; ASIZP
65576 (1, 288+ TL), CP 197. SET: ASIZP 65511 (1, 425 TL), CP 127. SCS: ASIZP 65622 (1, 415
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 71

TL), CD 226. Non-types and possible types: BMNH 1887.12.7.80 [1], BMNH 1889.6.30.8–9 [2],
NMI block reg. no. 114.1899 [1].
DISTINGUISHING FEATURES.— 1D II 9–11; P i18–i23; V 9–10; GR-I (inner) 1–2+7–9, GR-II
outer/inner 1–2+8/1–2+7–8; scales below midbase 1D 4.5–5.5, below 2D origin 7.0–7.5, over dis-
tance equal to pre-1D len. 36–40. Snout length 27–32% HL; orbit 24–30%; interorbital width
20–24%; upper jaw 29–36%; barbel 5–8%; greatest body depth 75–97%; interdorsal [1D-2D]
space variable 90–114% [2D rays rudimentary anteriorly]; length V 49–69%. Snout acutely point-
ed, extending short distance in front of mouth. Second spinous ray of 1D serrated, slightly pro-
duced (95–125% HL); outer V ray slightly produced. Upper jaw extends posteriorly to below mid-
dle of orbit. Head almost fully scaled except for gular and BR membranes; scales above suborbital
ridge thickened and modified; a stout conical tubercle at snout tip; body scales covered with need-
like spinules in convergent rows.
DISTRIBUTION.— Pacific coast of Japan from Hokkaido to East China Sea, and Taiwan from
e. coast to South China Sea. Published depth range 625–1180 m, but in Taiwan, specimens collect-
ed in 979–1268 m.
REMARKS.— Shao et al. (2008) first recorded C. nasutus from Taiwan and the South China
Sea based on the current specimens. This species is similar in many features to C. microps and
C. marginatus but has a larger, broader head, more strongly pointed snout armed with a stouter ter-
minal scute, V rays 9 or 10 (compared with 7 or 8), and an acute preopercular ridge and angular
preopercular margin.
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A
B
FIGURE 10. Coyphaenoides nasutus Gunther, 1877. A. ASIZP 64113, 338+ mm TL., fresh. B. ASIZP 65576, 288+ mm
TL, preserved.

Coryphaenoides rudis Günther, 1878
Figures 11A–C.
Coryphaenoides rudis Günther, 1878:24 (lectotype BMNH 1889.12.7.74; Pacific, n of Kermadec I., Chal-
lenger sta. 171, 600 fm [1097 m]. BMNH 1887.12.7.75–77 (paralectotypes: 3).— Shcherbachev and
Iwamoto, 1995:301 (Indian Ocean).— Iwamoto and Williams, 1999:170 (4 spec., nw. and se. Australia,
1120–1700 m).— Merrett and Iwamoto, 2000:754 (3 spec., New Caledonia region, 1315–1862 m).—
Iwamoto and Graham, 2001:467 (descr., 2 spec., NSW, Australia, 1050–1150m).— Shao et al., 2008: table
2 (1 spec., SCS, 1982 m; first record from Taiwan).
Macrourus paradoxus Smith and Radcliffe in Radcliffe, 1912:115–116, pl. 25, fig. 1 (holotype, USNM 72932,
585 mm TL, e. Palawan, Philippines, 9°13ʹ00ʺN, 118°51ʹ15ʺE, 1105 fm [2021 m]).
Nematonurus macrocephalus Maul, 1951:17–22, figs. 3, 4c (holotype, MMF [mounted spec.], 210 HL, taken
off Madeira).
MATERIAL EXAMINED (1 spec.).— SCS: ASIZP 66117 (1, 132.6 HL), CD 325, 1982 m. SET:
NMMB-P uncat. (1, 1240 TL), Taitung.
DISTINGUISHING FEATURES.— 1D II,9–11, P i19, V 9–11, inner GR-I about 10, outer GR-II
8–9 total, scales below 2D 6–7. (Proportional measurements of Taiwan specimen in square brack-
et): snout length [27] 23–29% HL; preoral [17]10–12%; orbit [20] 16–26%; interorbital [27]
26–30%; upper jaw [38] 37–43%; barbel [20]10–23%; greatest body depth 80–100%; height 1D
[76] 43–73%; length V [62] 50–111%. Snout low, bluntly rounded (in adults) to bluntly pointed
(juveniles), scarcely extending beyond mouth in adults; no stout angular ridges on head; suborbital
region vertical. Second 1D spine serrated (serrations reduced in largest specimens), slightly pro-
duced; outer V ray moderately produced. Upper jaw extends to below hind margin of orbit. Pre-
maxillary teeth in narrow tapered band, outer series enlarged; mandibular teeth in about 3 series
tapering to one. Head fully scaled except for gular and branchiostegal membranes; modified, thick-
ened scales on suborbital and small tubercles at snout tip and at lateral angles; body scales dense-
ly covered with small conical spinules in irregularly divergent rows. (Measurement and count data
partly from Shcherbachev and Iwamoto [1995], and Iwamoto and Williams [1999])
DISTRIBUTION.— Possibly worldwide in warm seas; depth range 600–3500 m, but usually
1000–2000 m. One of Taiwanese specimens collected in the South China Sea in 1982 m and anoth-
er one was taken from off Taitung as depth not less than 400 m.
REMARKS.— Our specimen is the first collected in Taiwan waters (recorded by Shao et al.
2008: table 2) and one additional specimen was collected from SW Taiwan off Taitung. Its pres-
ence is not surprising in that the holotype of C. paradoxus, a synonym, was captured off e. Luzon
Island in the Philippines.
Genus Hymenocephalus Giglioli, 1882
DISTINGUISHING FEATURES.— Snout high, relatively rounded, median nasal process forming a
weak snout tip (no horizontal platelike process mesially); paired nasal bone in broad contact along
median line, without wide gap around nostril cartilage; head much deeper than wide; body relative-
ly compressed; head mucous canals greatly expanded; head covering membranous and often trans-
parent; light organ long, small lens on chest anterior to pelvic-fin bases connected by a black streak
to round posterior lens immediately before anus; ventral striae well developed; fine black lines on
gular membrane oriented perpendicular to median line, not netlike; inner GR-I, lower limb 12–16;
spinous ray of 1D completely smooth; chin barbel present or absent.
REMARKS.— A genus of about 20 species, four of which are known from Taiwan. Most
species do not exceed about 20 cm TL. The highly developed luminescent organ system consisting
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 73

of a network of fine striations (“ventral striae”) on the cleithrum, chest, and belly regions is seen
in gadiforms only in this genus, its closely related genera Hymenogadus and Spicomacrurus, and
the monotypic genera Steindachneria Goode and Bean in Agassiz, 1888 and Lepidorhynchus
Richardson, 1846.The presence of two lens-like bodies of the light organ, one on the chest, the
other immediately before the anus, is unique among the grenadiers to Hymenocephalus,
Hymenogadus, and Spicomacrurus. The position of the lenses is similar to that of the ventral fos-
sae of some species of Coelorinchus, but so far as we can determine, there are no comparable struc-
tures in that or other grenadiers. The delicate, paper-thin head bones and membranous head integu-
ment are usually damaged during capture, adding to the difficulties of identifying specimens.
74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 11. Coryphaenoides rudis Günther, 1878. A. ASIZP 66117, 133 mm HL, fresh. B. NMMB-P uncat., 1240 mm
TL, fresh and C. same specimen, dorsal view of head.
A
B
C

Key to the Species of Hymenocephalus in Taiwan
1a. V 7–9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. V 11 or 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Barbel long, about 50% HL; orbit diameter 27–31% HL. . . . . . . . . . . . . . . . . . . . H. longiceps
2b. Barbel about 11–18% HL; orbit diameter 34–45% HL . . . . . . . . H. striatissimus striatissimus
3a. Barbel absent; orbit diameter 28–35% HL; color grayish brown in preservative, somewhat sil-
very in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lethonemus
3b. Barbel rudimentary, about 4% HL; orbit diameter 23% HL; color mostly blackish . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. papyraceus
Hymenocephalus lethonemus Jordan and Gilbert 1904
Hymenocephalus lethonemus Jordan and Gilbert, 1904:615, text-fig. (holotype, USNM 50936; Sagami Bay,
Japan, 120–265 fm [219–485 m]; paratypes, CAS-SU 8641 [3 spec.], USNM 51455 [3 spec.]).— Gilbert
and Hubbs, 1916:188–189 (22 spec., s. Japan, East China Sea, 197–440 fm [360–805 m]).— Okamura,
1970:54–56, pl. II, text-fig. 23 (83 spec., 85–140 mm TL; e. coast of s. Japan).— Yatou in Okamura et al.,
1982:143, 347, fig. 87 (p. 142) (7 spec., 175.5–233 mm TL; Kyushu-Palau Ridge).— Okamura in Masu-
da et al., 1984:94, pl. 80–G (compiled).— Okamura in Okamura and Kitajima, 1984:201, 358, fig. 142
(p. 200) (8 spec., 148–180 mm TL; East China Sea [Okinawa Trough]; 570–810 m).— Chiou et al.,
2004b:44, fig. 12 (13 spec., ne. Taiwan).— Shao et al., 2008: table 2 (21 spec., Taiwan [NET, SWT, SCS],
441–1040 m).
MATERIAL EXAMINED (36 spec.).— NET: ASIZP 61231 (1, 126 TL), Da-xi; ASZIP 64270 (2,
105+-108+ TL), CP 234, 547 m; ASZIP 66974 (12, 112–124 TL), Da-xi. SWT: ASIZP 65561 (1,
120+ TL) and ASIZP 65611 (1, 115 TL), CD 203, 634–866 m; ASIZP 65598 (3, 120–155 TL), CD
138, 441 m; ASIZP 65604 (1, 133 TL) and ASIZP 66946 (1, 330 TL), CD 134, 736–1040 m;
ASIZP 66731 (1, 97 TL), OCP 317, 515 m; ASIZP 66871 (5, 43–68 TL), OCP 312, 517 m; ASIZP
66872 (1, 78 TL), CD 311, 516 m; ASIZP 66873 (4, 50–80+ TL), OCP 313, 516 m; ASIZP 67582
(2, 75+-95+ TL), CP 348, 395 m. Other specimens: ASIZP 67976 (1, 151 TL), Aurora.
DISTINGUISHING FEATURES.— 1D II 9–11; P i13–i16; V 11 (rarely12); inner GR-I 21–25 total;
pyl. caeca 11–16. Head about 5–6 in TL; body depth about 8.0–9.5 TL. Snout high, pointed; mouth
large, upper jaw extending posteriorly to below hind margin of orbit; barbel absent; gill openings
wide, gill membrane free over isthmus; outer gill slit about as wide as orbit. Attains about 180 mm
TL. (Compiled from Okamura [1970] and Gilbert and Hubbs [1916]).
DISTRIBUTION.— Pacific coast of s. Japan, East China Sea off Kyushu Is., Japan, and Taiwan
(NET, SWT, SCS), in 360–1040 m.
REMARKS.— Okamura (1970:56) noted that among members of Hymenocephalus, this is the
deepest-occurring in Japan.
Hymenocephalus longiceps Smith and Radcliffe, 1912
Hymenocephalus longiceps Smith and Radcliffe in Radcliffe, 1912:111–112, pl. 23, fig. 3 (holotype, USNM
72928; off se. Luzon, Philippines, 201 fm [368 m]; paratypes, USNM 149297).— Yatou in Okamura et al.,
1982:141, 346, fig. 85 (p. 140) (5 spec., 135–225.5 TL; Kyushu-Palau Ridge, 420–555 m).— Okamura in
Masuda et al., 1984:93, pl. 80–D (compiled).— Okamura in Okamura and Kitajima, 1984:199, 357, fig.
140 (p. 198) (5 spec., 184–230 mm TL; East China Sea [Okinawa Trough], 353–490 m).— Shen et al.,
1993:171 (descr.).— Chiou et al., 2004b:37, 47 (in key, list).— Shao et al., 2008: table 2 (41 spec., Taiwan
[NET, SWT, SCS], 227–1211 m).
Hymenocephalus (Hymenocephalus) longiceps: Gilbert and Hubbs, 1920:525–526 (63 spec., South China Sea
off Hong Kong and Taiwan, Philippines, “East Indies,” 107–298 fm [196–545 m]).— Okamura, 1970:
45–47, pl. XIV (9 spec., 156–240 m TL; Pacific coast s. Japan, 300–500 m).
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 75

MATERIAL EXAMINED (47 spec.).— NET: ASIZP 61235 (1, 195 TL), Da-xi; ASIZP 61236 (6,
96–152 TL), Da-xi; ASIZP 66373 (1, 160 TL), Nan-fang-ao. SCS: ASIZP 65546 (1, 160+ TL), CD
137, 316–477 m; ASIZP 65554 (1, 200 TL) and ASIZP 65563 (1, 160 TL), CD 142, 227–335 m;
ASIZP 65589 (5, 125–155 TL) and ASIZP 65606 (4, 125+-170 TL), CD 136, 998–1211 m. SWT:
ASIZP 60321 (1, 128 TL), Dong-gang; ASZIP 62270 (2, 100+-130+ TL), Dong-gang. SCS: ASIZP
66269 (1, 170+ TL), ASIZP 66797 (13, 87+-118+ TL), and ASIZP 66879 (2, 63+-101+ TL), CD
311, 516 m; ASIZP 66739 (1, 100+ TL), CP 315, 509 m; ASIZP 66836 (1, 26.1 HL, 110+ TL) and
ASIZP 66867 (1, 88 + TL), CP 314, 509 m; ASIZP 66837 (1, 168+-177+TL), CD 311, 516 m.
Other specimens: ASIZP 67841 (1, 200 TL), Aurora, 358–342 m; ASIZP 67949 (1, 90 TL), Auro-
ra, 269–277 m; ASIZP 68031 (1, 120 TL), Aurora, 500–524 m; ASIZP 68192 (1, 166 TL), Auroa,
442–431.
DISTINGUISHING FEATURES.— 1D II 9–10; P i13–i17; V 8; inner GR-I 18–23 total; pyl. caeca
18–22. Orbit 27–31% HL; upper jaw 53–59%. Head rather low, its length about 5–6.5 in TL; body
depth about 7–8 in TL. Snout low, broadly rounded, not projecting beyond mouth; barbel long,
more than orbit, about 2 in HL. Attains about 240 mm TL. (Compiled from Okamura, 1970 and
Gilbert and Hubbs, 1916).
DISTRIBUTION.— Known from s. Japan to South China Sea, Philippines, Sulu Sea, Bohol Sea,
and Celebes Sea, in 196–555 m. The species was taken in 227–1211 m depth off Taiwan.
REMARKS.— The long barbel and low, rounded snout distinguish this species from all others
of the subgenus from Taiwan. It is closely similar to H. longibarbis Günther, 1887 and the two may
be synonymous.
Hymenocephalus papyraceus Jordan and Gilbert, 1904
Figure 12.
Hymenocephalus papyraceus Jordan and Gilbert, 1904:614. fig. (holotype USNM 50935, 147 mm TL; Saga-
mi Bay, Japan, 120–265 fm [219–485 m], Albatross sta. 3697).— Okamura, 1970:56–58, pl. XVII, text-
fig. 24 (1 spec., 95 mm TL, Pacific coast s. Japan).— Sazonov, 1993:117–121 (fig.) (1 spec., East China
Sea; 800–826 m).
Hymenocephalus (Papyrocephalus) papyraceus: Gilbert and Hubbs, 1920:539 (included as one of three
species in new subgenus).
Hymenocephalus papiraceus: Okamura in Masuda et al., 1984:94, pl. 344–I (species name misspelled; com-
piled).
MATERIAL EXAMINED.— NET: NMMB-P9121 (1, 25.7 HL), Da-xi.
DISTINGUISHING FEATURES.— 1D II,10; P i14; V 11; GR-I (outer/inner) 0+16/5+17, GR-II
5+17/4+16. Orbit 23% HL, 1.4 into postorbital; interorbital 35%; internasal width 23%; suborbital
19%; postorbital 54%; orbit to preopercle angle 54%; upper jaw 47%; barbel 3.5%; 1D-2D inter-
space 60%; P length 70%; V length 62%; length outer V to A origin 54%; pre-anal length 144%;
length outer gill slit 27%. Head relatively deep and broad; orbit not included in dorsal profile; nape
high, beginning slightly behind posterior edge of orbit. Snout with high median nasal ridge, flexi-
ble, pointed tip projecting slightly beyond mouth. Posterior end of preopercle ridge narrowly chis-
el-shaped (or flathead shovel-shaped). Barbel small but distinct. Head color mostly black, but dor-
sal head bones and snout transparent. Entire trunk to over A origin deeply and heavily peppered;
remainder of body covered with large melanophores. Ground color medium brown; A base black
anteriorly. Gular and BR membranes black. Attains about 150 mm TL
REMARKS.— Hymenocephalus papyraceus is apparently a rare species, having been recorded
only three times previously: the holotype (and only type specimen), a small specimen in poor con-
dition reported by Okamura (1970), and the East China Sea specimen of Sazonov (1993). The Tai-
wan specimen is the first record from outside Japanese waters.
76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3

Hymenocephalus striatissimus striatissimus Jordan and Gilbert, 1904
Hymenocephalus striatissimus Jordan and Gilbert, 1904:612–613, text-fig. p. 613 (holotype, USNM 50934.
Suruga Bay, Japan, 167 fm [305 m]; paratype, CAS-SU 8549 [1]).— Gilbert and Hubbs, 1916:187–188
(22 spec., Pacific coast s. Japan and East China Sea; 360–543 m).— Yatou in Okamura et al., 1982:143,
346, fig. 86 (p. 142) (7 spec., 101–194.5 mm TL; East China Sea [Okinawa Trough], 300–555 m).— Oka-
mura in Masuda et al., 1984:93, pl. 80–E (compiled).— Okamura in Okamura and Kitajima, 1984:199,
358, fig. 141 (p. 198) (25 spec., 148–180 mm TL; Kyushu-Palau Ridge, 425–570 m).— Shen et al.,
1993:171 (descr.).— Chiou et al., 2004b:37, 47 (in key, list).
Hymenocephalus striatissimus striatissimus: Gilbert and Hubbs, 1920:529–530 (32 spec., South China Sea off
Hong Kong, Taiwan, and Luzon, Philippines; 380–494 m; 3 subspecies recognized, including one new
[H. s. aeger]).— Okamura, 1970:48–50, pl. XV (90 spec., 115–200 mm TL; Pacific coast s. Japan and East
China Sea off Kagoshima Prefecture; 300–540 m; new subspecies described [H. s. hachijoensis]).— Shao
et al., 2008: table 2 (34 spec., Taiwan [NET, ET, SCS], 100–1188 m).
MATERIAL EXAMINED (42 spec.).— NET: ASIZP 57973 (1, 206 TL), Nan-fang-ao; ASIZP
61233 (1, 184 TL), Nan-fang-ao; ASIZP 61234 (13, 112–162 TL), Da-xi; ASIZP 64247 (1, 110+
TL), CP 234, 547 m; ASIZP 65564 (1, 125 TL), CP 124, 1129–1165 m; ASIZP 65635 (1, 207 TL),
CD 209, 508–522 m; ASIZP 70731 (1, 115 TL), Da-xi. ET: ASIZP 65524 (8, 145–200 TL), ASIZP
65528 (2, 160–170 TL), ASIZP 65536 (1, 161 TL), ASIZP 65600 (1, 162+ TL), and ASIZP 65619
(1, 172 TL), CD 210, 445–1185 m; ASIZP 65553 (1, 93+ TL), CD 199, 1134–1188 m. SCS: ASIZP
65677 (1, 140 TL), CD 311, 516 m; ASIZP 66834 (1, 117 TL), CP 314, 506 m. Other locality:
ASIZP 67873 (1, 142 TL), Aurora, 506–542 m; ASIZP 67972 (1, 90 TL), Aurora, 507–540 m;
ASIZP 68189 (1, 117 TL), Aurora, 431–442 m; ASIZP 68414 (4, 70–110 TL), Aurora, 507–540 m.
DISTINGUISHING FEATURES.— 1D II 8–10; P i11–i16; V 8 (rarely 7 or 9); inner GR-I 16–22
total; pyl. caeca 10–17. Orbit circular, large 34–45% HL, 0.9–1.0 into postorbital; upper jaw
48–59%. Head deep, length about 6–7 in TL; body depth about 8–9 in TL. Snout bluntly rounded
but with short terminal point, barely projecting beyond mouth. Upper jaw extends to vertical
through posterior margin of orbit. Barbel about 1⁄3orbit. Attains about 200 mm TL.
DISTRIBUTION.— South China Sea off Hong Kong, Taiwan, and Luzon, Philippines, and ne.
Taiwan, in 445–1188 m, but mostly between about 300 and 550 m.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 77
FIGURE 12. Hymenocephalus papyraceus Jordan and Gilbert, 1904. NMMB-P9121, 25.7 mm HL, preserved, photo
reserved.

REMARKS.— Gilbert and Hubbs (1920) recognized three subspecies of this species: H. striatis-
simus striatissimus, H. striatissimus aeger, and H. striatissimus torvus. The last was originally
described as a full species by Smith and Radcliffe (in Radcliffe 1912). Okamura (1970:50–54)
described H. s. hachijoensis from two specimens taken off Hachijo, a group of remote islands about
180 nautical miles s. of Tokyo; Okamura in Masuda et al. (1984:93) elevated the taxon to full
species. Sazonov (1994:101–102) later recorded two additional specimens from the Northwest
Pacific Ridge (Emperor seamounts) and one from the Kyushu-Palau Ridge. It is distinguished from
H. s. striatissimus by “the longer barbel, the larger head, the smaller eye, and the lower snout”
(Okamura 1970:53).
Genus Hymenogadus Gilbert and Hubbs, 1920
DISTINGUISHING FEATURES.— Snout low, relatively pointed and protruding; paired nasal bones
in broad contact along median line, without gap around nostril cartilage; head about as broad as
high; body and head somewhat cylindrical; head mucous canals moderately developed; head cov-
ering mostly transparent; light organ long, small lens on chest anterior to pelvic-fin bases connect-
ed by a black streak to round posterior lens immediately before anus; ventral striae well developed;
fine black lines on gular membrane oriented perpendicular to median line, not netlike; inner GR-I,
lower limb 10–16; spinous ray of 1D weakly serrated; chin barbel present.
REMARKS.— Aside from the low, cylindrical body and head, pointed snout, fewer gill rakers,
and the presence of serrations along the leading edge of the first dorsal fin, this genus is otherwise
scarcely distinguishable from Hymenocephalus, in which it was originally included as a subgenus.
Aside from the widely distributed H. gracilis, the genus contains only H. tenuis, which is known
only from the single original record off Hawaii.
Hymenogadus gracilis Gilbert and Hubbs, 1920
Hymenocephalus (Hymenogadus) gracilis Gilbert and Hubbs, 1920:522–525, fig. 31 (holotype, USNM
78227, 96 TL, off s. Luzon, Philippines, 13 28ʹ45ʺN, 121 01ʹ12ʺE, 162 fm [296 m]).— Marshall and
Iwamoto in Marshall, 1973:602–604, fig. 31 (19 spec., nw. Pacific, e. and w. Atlantic; 342–618 m).—
Iwamoto and Merrett, 1997:518, fig. 20b.— Shao et al., 2008: table 2 (3 spec., Taiwan [NET, SCS],
100–950 m)
Hymenogadus gracilis: Okamura, 1970:61–63, pl. XVIII, text-fig. 27 (60 spec., 91–128 mm TL; s. Japan,
300–500 m).— Okamura in Masuda et al., 1984:94, pl. 80H (compiled).— Okamura in Okamura and Kita-
jima, 1984:201, 359, fig. 143 (p. 200) (3 spec., Okinawa Trough [East China Sea], 295–385 m).— Chiou
et al., 2004b:44–45, fig. 13 (2 spec., ne. Taiwan).
MATERIAL EXAMINED (6 spec.).— NET: ASIZP 61229 (1, 118 TL), Nan-fang-ao; ASIZP
61230 (1, 120 TL), Da-xi; ASIZP 65565 (1, 60 TL), Da-xi; ASIZP 70278 (1, 105 TL), Da-xi. Other
specimens: ASIZP 68334 (2, 96–112 TL), Aurora, 357–367 m.
DISTINGUISHING FEATURES.— 1D II9–11; P i14–i17; V 7–9, usually 8; inner GR-I 14–18 total.
Body depth 50–60% HL, about 9–10 in TL; barbel long, 20–30% HL. Attains about 130 mm TL.
DISTRIBUTION.— Probably circumglobal in warm seas, but not central e. Pacific, central e.
Atlantic, and central Pacific (but see Remarks), in 300–450 m.
REMARKS.— The taxonomic status of the Hawaiian species H. tenuis Gilbert and Hubbs, 1917
has yet to be resolved. It may be a synonym of H. gracilis. Okamura (1970:58) recognized
Hymenogadus as a full genus with two included species, H. gracilis and H. kuronumai, and by
implication, also H. tenuis. Okamura also placed H. kuronumai into its own subgenus Spicomacru-
rus Okamura, 1970, a taxon that Iwamoto et al. (2011) elevated to full generic status.
Hymenogadus gracilis appears to be more characteristic of oceanic elevations, such as islands,
78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3

seamounts, and ridges, than to continental landmasses, and the terete body form is suggestive of a
more pelagic, active-swimming lifestyle (not benthic, as proposed by Okamura 1970:60). Surpris-
ingly, the species was not recorded by Okamura (in Okamura et al. 1982) from the Kyushu-Palau
Ridge. Pelagic captures have been reported (Sazonov and Iwamoto 1992) in the upper 300 m over
bottom depths of >1,000 m.
Genus Kumba Marshall, 1973
DISTINGUISHING FEATURES.— BR 7; anus in middle 1⁄3of space between A and V, usually clos-
er to the latter. Luminescent organ with one gland and lens immediately anterior to anus. Most dor-
sal surface of snout and almost entire ventral surfaces of snout, suborbital, and lower jaw naked.
No terminal or lateral snout scute. V 8–12; GR-I (inner) 10–14 total. (After Iwamoto and Sazonov
1994.)
REMARKS.— The genus was revised by Iwamoto and Sazonov (1994) to include nine species,
two of which were described as new. The species are known from few and widely scattered spec-
imens. Three species are recorded from Taiwan, two of these were newly recorded from the area
by Shao et al. (2008).
Key to the Species of Kumba in Taiwan
(Adapted from Iwamoto and Sazonov, 1994)
1a. Scaleless areas on dorsal surface of snout extending posteriorly beyond lateral nasal angles;
V 9–10; orbit 37–43% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. punctulata
1b. Scaleless areas on dorsal surface of snout extending posteriorly only to lateral nasal angles;
V 10–13; orbit 26–43% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. No pigment spots along A; barbel short, about 7% HL; orbit 26–27% HL, equal or less than
interorbital width; V 10–11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. gymnorhynchus
2b. Three black pigment spots above mid-length of A; barbel 21–33% HL; orbit 36–43% HL, much
greater than interorbital width; V 11–13. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. japonica
Kumba gymnorhynchus Iwamoto and Sazonov, 1994
Kumba gymnorhynchus Iwamoto and Sazonov, 1994:229, figs. 3–4 (holotype, CAS 77313 [ex ZMMU
P.17765]; Indian Ocean, West Australian Ridge (Broken Ridge), 30°46ʹS, 93°20ʹE, 1260–1370 m;
1 paratype, ZMMU P.17766).— Shao et al., 2008: table 2 (1 spec., Taiwan [SCS], 736–1040 m; first record
from Taiwan and SCS).— Iwamoto et al., 2009:48–49, fig. 5 (data and photograph of Taiwan spec., ASIZP
65527).
MATERIAL EXAMINED.— SCS: ASIZP 65527 (1, 316 TL), CD 134, 1260–1370 m.
DISTINGUISHING FEATURES [ASIZP 65527 in square brackets; ranges from types and NMV
23944].— 1D II [10]11; P [i18] i20–i24; V [8] 10–11; inner GR-I [10] 12–13 total, inner GR-II
[10] 12–13 total. Snout [27] 27–29% HL; orbit about [26] 25–26%; interorbital [24] 24–31%; sub-
orbital [16] 16–17%; postorbital [40] 48–52%; orbit to preopercle [50] 48–50%; upper jaw [41]
39–43%; barbel [12] about 7–11%; length isthmus to anal-fin origin [44] 58–65%. Head relative-
ly deep and compressed; snout blunt, scarcely protruding beyond wide mouth; upper jaw extends
to below posterior 1⁄3of orbit; suborbital region flat, without sharp ridge. Spinules on scales short,
conical, aligned in 1–3 comblike rows; dorsal snout surface naked to lateral nasal angles but not
posteriorly. Spinous second ray of 1D finely serrated. Attains at least 400 mm TL.
DISTRIBUTION.— Originally described from two specimens taken on the West Australian (Bro-
ken) Ridge in the e. Indian Ocean in 1260–1370 m. Our specimen was collected from the South
China Sea off Taiwan in 736–1040 m.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 79

REMARKS.— The Taiwan specimen, as reported by Shao et al. (2008), represents the first
record of the species from the w. Pacific and only the fourth known specimen. The two type spec-
imens were taken off the West Australian [Broken] Ridge; a third specimen (NMV 23944) was
taken off Albany in Western Australia. The species should be expected in intervening areas of the
Pacific and Indian oceans. In the Taiwan specimen the counts of the pelvic and pectoral fin rays,
and the inner gill rakers on first and second gill arches, and the distance between the isthmus and
anal-fin origin were low and must be confirmed when other specimens are captured from the area.
Kumba japonica (Matsubara, 1943)
Lionurus japonicus Matsubara, 1943:149, fig. 9 (holotype, FAKU 1951; Kumano-Nada, Japan [not seen in
2007]; 1 paratype, FAKU 1938 [not seen in 2007]).
Nezumia japonicus: Okamura, 1970:88–91, pl. XIX, text-fig. 39 (5 spec., 135–157 mm TL; s. Japan).
Ventrifossa japonica: Okamura in Okamura et al., 1982:147, 349 (10 spec.; s. Japan and Kyushu-Palau Ridge,
550–710 m).— Okamura in Masuda et al., 1984:94, pl. 81–F (compiled).— Nakabo, 2002:421 (compiled).
Kumba japonica: Iwamoto and Sazonov, 1994:231.— Chiou et al., 2004b:45, fig.14 (2 spec.; sw. Taiwan).—
Shao et al., 2008: table 2 (3 spec.; Taiwan [NET, SWT], 100–600 m).
MATERIAL EXAMINED (10 spec.).— NET: ASIZP 66371 (1, 93 TL), Nan-fang-ao; ASZIP
66941 (1, 140 TL), CD 210, 445+1185 m; ASIZP 70681 (1, 163 TL), Da-xi. SWT: ASIZP 61240
(1, 162 TL), Dong-gang; ASIZP 61241 (1, 156 TL), Dong-gang. Other materials: Japan: NSMT-
P 58943 (1, 15.8 HL, 115 TL), 65441 (1, 17.4 HL, 113 TL), 65731 (1, 15.2 HL, 110 TL), 65733
(1, 16.0 HL, 99+ TL), 91535 (1, 16.4 HL, 111 TL).
DISTINGUISHING FEATURES.— 1D II 8–11; P i16–i22; V 9–12; inner GR-I 12–14 total; pyl.
caeca 40–52. Snout 21–30% HL; preoral 16–22%; internasal width 17–21%; orbit 36–43%;
interorbital 22–28%; upper jaw 37–40%; barbel 18–27%; outer gill slit 17–24%; pre-A 142–155%;
isthmus to A 78–96%; body depth 63–89%; height 1D 83–88%; 1D-2D interspace 63–86%; length
outer V ray 89–171%. Snout bluntly pointed, relatively high, much shorter than orbit diameter;
suborbital ridge prominent; upper jaw extends to below posterior ½ to ¼ of orbit; barbel about two-
thirds of orbit. Underside of head entirely naked; naked areas dorsally on snout extend only to
transverse line crossing lateral snout angles; pores on mandible large and prominent. V origin under
posterior margin of operculum; P origin slightly in advance of 1D origin; A origin behind 1D. Anus
located about midway between A origin and inner V bases; ADW rather large, at or slightly ahead
of line connecting inner V bases. Spinules on scales short, conical, aligned in 5–8 parallel crest-
like rows. Spinous second ray of 1D sparsely serrated. Three small black blotches above mid-
length of A. Species small, probably not attaining more than 170 mm TL.
DISTRIBUTION.— Southern Japan, Kyushu-Palau Ridge, and Taiwan, 550–710 m (Taiwan
records from 100 to 600 m).
REMARKS.— The generic placement of this enigmatic species has been problemmatic; its pre-
vious allocation to three different genera reflects this. One of the current authors (HCH) visited the
FAKU collections in Maizuru in 2007 but was unable to locate the type specimens of this species.
The three small black blotches (faint in some specimens) above the A fin are unique and highly
characteristic of this species.
Kumba punctulata Iwamoto and Sazonov, 1994
Figures 13A–B.
Kumba punctulata Iwamoto and Sazonov, 1994:233–234, figs. 6–7 (holotype, MNHN 1994–0034; off New
Caledonia, 20°54ʹS, 168°21ʹ02ʺE, 530 m; 1 paratype, ZMMU P.17762 (13.4 HL, 89+ TL; Bismark Sea off
New Guinea, 5°20.9ʹS, 146°16ʹE; 0–1000 m).— Iwamoto and Merrett, 1997:526, fig. 23 (holotype listed
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from New Caledonia).— Merrett and Iwamoto, 2000:764 (3 spec.; Vanuatu, 541–577 m).— Shao et al.,
2008: table 2 (17 spec.; Taiwan [SCS], 509–516 m. first record for Taiwan).
MATERIAL EXAMINED (17 spec.).— SCS: ASIZP 66816 (4, 90+-131 TL), CP 315, 509 m;
ASIZP 66861 (1, 95 TL) and ASIZP 66891 (3, 82+-128 TL), OCP 313, 513 m; ASIZP 66877 (2,
105+-118+ TL), CP 314, 506 m; ASIZP 66902 (3, 105–107+ TL), CP 316, 514 m; ASIZP 66942
(1, 90+ TL), CD 311, 516 m; ASIZP 66943 (3, 83+-103+ TL), OCP 317, 515 m.
DISTINGUISHING FEATURES.— 1D II 9–11; P i18–i22; V 9–10 (rarely 11); inner GR-I 12–13
total. Snout 20–29% HL; orbit 37–43%; interorbital (25)28–35%; upper jaw 40–51%; barbel
13–20%. Snout bluntly pointed, relatively high, much shorter than orbit diameter; suborbital ridge
relatively flat, not sharply angular in cross section; upper jaw extends to below of middle one-third
orbit; barbel about 1.4–2.2 times into snout length. Underside of head entirely naked; naked areas
dorsally on snout extend only to transverse line connecting lateral snout angles. Spinules on scales
short, conical, aligned in 5–8 parallel crest-like rows. Spinous second ray of 1D sparsely serrated.
Origin of V under opercle, A origin under anterior half of 1D. A small species, probably not attain-
ing more than 150 mm TL.
DISTRIBUTION.— Known from relatively few captures off Vanuatu and New Caledonia in sw.
Pacific, but numerous specimens were collected in six trawls from the South China Sea off Taiwan
in 509–516 m.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 81
FIGURE 13. Kumba punctulata Iwamoto and Sazonov, 1994. A. ASIZP 66877, 1 of 2, 105 mm TL fresh. B. ASIZP 66816,
1 of 4, 131 mm TL, preserved.
A
B

REMARKS.— Our Taiwan specimens (as listed by Shao et al., 2008: table 2) represent the first
record of the species from this region and from the nw. Pacific.
Genus Kuronezumia Iwamoto, 1974
DISTINGUISHING FEATURES.— BR 7. Body and head compressed and deep; snout rounded in
profile, almost entirely covered with small uniform, finely spinulated scales; suborbital region ver-
tical, without angular midlateral ridge, covered with small unmodified scales. Teeth in broad villi-
form bands in both jaws, outer series of upper jaw slightly enlarged. GR-I 8–11 total. Body scales
small, adherent, densely covered with long slender spinules. Anus removed from A origin, closer
to V bases; a dermal light organ between V bases. Color overall from gray to brown to swarthy;
fins uniformly dusky to black. (After Sazonov and Iwamoto, 1994:65–65.)
REMARKS.— Seven species currently recognized, only one known from Taiwan. The genus
was reviewed by Shcherbachev et al. (1992).
Kuronezumia dara (Gilbert and Hubbs, 1916)
Figure 14.
Lionurus darus Gilbert and Hubbs, 1916:197–199, pl. 10, fig. 1 (holotype, 132 mm TL, USNM 76867; Suru-
ga Gulf, Japan, 35°06ʹN, 138°40ʹ10ʺE, 197 fm [360 m]).
Nezumia darus: Okamura, 1970:101–102 (description from Gilbert and Hubbs, 1916).— Okamura in Okamu-
ra et al., 1982:161, 349, fig. 95 (p. 160) (2 spec., 130–144 mm TL; Tosa Bay, 355–605 m).— Okamura
in Masuda et al., 1984:95, fig. 81–J (compiled).— Okamura in Okamura and Kitajima, 1984:217, 363,
fig. 153 (p. 216) (2 spec., 220–318 mm TL; East China Sea [Okinawa Trough], 560–692 m).— Nakabo,
2002:424 (compiled).
Kuronezumia darus: Shao et al., 2008: table 2 (1 spec., Taiwan [SWT], 280–452 m; first record from Taiwan).
Kuronezumia dara: Shcherbachev et al., 1992:100–101(mentioned, no additional specimens).
MATERIAL EXAMINED (1 spec.).— SWT: ASIZP 65514 (1, 482 TL), CD 140, 280–452 m.
Other specimens: Japan: BSKU 27666 (1, 49.8 HL, 255+ TL), 26326 (1, 34.6 HL, 209+ TL),
44828 (1, 42.4 HL, 218+ TL), 45036 (60.0 HL, 376+ TL); HUMZ uncat. (1, 22.0 HL, 138+ TL).
DISTINGUISHING FEATURES.— 1D II, 9–10; P i21–i23; V 10–12, usually11; GR-I (outer/inner)
0+(1–8) / 2+(8–9) , GR-II 1+(7–8) / 1+(7–9); scale rows below 1D origin 12–15, below 2D origin
10.5–11.0; below mid-base 1D 8.5–10; over distance equal to pre-1D 35–42. Snout 24–30% HL;
preoral length 15–19%; internasal width 18–19; orbit 25–33%; interorbital 24–25; postorbital
length 44–49%; orbit to angle of preopercle 36–42%; upper jaw 30–35%; barbel 20–27%; length
outer gill slit 16–20%; greatest body depth 85–97%; pre-A length 139–152%; isthmus to A
57–79%; 1D height 97–98% (3 spec.); 1D base length 28–31; 1D-2D interspace 23–41%; length
outer V ray 68–87%. Head compressed, much deeper than wide; snout low, bluntly rounded,
scarcely protruding beyond large mouth, lacking stout spinous tubercle at tip; upper jaw extends
posteriorly about to below mid-orbit. Barbel moderately long, slender, about 0.5–0.8 of orbit.
Scales small, densely covered with slender, conical spinules giving velvety feel to body surfaces;
scale ridges on head not especially developed, head contours smoothly rounded. Color medium
brown to swarthy, juveniles blackish overall, fins dark (usually black). A moderately large species,
probably attaining about 500 mm TL; the current Taiwan specimen is the largest recorded.
DISTRIBUTION.— Pacific coast of s. Japan in 360–692 m and SCS off Taiwan in 280–452 m.
REMARKS.— Shao et al. (2008) first recorded the species from Taiwan and the South China
Sea based on the ASIZP specimen. Kuronezumia dara is quite similar in many respects to K. bubo-
nis Iwamoto, 1974, which has been recorded from the Atlantic, Pacific, and Indian oceans, includ-
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ing the South China Sea off Vietnam (Shcherbachev et al. 1992:99–100). A notable difference is
the tubercular swelling housing the light organ in K. bubonis, which is absent in K. dara.
The species-group name darus has been used by most Japanese authors even after the name
had been transferred to the genus Kuronezumia by Shcherbachev et al. (1992), to which transfer
those Japanese authors agree. Gilbert and Hubbs (1916) originally described the species in the mas-
culine genus Lionurus, taking the Japanese term dara and latinizing it to darus to agree in gender
with the genus name. When the specific name was combined with the feminine genus
Kuronezumia, under Article 31.2 of the International Code of Zoological Nomenclature the
“species-group name, if it is or ends in a Latin or latinized adjective or participle in the nominative
singular, must agree in gender with the generic name with which it is at any time combined” (Inter-
national Commission on Zoological Nomenclature 1999:38). Thus, the masculine species-group
name darus must be changed to the feminine dara.
Genus Lucigadus Gilbert and Hubbs, 1920
Lucigadus Gilbert and Hubbs, 1920:553 (as subgenus of Ventrifossa; type species Macrourus lucifer Smith
and Radcliffe, 1912, by original designation).— Sazonov, 1985:17 (elevated subgenus Lucigadus to genus
level).
Lucigadella Gilbert and Hubbs, 1920:552 (as subgenus of Ventrifossa; type species Macrourus nigromargina-
tus Smith and Radcliffe, 1912, by original designation)
DISTINGUISHING FEATURES.— BR 7; chin barbel present; spinous ray of 1D serrated; anus
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 83
FIGURE 14. Xuronezumia dara (Gilbert and Hubbs, 1916). ASIZP 65514, 482 mm TL. A. lateral view.
B. dorsal view.
A
B

removed from A, closer to V bases; light organ well developed, two dermal windows, one before
anus, the second between V bases; ventral region of body appearing to have swung far forward so
that gill membranes unite below or forward of preopercle, pelvic fins below opercle; suborbital
shelf covered with several rows of small relatively unmodified scales, with no sharp ridges; under-
side of snout mostly or completely scaled; spinules on body scales usually aligned in parallel rows;
teeth in both jaws small, in tapered bands, premaxillary band not reaching beyond posterior edge
of maxillary process. (Adapted from Iwamoto and Merrett 1997:526).
REMARKS.— Seven species, only one in Taiwan. Most species smaller than 20 cm TL.
Lucigadus nigromarginatus (Smith and Radcliffe, 1912)
Macrourus nigromarginatus Smith and Radcliffe in Radcliffe, 1912:114, Pl. 24, fig. 2 (holotype, USNM
72930; near Simaluc I., Philippines, 5°33ʹ15ʺN, 120°15ʹ30ʺE, Albatross sta. 5569, 303 fm [554 m]).
Lionurus (Nezumia)nigromaculatus: Gilbert and Hubbs, 1916:145, 192 (name only, mispelled).
Ventrifossa (Lucigadella) nigromarginata: Gilbert and Hubbs, 1920:552–553 (44 spec., Philippines, Borneo,
Celebes, Java Sea, in 135–392 fm [247–717 m]; new subgenus erected).
Ventrifossa nigromarginata: Shen et al., 1993:172 (descr.).— Chiou et al., 2004b:37, 47 (in key, listed from
Taiwan).
Lucigadus nigromarginatus: Iwamoto and Williams, 1999:185 (mentioned).— Shao et al., 2008: table 2 (2
spec., Taiwan [NET, SWT], 100–600 m).
Lucigadus lucifer (non Smith and Radcliffe): Chiou et al., 2004b:46–47, fig. 15 (incorrect identification; 1
spec. from Taiwan).
MATERIAL EXAMINED (9 Taiwan spec.).— NET: ASIZP 57531 (1, 193 TL), Hsiao-liou-chiou;
ASIZP 61308 (1, 110 TL), Nan-fang-ao. SWT: ASIZP (1, 158 TL), CD 142, 227–335 m; ASIZP
66322 (1, 153 TL), Dong-gang; ASIZP 67599 (1, 133 TL, 13 HL), CP 347, 305 m. SCS: ASIZP
65547 (2, 105–155 TL), CD 134, 736–1040 m. Other specimens:ASIZP 67968 (1, 149 TL), Auro-
ra, 507–540 m. The holotype (USNM 72930) and 9 paratypes were previously examined (by TI) at
the USNM. ASIZP 61630 (1, 135 TL), New Caledonia, 385–401 m.
DISTINGUISHING FEATURES.— 1D II,9–11; P i18–i21; V 10–11; inner GR-I 10–12 total, outer
GR-II 9–11 total; scales below 1D 14–16, below 2D 7–9.5, lateral line scales over distance equal
to pre-1D length 37–42. Snout length 24–28% HL; ventral length of snout 17–20%; orbit 33–36%;
interorbital 22–26%; upper jaw 33–39%; barbel 20–26%; outer gill slit 18–21%; body depth
85–94%; height 1D 92–107%. Snout high, smoothly and bluntly rounded; nape with moderate
arch; upper jaw extends posteriorly to below mid-orbit; gill openings wide, extending forward
under or anterior to posterior margin of orbit; scales small, covered with small, recurved conical
spinules in parallel rows, no modified thickened scales on head ridges nor tubercular scutes on
snout; 1D with black blotch, anterior portion of A black, no blotches or marking on body. A small
species attaining approximately 180 mm TL.
DISTRIBUTION.— Originally recorded from Philippines and Indonesia in 266–718 m; Taiwan
records from 100–600 m.
REMARKS.— Taiwan appears to be the n. limit of Lucigadus nigromarginatus. The species is
apparently not uncommon in the Philippines and the Indo-Malaysian Archipelago.
Genus Macrosmia Merrett, Sazonov and Shcherbachev, 1983
Macrosmia Merrett, Sazonov and Shcherbachev, 1983 (type species Macrosmia phalacra Merrett, Sazonov
and Shcherbachev, 1983, by original designation).
DISTINGUISHING FEATURES.— BR 7; V 11–12; pyl. caeca 15–18. Chin barbel present; spinous
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ray of 1D weakly serrated; anus in small periproct immediately before A; no light organ; V origin
anterior to that of P; A origin below hind margin of 1D; head squamation reduced, mostly lacking
on snout and underside of head, no scaled ridges on head and snout; teeth in bands in both jaws;
olfactory organ massive, posterior nostril about equal to diameter of pupil, 4–5 times in HL.
(Adapted from Merrett and Iwamoto 2000:767).
REMARKS.— Only the single species known. Relationships obscure, but probably most close-
ly related to Asthenomacrurus or Pseudonezumia. Broad distribution, having been captured in the
e. North Atlantic, w. South Atlantic, se. Indian Ocean, and w. Pacific Ocean.
Macrosmia phalacra Merrett, Sazonov and Shcherbachev, 1983
Figures 15A–B.
Macrosmia phalacra Merrett, Sazonov and Shcherbachev, 1983:554, figs. 1–2 (holotype, ZMMU P.15390,
31.5 mm HL, 217 mm TL; Ninety East Ridge, se. Indian Ocean, 17°04.8ʹS, 88°09.0ʹE, 1650–1660 m;
paratype, BMNH 1980.12.31.2, 28.5 HL, 177+ TL; Canary Passage, e. North Atlantic).— Merrett and
Iwamoto, 2000:767–768, fig. 25 (1 spec., 207+ TL; Vanuatu, 1160–1220 m).— Shao et al., 2008: table 2
(8 spec., Taiwan [SCS], 1098–1293 m; first record from Taiwan).— Melo et al., 2010:35 (off Brazil).
MATERIAL EXAMINED (9 spec.).— SCS: ASIZP 66730 (1, 152+ TL) and ASIZP 66919 (3,
120+-145+ TL) and CAS 224493 (ex ASIZP 66919)(2, 120+-178 TL), CD 322, 1098 m; ASIZP
66775 (1, 150+ TL), CD 324, 1293 m. Other material. ASIZP 68275 (1, 128 TL), Aurora,
1262–1360 m.
DISTINGUISHING FEATURES.— 1D II,9; P i20–i21; V 11–12; total GR-I (outer/inner) 8/10, GR-
II 11/10; scales below mid-1D 4, below 2D 6, lateral line scales over distance equal to pre-1D
length about 29. Snout length 28–33% HL; preoral 20–22%; interorbital 22–25%; posterior nostril
about 16%; orbit 23–29%; suborbital 13–21; orbit-preopercle 45–48%; upper jaw 30–41%; barbel
21–26%. Snout blunt, high; upper jaw extends to below mid-orbit; orbit slightly less than snout
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 85
FIGURE 15. Macrosmia phalacra Merrett, Sazonov and Shcherbachev, 1983. A. ASIZP 66919, 145+ mm TL, fresh.
B. CAS 224493, 178 mm TL, preserved.
A
B

length; suborbital region vertical and rounded; gill openings wide, extending forward to under pos-
terior margin of orbit; scales small, deciduous, entirely absent on underside of head and above
snout; pores of cephalic lateral-line system well developed. Coloration mostly black to dark brown.
Attains about 220 mm TL.
DISTRIBUTION.— Originally described from the ne. Atlantic and se. Indian Ocean, and subse-
quently recorded off Vanuatu in the sw. Pacific and off Brazil in the sw. Atlantic, in 1060–1699 m.
Our Taiwan specimens are the first from this area and were taken at depths of 1098–1293 m.
REMARKS.— Macrosmia phalacra is turning out to be much more widely distributed than
thought when originally described. The lack of specimens from the area between the North Atlantic
and e. Indian Ocean leaves a perplexing distributional gap, but it may simply represent a collect-
ing artifact.
Genus Malacocephalus Günther, 1862
Malacocephalus Günther, 1862 (type species Macrourus laevis Lowe, 1843 by monotypy).
DISTINGUISHING FEATURES.— BR 7; V usually 8 or 9; inner GR-I 9–14; pyl. caeca 50–100 or
more. Snout smoothly rounded; sides of head relatively compressed and vertical; mouth large, usu-
ally >45% HL, upper jaws extend to or beyond vertical through posterior edge of orbits; premax-
illary teeth in two rows to moderate band, with outer series enlarged; dentary teeth large, widely
spaced, in one row laterally; chin barbel present; spinous ray of 1D smooth (subgenus Malaco-
cephalus) or serrated (subgenus Pawnurus); periproct in middle third of space between V and A,
usually closer to V; two dermal windows of light organ, one between V bases, the other within
naked periproct region and before anus. Scales densely covered with fine needlelike spinules giv-
ing velvety feel to surface; no coarsely scaled ridges or scutes on head and snout; lowermost BR
scaled. Gill membranes narrowly united over isthmus, with a free fold; gill openings extend for-
wards under orbits. Two short, broad retia and two wide but short gas glands.
REMARKS.— Two subgenera (Malacocephalus and Pawnurus) and seven species known, but
two or three species of subgenus Malacocephalus may be synonyms of the widely distributed
M. laevis.
Malacocephalus nipponensis Gilbert and Hubbs, 1916
Malacocephalus nipponensis Gilbert and Hubbs, 1916:189–191, pl. 9, fig. 2 (holotype, USNM 76866, 460
mm TL; e. coast Japan, Albatross st. 4967, 244–253 fm [446–463 m].— Okamura in Okamura et al.,
1982:145, 347, fig. 88 (p. 144) (1 spec., 472 mm TL, Kyushu-Palau Ridge, 453 m).— Okamura in Masu-
da et al., 1984:94, pl. 80–J (compiled).— Okamura in Okamura et al., 1984:144, 360, fig. 145 (p. 202)
(18 spec., 270–480 mm TL, East China Sea [Okinawa Trough], 420–550 m).— Shen et al., 1993:172
(descr.).— Chiou et al., 2004b:37, 47 (in key, list).— Shao et al., 2008: table 2 (2 spec., Taiwan [SCS],
979–1268 m, first record for Taiwan).
Malacocephalus laevis: Okamura, 1970:69–73, pl. IV, text-fig. 29–31(56 spec., 255–520 mm TL; Pacific
coast of s. Japan, 350–500 m).— Shao et al., 2008: table 2 (9 spec., NET, ET, SWT).
MATERIAL EXAMINED (20 spec.).— NET: ASIZP 60015 (1, 445 TL), Da-xi; ASIZP 61312 (1,
336 TL; Nan-fang-ao; ASIZP 61313 (3, 184–252 TL), Nan-fang-ao; ASIZP 65517 (1, 68 HL, 405
TL), CD 210; ASIZP 70229 (1, 287 TL), Da-xi. SWT: ASIZP 61314 (1, 172 TL), Dong-gang;
ASIZP 62331 (2, 250+-305+ TL), Fong-gang, 200 m; ASIZP 65517 (1, 405 TL), CD 210,
445–1185 m; ASIZP 65597 (2, 360–370 TL), CD 137, 316–477 m; ASIZP 70615 (1, 196 TL),
Dong-gang). SCS: ASIZP 58031 (2, 361–389 TL), Tong-sa Islands); ASIZP 66277 (1, 220+ TL),
OCP 312, 517 m; ASIZP 66745 (1, 178+ TL), CP 314, 506 m; FRIP 0669 (1, 302 TL), FRI, 630
m; NMMSTP 0907 (1, 290+ TL), Tong-sha Islands, 515 m.
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DISTINGUISHING FEATURES.— 1D II,10–14; P i16–i22; V 9 (8–10); inner GR-I 2–4+7–9; pyl.
caeca 107–130. Snout 23–28%; orbit 29–37%; suborbital 11–14%; interorbital 26–31%; orbit-pre-
opercle 42–50%; upper jaw 45–56%; barbel 18–22%. Snout bluntly pointed; mouth large, upper
jaw extends to below hind margin of orbit; orbit large, greater than snout length; suborbital region
vertical and smoothly curved; gill openings wide, extend forward to under posterior margin of
orbit. Scales small, beset with fine, slender spinules; scales uniformly and smoothly cover head and
body; no coarsely modified scales on ridges of head or tip of snout, lower branchiostegal rays
scaled, but gular membrane naked. Light organ with anterior dermal window relatively small and
round, situated between or slightly anterior to V bases; periproct enclosing posterior dermal win-
dow and urogenital openings located between V fins, far removed from A origin. Coloration gray
to swarthy with silvery sides, blackish ventrally on head and over abdomen; fins dark dusky to
black. Attains >520 mm TL. (Mostly after Okamura 1970 and from Taiwan specimens.)
DISTRIBUTION.— Pacific coast of s. Japan, East China Sea [Okinawa Trough], Kyushu-Palau
Ridge, and Taiwan off ne., sw. coast and in South China Sea in 316–1185 m.
REMARKS.— Malacocephalus nipponensis was first synonymized into M. laevis Lowe, 1843
by Okamura (1970), but later (Okamura in Okamura and Kitajima 1982; Okamura in Okamura et
al. 1984) considered it a valid species based on the absence of scales on the gular membrane (usu-
ally present in M. laevis) and the small round anterior dermal window (bean-shaped in M. laevis).
Iwamoto (1979:149) suggested that M. laevis, M. nipponensis, and M. hawaiiensis Gilbert, 1905
may eventually prove to be the same. Although specimens of this species are available in many col-
lections around the world, no one has yet to comprehensively study these collections. A molecular
study may prove the easiest and the most-effective approach to resolving this question of how
many species are involved in this clade. We have taken a conservative approach and treat these
northwestern Pacific specimens as M. nipponensis, following Okamura.
Genus Mataeocephalus Berg, 1898
Mataeocephalus Berg, 1898:41 (replacement name for Coelocephalus Gilbert and Cramer, 1897, preoccu-
pied).
Coelocephalus Gilbert and Cramer, 1897:422 (non Agassiz, 1843) (type species Coelocephalus acipenser-
ineus Gilbert and Cramer, 1897, by monotypy.)
DISTINGUISHING FEATURES.— BR 6 or 7; V 7–9; inner GR-I 6–8, none or few rudiments in
outer series; pyl. caeca 8–20. Snout pointed, armed with two tubercular scutes at tip, underside
fully scaled to entirely naked; snout protruding well beyond small, almost inferior mouth; upper
jaws about 1⁄3of HL; chin barbel short; suborbital ridge sharp, angular in cross section, strongly
armed with modified scutelike scales. Gill slits restricted by membranes across upper and lower
arms; gill membranes broadly attached to isthmus, restricting opercular opening; gular and BR
membranes naked. Dentition in both jaws in broad short bands, outer premaxillary series slightly
enlarged. Spinous ray of 1D with rudimentary or well-developed serrations. Periproct moderately
large, located within middle one-third of space between V and A (but closer to A in one species),
rudimentary light organ developed before anus.
REMARKS.— Sazonov et al. (2003) provided a comprehensive review of the genus and
described two new species. They also removed Macrourus hyostomus Smith and Radcliffe, 1912
from either Coryphaenoides or Hyostomus and included it as a subgenus of Mataeocephalus. Six
species, two of which are found in Taiwan waters, although two others (M. adustus and
M. accipenserinus) could be expected, based on their presence in the SCS, and for the latter, in n.
Philippine Islands.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 87

Key to Species of Mataeocephalus in Taiwan
1a. BR 6; body scales densely covered with slender somewhat flattened, reclined spinules in some-
what convergent to divergent rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. hyostomus
1b. BR 7; body scales with needle-like spinules in 12–18 parallel rows, middle row slightly
enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. cristatus
Mataeocephalus (Mataeocephalus) cristatus Sazonov, Shcherbachev and Iwamoto, 2003
Figures 16A–C.
Mataeocephalus cristatus Sazonov, Shcherbachev and Iwamoto, 2003:290–291, figs. 3–4 (holotype, 48 mm
HL, 215 mm TL; ZMMU P.15345, Ninety East Ridge, 11°31ʹS, 88°55ʹE, 1600–1700 m;19 paratypes,
152–271 mm TL; w. tropical Pacific and Indian Ocean, 1000–1720 m).— Shao et al., 2008: table 2 (2
spec., Taiwan [SCS], 227–1010 m; first record from Taiwan).
MATERIAL EXAMINED (3 spec.).— SCS: ASIZP 66077 (1, 188 TL) and ASIZP 66912 (1, 216+
TL), CD 322, 1098 m. SET: ASIZP 67386 (1, 52 HL, 262 TL), CP 366, 1032 m.
DISTINGUISHING FEATURES.— BR 7; P i17–i21; V 7. Snout 35–41% HL, 1.4–2.0 times larger
than orbit; orbit 21–25% HL, about equal to interorbital space; orbit to angle of preopercle 33–38%
HL. Snout depressed, narrowly pointed in lateral view, broadly triangular in dorsal view, tipped
with stout spiny scutes; mouth small, underslung, U-shaped; underside of head fully scaled; spin-
ous second ray of 1D weakly serrated, slightly prolonged, its height usually 60–84% HL; body
scales with needle-like spinules in 12–18 parallel rows, middle row slightly enlarged; window of
light organ absent; periproct relatively small, about midway between V and A. Attains at least 271
mm TL.
DISTRIBUTION.— Broadly distributed in the w. tropical Pacific and Indian Ocean in
1000–1720 m. Our two specimens were collected from the South China Sea off Taiwan in
1032–1098 m.
REMARKS.— Shao et al. (2008: table 2) first recorded the species from Taiwan based on the
current specimens. Mataeocephalus cristatus is quite distinct from M. hyostomus, its congener in
Taiwan, which is classified in a separate subgenus. The two species can be distinguished by a com-
bination of characters including BR 7 (vs. 6 in M. hyostomus), V ray count (7 vs. 7–8, rarely 9, in
M. hyostomus), height of spinous 1D ray (<HL in M. cristatus, greatly elongated, 145–295% HL
in M. hyostomus), snout length (35–41% HL vs. 30–36%), orbit-preopercle (33–38% HL vs.
40–44%), and length upper jaw (20–27% HL vs. 28–31%). Mataeocephalus accipenserinus and
M. adustus are distinguished from M. cristatus by their naked underside of snout. (Mostly adapted
from Sazonov et al., 2003.)
Mataeocephalus (Hyostomus)hyostomus (Smith and Radcliffe, 1912)
Figures 17A–B.
Macrourus hyostomus Smith and Radcliffe in Radcliffe, 1912:121–122, pl. 27, fig. 1 (holotype, 280 mm TL,
USNM 72938, Lagonoy Gulf, Luzon I., Philippines; Albatross sta. 5470, 560 fm [1024 m]; paratypes from
Sibuku Bay, Borneo, 750 m, and Buton Strait, Celebes, 1022 m).
Coryphaenoides (Hyomacrurus) hyostomus: Gilbert and Hubbs, 1920:422–424 (redescription of types;
described new subgenus Hyomacrurus).
Hyomacrurus hyostomus: Marshall, 1973:565 (listed).
Mataeocephalus (Hyomacrurus) hyostomus: Sazonov et al., 2003:294–296, fig. 6 (7 spec. [including holo-
type], 36.5–65.5 HL, 160+–283 TL; Philippines, 760–1100 m).— Shao et al., 2008: table 2 (2 spec., Tai-
wan [SWT, SCS], 227–1040 m; first record from Taiwan).
88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 89
FIGURE 16. Mataeocephalus (Mataeocephalus) cristatus Sazonov, Shcherbachev and Iwamoto, 2003, ASIZP 66386, 262
mm TL. A. lateral view. B. dorsal view of head.
FIGURE 17. Mataeocephalus (Hyostomus) hyostomus (Smith and Radcliffe, 1912), ASIZP65550, 220 mm TL.
A. lateral view. B. dorsal view of head.
A
B
A
B

MATERIAL EXAMINED (2 spec.).— SCS: ASIZP 65541 (1, 180+ TL), CD 142, 227–335 m;
ASIZP 65550 (1, 220 TL), CD 134, 736–1040 m.
DISTINGUISHING FEATURES.— BR 6; P i16–i20; V 7–8; inner GR-I 5–8; scales below mid-1D
5.0–7.5; pyl. caeca 8–13. Snout 30–36% HL; orbit 22–25%; interorbital 22–26%; orbit to angle of
preopercle 40–44%. Snout broad, moderately pointed and protruding; upper jaws extend posterior-
ly to below posterior 1⁄4to 1⁄3of orbit; spinous second ray of 1D weakly serrated near base, pro-
longed, 1.5 or more times HL; body scales densely covered with slender, slightly flattened, reclined
spinules in somewhat convergent to divergent rows; underside of head almost fully scaled; window
of light organ scarcely discernible from exterior; periproct relatively moderate in size, situated
between V and A but closer to former. Attains at least 283 mm TL. (Adapted from Sazonov et al.
2003.)
DISTRIBUTION.— Known only from the Philippines, Indonesia, and Taiwan in 227–1100 m.
REMARKS.— Our two specimens collected in the South China Sea in 227–1040 m were first
recorded from Taiwan and the South China Sea by Shao et al. (2008: table 2). Mataeocephalus
hyostomus is closely similar to M. kotlyari Sazonov, Shcherbachev, and Iwamoto, 2003, which is
found in the sw. Pacific, but M. hyostomus has more pyl. caeca (16–19 cf. 8–13 in M. kotlyari),
V 7, rarely 6 vs. 7–8, rarely 9, and shorter 1D (88–119% HL vs. 124–295%).
Genus Nezumia Jordan, 1904
Nezumia Jordan, 1904:620 (type species Nezumia condylura Jordan and Gilbert, 1904, by original designa-
tion).
DISTINGUISHING FEATURES.— BR 7; V 7–17 or more; inner GR-I usually 12 or fewer; mouth
subterminal, upper jaw usually <40% HL; chin barbel present; spinous ray of 1D serrated; anus
removed from A, closer to V bases; periproct located between V and A, usually closer to V; light
organ well developed, a small dermal window between V bases; V base usually under P base; sub-
orbital shelf with two rows of coarsely modified scutelike scales; underside of snout scaled or
naked; spinules on body scales variable, from conical to broadly triangular, in parallel to widely
divergent rows; teeth in both jaws in tapered bands, outer premaxillary tooth series slightly
enlarged and not extending beyond maxillary process. Ground color usually light brown to black-
ish to bluish; ventral and abdominal area of trunk generally dark, often underlain with blue integu-
ment.
REMARKS.— A diverse genus with approximately 48 species, but some of these may be allo-
cated to other genera in future; others are yet to be described. There are five species represented in
Taiwan waters.
Key to the Species of Nezumia in Taiwan
1a. V 8–10; 1D usually overall dusky to blackish, without prominent black blotch . . . . . . . . . . . 2
1b. V 13–17; 1D with distinct black distal tip contrasting with pale proximal part of fin . . . . . . 4
2a. Underside of head mostly scaled except under snout; length second spinous 1D ray usually
< HL; body scales with narrowly triangular to broadly lanceolate spinules. . . . . . . . . . . . . . 3
2b. Underside of head almost entirely naked; length second spinous 1D ray > 95% HL; body scales
densely covered with needle-like spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. spinosa
3a. V 8 or 9, rarely 10; underside of snout naked only along median line . . . . . . . . . . . N. proxima
3b. V 10; underside of snout virtually entirely naked . . . . . . . . . . . . . . . . . . . . . . . . . N. cf. coheni
90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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4a. V 13–15; scale rows below mid-1D 7.0–7.5, below 2D origin 8.5–10.0; length outer V ray
66–127% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. evides
4b. V 13–17; scale rows below mid-1D 8.0–9.5, below 2D origin 10.0–13.0; length outer V ray
68–81% HL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. condylura
Nezumia condylura Jordan and Gilbert, 1904
Nezumia condylura Jordan and Gilbert in Jordan and Starks, 1904:620–621, pl. 4, fig. 2 (holotype, 195 mm
TL, USNM 50937; Suruga Bay, Japan, 207–257 fm [379–470 m]; paratypes, CAS-SU 8551).— Okamu-
ra, 1970:85–88, pl. VII, text-fig. 38 (71 spec., 153–207 mm TL; Pacific coast s. Japan, 360–720 m).—
Okamura in Okamura et al., 1982:161, 350, fig. 96 (p. 160) (7 spec., 152–200 mm TL; Kyushu-Palau
Ridge).— Okamura in Masuda et al., 1984:95, pl. 81–G (compiled).— Okamura in Okamura and Kita-
jima, 1984:215, 362, fig. 151 (5 spec., 170–204 mm TL; Okinawa Trough [East China Sea], 510–760
m).— Iwamoto, 1990:269–370, fig. 625–626 (descr. compiled).— Chiou et al., 2004b:45, fig.16 (25
spec.; Taiwan [NET, ET, SCS], 400–1211 m).
Lionurus condylura: Gilbert and Hubbs, 1916:195–197 (29 spec., 103–204 mm TL; Pacific coast s. Japan,
197–440 m).
MATERIAL EXAMINED (33 spec.).— NET: ASIZP 61238 (1, 186 TL), Nan-fang-ao; ASIZP
61239 (8, 116–124 TL), Nan-fang-ao; ASIZP 64104 (1, 193 TL), CP 235, 764 m; ASIZP 65552 (1,
110 TL) and ASIZP 65607 (2, 114+-162+ TL); CD 210, 445–1185 m; ASIZP 65625 (2, 139–153
TL), CD 211, 517–529 m; ASIZP 65631 (1, 168 TL), CD 209, 508–522 m; ASIZP 65638 (1, 164
TL), CD 214, 488–1027 m; ASZIP 65643 (1, 132 TL), Da-xi; ASIZP 65663 (1, 192 TL), Da-xi;
ASIZP 70683 (3, 25–28 HL), Da-xi. SCS: ASZIP 58022 (2, 131–133 TL), Tong-sha Islands;
ASIZP 65587 (1, 155+ TL), CD 136, 998–1211 m; ASIZP 66803 (3, 27.8–33.0 HL, 160+-210+
TL); CD 320, 731 m; ASIZP 66831 (2, 20–23.6 HL, 105+-117+ TL); CP 315, 509 m; ASIZP 66894
(2, 16.8–22.8 HL, 82+-130+ TL), OCP 312, 517 m. Other specimen: ASIZP 68038 (1, 120+ TL;
Aurora, 500–524 m).
DISTINGUISHING FEATURES.— 1D II,10–13; P i18–i22; V 13–17; inner GR-I 8–11 total; scale
rows below midbase 1D 8–9.5, below 2D 10–13; pyl. caeca 25–28. Snout length 27–30% HL;
interorbital 23–27%; orbit 29–36%; upper jaw 29–33%; barbel 14–21%. Body relatively deep,
about equal to HL; head 6.4–6.7 in TL; snout short, high, projecting slightly beyond mouth, with
stout tubercles at tip and lateral angles; mouth rather small, upper jaw extends posteriorly to under
posterior margin of pupil; barbel well developed, about 2⁄3orbit diameter; suborbital ridge sharp,
beset with two rows of thickened scales; underside of head naked except along peopercle; pores of
cephalic lateralis system prominent; body scales densely covered with short, conical spinules
arranged in 4–16 parallel rows. Spinous 1D ray about equal to HL, with slender, widely spaced ser-
rations. Anterior dermal window of light organ small, situated between inner V-fin bases; anus clos-
er to A than to outer base of V. Color yellowish gray to grayish brown, trunk, gill and gular mem-
branes purplish-black; mouth and gill cavities dark; 1D with black tip, V and anterior portion of
A black, other fins pale. Attains about 210 mm TL.
DISTRIBUTION.— Pacific coast of s. Japan, East China Sea, and South China Sea in 360–910
m, and off Taiwan in 400–1211 m.
REMARKS.— Differences between N. condylura and N. evides are slight and need to be more
adequately determined. Iwamoto (1990:281) differentiated the two based on V fin-ray count, length
P, and position of anus relative to V and A, but in the same volume (1990:270) said “this species
may be the same as N. propinqua.” Okamura (1984) cited “the shorter distance between the vent
and the origin of anal fin, dense outer premaxillary teeth, etc.” to distinguish
N. condylura from N. propinqua.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 91

Nezumia evides (Gilbert and Hubbs, 1920)
Figure 18.
Lionurus evides Gilbert and Hubbs, 1920:557, fig. 39 (holotype, USNM 78231; near Sibuko Bay [Celebes
Sea], Borneo, 4°12ʹ10ʺN, 118°38ʹ08ʺE, 260 fm [475 m]; 7 paratypes).
Nezumia evides: Iwamoto, 1990:262, fig. 614 (in key).— Iwamoto and Williams, 1999:201 (mentioned).—
Shao et al., 2008: table 2 (5 spec., Taiwan [NET, SCS], 488–1027 m; first record from Taiwan).
MATERIAL EXAMINED (17 spec.).— NET: ASIZP 65641 (4, 147–175+ TL), CP 214, 488–1027
m; ASIZP 70699 (2, 156–162 TL), Da-xi. SCS: ASIZP 66830 (1, 127+ TL), OCP 317, 515 m.
Other specimens: CAS-SU 25467 (4 paratypes, 19.6–27.5 HL, 104+-146+ TL), off Borneo, 475
m; CAS-SU 25468 (3 paratypes, 22.8–23.5 HL, 12–132+ TL), Molucca Sea off Halmahera [Gillo-
lo], 545 m; and Aberdeen Fishery Station, Hong Kong uncat. (3, 22.3–27.5 HL, 120–160 TL), SCS
s. of Hainan Is., 200–400 fm [366–732 m].
DISTINGUISHING FEATURES.— 1D II,9–13; P i16–i21; V 13–15; inner GR-I 8–11 total; scale
rows below midbase 1D 7.0–7.5, below 2D 8.5–10; pyl. caeca about 25. Snout length 27–33% HL;
interorbital 24–29%; orbit 31–36%; orbit to angle of preopercle 32–36%; upper jaw 29–34%; bar-
bel 13–18%; length outer ray V 83–127%. Body relatively deep, 82–92% HL; head 4.1–5.8 in TL;
snout short, high, projecting slightly beyond mouth, with stout tubercles at tip and lateral angles;
mouth rather small, upper jaw extends posteriorly to under posterior margin of pupil; barbel well
developed, about 2⁄5to 1⁄2orbit diameter; suborbital ridge with coarsely modified scales; underside
of head to end of upper jaw and mandibular rami naked; body scales densely covered with short,
conical spinules arranged in 8–13 parallel rows. Spinous 1D ray about equal to HL, denticulation
on leading edge widely spaced; outer V ray extends to 10th to 20th A ray. Anterior dermal window
of light organ small, situated between inner V bases; anus between V and A, but usually closer to
inner V bases. Color in alcohol overall brownish, opercle and abdomen blackish with silvery sheen,
abdomen underlain with bluish; mouth and gill cavities dark; 1D with distinct black tip. A small
species, probably not much >160 mm TL.
DISTRIBUTION.— From Taiwan, South China Sea, Philippines, Celebes Sea, Molucca Sea, in
475–1027 m; the Taiwan specimens were collected in 517 m (NET) and 488–1027 m (SCS).
REMARKS.— Our specimens represent the northernmost record, and the first record (as report-
ed by Shao et al. 2008) from Taiwan and the South China Sea. Nezumia evides closely resembles
N. condylura and N. propinqua in having similar head shape, high V counts, naked underside of
snout and mandibles, black-tipped 1D, and scale size and spinulation. The slightly lower counts of
V rays, somewhat longer outer V ray, slightly fewer scales below mid-1D and below 2D origin
(7–7.5 vs. 8–9.5 and 8.5–10 vs. 10–13, respectively), shorter body (HL 4.1–5.9 in TL vs. 6.4–6.7),
and possibly smaller size attained distinguish N. evides from the two.
92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 18. Nezumia evides (Gilbert and Hubbs, 1920). ASIZP 66830, 127 mm TL, fresh.

Nezumia proxima (Smith and Radcliffe, 1912)
Figure 19.
Macrourus proximus Smith and Radcliffe in Radcliffe, 1912:119–120, pl. 26, fig. 2 (holotype, 292 mm TL,
USNM 72936; Sogod Bay, Leyte I., Philippines, 10°12ʹ00ʺN, 125°04ʹ10ʺE, 502 fm [918 m]; paratypes,
USNM 135338–39).
Macrourus nasutus [non Coryphaenoides nasutus Günther, 1877]: Jordan and Gilbert, 1904:618 (spec. from
off Izu, Japan).
Lionurus proximus: Gilbert and Hubbs, 1916:201–202 (11 spec. [excluding 3 types]; East China Sea, Pacific
coast of s. Japan, 361–544 fm [660–995 m]).
Lionurus (subgenus Nezumia) proximus: Gilbert and Hubbs, 1920:554 (3 spec. [including holotype], Philip-
pines).
Lionurus abei Matsubara, 1943:146–147, fig. 7, 8 (holotype, 160.7 mm TL, cat. no. 4909, Kumano-nada,
Japan; “Sigenkagaku Fish Coll. No. 7”).
Nezumia proximus: Okamura, 1970:94–98, pl. XXI, text-fig. 41(17 spec., 174–370 mm TL; e. coast s. Japan,
East China Sea; 420–910 m; synonymized Lionurus spinosus Gilbert and Hubbs, 1916 with N. proxima).
Nezumia proxima: Okamura in Okamura et al., 1982:163, 351, fig. 98 (p. 162) (1 spec., 160 mm TL; Kyushu-
Palau Ridge).— Sawada in Amaoka et al., 1983:107, 192, fig. 58 (p. 106) (4 spec., 269.6–346.4 mm TL;
Ibaragi [central Honshu, ca. 37°N], Japan, 1100 m).— Okamura in Masuda et al., 1984:95, pl. 81–I (com-
piled).— Okamura in Okamura and Kitajima, 1984:217, 362, fig. 152 (p. 216) (8 spec., 186–350 mm TL;
Okinawa Trough [East China Sea], 820–932 m).— Iwamoto, 1990:281–282, fig. 645–646 (compiled).—
Nakabo, 2002:424 (compiled).— Shao et al., 2008: table 2 (2 spec., Taiwan [ET, SCS], 488–1098 m; first
record from Taiwan).
MATERIAL EXAMINED (2 spec.).— NET: ASIZP 65632 (1, 377+ TL), CP 214, 488–1027 m.
SCS: ASIZP 66187 (1, 32 HL, 163+ TL); CD 322, 1098 m. SWT: ASIZP 65529 (1, 282+ TL), CD
142, 227–335 m.
DISTINGUISHING FEATURES.— 1D II,9–11; P i17–i21; V 9, rarely 10; inner GR-I 9–11 total;
scale rows below midbase 1D 5–7.5, below 2D 7–8.5; pyl. caeca 24–31. Snout length 23–31% HL;
interorbital 18–25%; orbit 28–33%; suborbital 14–17%; upper jaw 31–34%; barbel 15–23%. Body
relatively slender, compressed; head in TL about 6–7 times; snout conical, protruding well beyond
mouth, with stout tubercles at tip and lateral angles; suborbital ridge sharp and beset with stout
modified scales; underside of head mostly scaled except for broad naked median swath on under-
side of snout; body scales densely covered with recumbent narrowly triangular to lanceolate spin-
ules. Long spinous ray of 1D about 70–100% HL, serrations strong, sharp, widely spaced. Anteri-
or dermal window of light organ small, situated between inner V-fin bases; anus about midway
between V and A. Color overall dark brown, abdominal region bluish to black; gill cover, gill mem-
branes black; mouth dusky; fins dark dusky to black. Attains at least 380 mm TL.
DISTRIBUTION.— Broadly distributed from east-central Japan and East China Sea to South
China Sea off Taiwan and the Philippines; recorded depth range 355–1100 m. The sw. Taiwan spec-
imen was taken in 227–335 m; the ne. Taiwan specimen in 488–1027 m, and the SCS specimen in
1098 m.
REMARKS.— The 133 mm specimen listed in the table in Gilbert and Hubbs (1916:202) as
N. proxima from Albatross sta. 4918 is deposited at CAS [cat. no. CAS-SU 22941]; we have deter-
mined that it represents N. spinosa, a species that is closely similar to N. proxima and with which
it has been mistaken in the past. However, the underside of head of N. spinosa is almost wholly
naked, compared with only the median portion of snout in front of the mouth naked in N. proxima.
The spinous 1D ray is also longer in N. spinosa, the scale spinules needle-like and not flattened
(viz., not lanceolate or spear-shaped), and the outer premaxillary dentition is larger than in N. prox-
ima.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 93

Nezumia spinosa (Gilbert and Hubbs, 1916)
Figures 20A–B.
Lionurus spinosus Gilbert and Hubbs, 1916:199–200, pl. 10, fig.2 (holotype, USNM 76868, East China Sea
off Japan, 427 fm [781 m]).— Gilbert and Hubbs 1920:554 (4 spec., off Luzon, Philippines).— Okamu-
ra, 1970:94 (in part; in synonymy of Nezumia proximus).
Nezumia spinosa: Okamura in Masuda et al., 1984:95, pl. 345–A (compiled).— Iwamoto, 1990:254, fig. 582
(in key).— Iwamoto and Anderson, 1994:18–19 (22 spec., 26.3–44.2 HL; 117–255 mm TL, s. Africa and
Mozambique, 560–1000 m).— Iwamoto and Merrett, 1997:542–545, fig. 29b (8 spec., 20–54.2 HL,
133–319 TL; sw. Pacific, Japan, South China Sea, 660–900 m).— Iwamoto and Williams, 1999:
202–204, fig. 43b (8 spec., 33.3–51.1 HL, 140–260+ TL, Western Australia, 420–853 m).— Merrett and
Iwamoto, 2000:772–773 (1 spec., 45 HL, 250+ TL; Norfolk Ridge [sw. Pacific], 640–740 m).— Shao et
al., 2008: table 2 (3 spec., Taiwan [SWT, SCS], 316–720 m; first record from Taiwan).
MATERIAL EXAMINED (6 spec.).— SWT: ASIZP 65545 (1, 170 TL), CD 137, 316–477 m;
ASIZP 67586 (2, 105+-160+TL), CP 338, 569 m; ASIZP 67591 (1, 34 HL, 159+ TL); CP 338, 569
m. SCS: ASIZP 66747 (1, 216+ TL), CD 320, 731 m; ASIZP 66817(1, 127+ TL), CP 315, 509 m.
DISTINGUISHING FEATURES.— 1D II,9–10 (rarely 8 or 12); P i18–i22; V 8–9; inner GR-I 9–12
total; scale rows below midbase 1D 6.5–9, below 2D 7.5–11; lateral line scales over distance equal
to predorsal length 35–42; pyl. caeca 15–16 (13–14 fide Iwamoto and Anderson 1994:18). Snout
94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 19. Nezurnia proxima (Smith and Radcliffe, 1912), ASIZP 66187, 163+ mm TL, fresh (A) and preserved (B).
A
B

length 28–32% HL; preoral length 18–29%; interorbital 18–24%; orbit 29–31%; orbit to preoper-
cle 34–40%; suborbital 12–14%; upper jaw 27–33%; barbel 9–18%. Body relatively slender, com-
pressed; head about 5–6 in TL; snout conical, protruding well beyond mouth, with stout tubercles
at tip and lateral angles; suborbital ridge sharp and beset with stout modified scales; underside of
head almost completely naked, with large pores; body scales densely covered with long, needle-
like spinules in parallel to convergent rows. Long spinous ray of 1D 96–156% HL, serrations wide-
ly spaced. Anterior dermal window of light organ prominent, situated between V-fin bases. Color
overall light brown, blackish over abdomen; gill cover, gill membranes dark; mouth and gill cavi-
ties black; 1D blackish proximally, pale to whitish distally, V black, P and A dusky. Attains about
320 mm TL.
DISTRIBUTION.— A widespread Indo-Pacific species; s. Japan through Philippines, Indonesia,
s. to New Caledonia region, w. coast of Australia, and s. Africa, in 316–1000 m. Our Taiwan spec-
imens were taken in 316–731 m.
REMARKS.— Shao et al. (2008) first recorded the species from Taiwan based on the current
specimens.
Nezumia cf. coheni [sensu Iwamoto and Merrett, 1997]
Figure 21A–C.
Nezumia coheni (not of Iwamoto and Merrett, 1997): Shao et al., 2008: table 2 (3 spec., SWT, 1305 m; first
record from Taiwan).
MATERIAL EXAMINED (3 spec.).— SWT: ASIZP 63791 (3, 290+-333+ TL), CD 192, 1305 m.
DISTINGUISHING FEATURES.— 1D II,9–11; P i19–i21; V9–10; GR-I (inner) 2+7, total GR-II
(outer /inner) 8–9 / 9–10; scale rows below 2D origin 7.0–8.5, under midbase of 1D 6.5; lateral-
line scales over distance equal to predorsal length 33–35. Snout conical with smoothly rounded
dorsal and ventral profiles, length 31–33% HL, interorbital flat, its width 22–24%, orbit 27–32%,
suborbital almost vertical, gently rounded, depth 17–19%; upper jaw 33–35%; barbel 15–20%;
outer gill slit 12–16%; pre-A length 165–186%; outer V to A origin 44–50%; body depth about
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 95
FIGURE 20. Nezumia spinosa (Gilbert and Hubbs, 1916). A. ASIZP 67586, 105 mm TL, fresh. B. ASIZP66817, 127+
mm TL, preserved.
A
B

77–92%; 1D-2D interspace 38–52%; height 1D 94–98%; length 1D base 24–30%; length P
51–61%; length outer V 49–54%. Underside of snout mostly naked; underside of head becoming
increasingly covered with scales ventrally and posteriorly from about vertical through nasal fossa;
pores and sensory papillae of cephalic lateralis system densely arrayed on underside of head, espe-
cially under snout and mandibles, producing a roughened surface texture; scales densely covered
with broadly lanceolate spinules. All fins dark, blackish; dermal window of light organ relatively
small and located between inner margins of V bases. Color in alcohol overall dark brownish;
abdomen, gill covers, and gill membranes blackish; mouth dark but somewhat pale along outer
margins; gums pale. Attains at least 333 mm TL.
DISTRIBUTION.— Known from the South China Sea off the sw. coast of Taiwan in 1305 m.
REMARKS.— Shao et al. (2008) first recorded these specimens as Nezumia coheni Iwamoto
and Merrett, 1997, but on closer examination, certain character disparities suggested that the Tai-
wan specimens may represent a different species. The pelvic fin-ray count of 9 or 10 contrasts with
96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 21. Nezumia cf. coheni. A. ASIZP 63791, 1 of 3, 333+ mm TL, fresh. B-C. ASIZP 63791, 1 of 3, 290+ mm TK.
B. lateral view. C. lateral view of left head.
B
A
C

the usually 11 (rarely 9 or 10) of N. coheni. Certain proportional measurements differed slightly:
suborbital depth 17–19% HL cf. 15–16% in N. coheni; preanal length 165–186% cf 143–166%;
and barbel length into orbit diameter 1.3–1.6 times cf. 1.6–2.0 times. The spinules on body scales
also have rather broadly convex tips in the Taiwan specimens compared with the more attenuated
and sharp tips of N. coheni. Although the three Taiwan specimens are in excellent condition, they
were all collected together and are all of relatively the same size and likely do not show the range
of variation in the species. The similarities in most features suggest a close relationship between
N. coheni and these Taiwan specimens, if indeed they are different. We must await collection and
study of more specimens of different sizes to confirm or reject the notion of their being different
species.
Genus Pseudocetonurus Sazonov and Shcherbachev, 1982
Pseudocetonurus Sazonov and Shcherbachev, 1982:5–6 (type species Pseudocetonurus septifer Sazonov and
Shcherbachev, 1982 by monotypy).
DISTINGUISHING FEATURES.— BR 7; V 9 or 10, rarely 8; inner GR-I 13–18 total; pyl. caeca
22–34. Head enlarged, notably broad and deep, preopercle and suborbital bones deep and large;
orbit small in adults, proportionally larger in young, diameter 19–31% HL; interorbital width
33–44% HL. Snout high, bluntly pointed, little protruding beyond mouth; mouth large, upper jaw
41–49% HL; chin barbel small, 7–10%; gill opening wide, extending ventrally to below end of
maxilla. Spinous ray of 1D smooth proximally, finely serrated distally; abdominal area short,
periproct between V and A, closer to V bases. Light organ developed: a small round dermal win-
dow between V bases. Head fully scaled except for gular and gill membranes; small, awl-shaped
spinules on body scales, no reticulations on anterior field; grooved lateral line scales over trunk
present or absent. Teeth small, in narrow band on premaxillary, in single series in dentary. Ground
color black to dark brown overall.
REMARKS.— This genus is closely similar to Ventrifossa and would likely be categorized in
that taxon except for its very dark color, greatly expanded head, and commensurately expanded
bones of the opercular series. Only the single species known.
Pseudocetonurus cf. septifer [sensu Sazonov and Shcherbachev, 1982]
Pseudocetonurus septifer: Chiou et al., 2004b:46, fig. 17 (1 spec., ASIZP 61237, 147 mm TL; Da-xi, Taiwan
[NET]).— Shao et al., 2008: table 2 (3 spec., Taiwan [NET]).
MATERIAL EXAMINED (1 spec.).— NET: ASIZP 61237 (1, 147 TL), Da-xi.
DISTINGUISHING FEATURES.— 1D II,8–12, usually 9 or 10; P i16–i20; GR-I (outer/inner)
7–12/13–18, GR-II 14–18/14–17. Snout length 25–34% HL; preoral length 19–27%; postorbital
47–59%; orbit to preopercle 53–64%; suborbital 19–26%; outer gill slit 23–31; body depth 75–95;
V-A 12–38%; height 1D 52–66%; length V 41–63%. Head about 5.0–5.5 in TL; greatest width
about 1.5 into greatest depth; interorbital about 2.3–3.0 into HL; suborbital deep, almost vertical
and lacking modified scutelike scales; Mouth large, upper jaw extends to below midorbit. Barbel
almost rudimentary, <0.5 orbit. Sensory canals of head greatly expanded, but sensory pores small;
free neuromasts poorly developed. Grooved lateral line present.
DISTRIBUTIONs.— So far known only from the nw. Pacific off Taiwan.
REMARKS.— Based on the presence of a grooved lateral line, this specimen is probably an
undescribed species. The only other member of the genus, P. septifer Sazonov and Shcherbachev,
1982, known from the Saly-Gomez and Nazca ridges in the southeastern Pacific and off Hawaii,
lacks a grooved lateral line.The Taiwan species was first recorded as P. septifer by Chiou et al.
(2004b) and subsequently by Shao et al. (2008).
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 97

Genus Pseudonezumia Okamura, 1970
Pseudonezumia Okamura, 1970:38–39 (type-species Pseudonezumia japonicus Okamura, 1970 by original
designation).
Paracetonurus Marshall, 1973:615–616 (type-species Macrourus parvipes Smith and Radcliffe, 1912 by orig-
inal designation).
DISTINGUISHING FEATURES.— BR 7 or 8; V 6, rarely 5 or 7, situated below posterior end of
head; P 16–18; anus slightly removed from A; head much deeper than wide (P. japonica) to rela-
tively broad and deep (other spp.); sensory canals expanded, well developed; head and body almost
fully scaled; body scales small, exposed field with erect awl-shaped spinules in subparallel to
slightly divergent rows, reticulations on scales; suborbital ridge without modified scales; snout
broad, tip and lateral angles lacking tubercles; teeth in narrow bands, somewhat enlarged in outer
premaxillary and inner dentary series. No apparent light organ developed.
REMARKS.— We are treating the genus Paracetonurus Marshall, 1973 as a synonym, although
this has not been adequately confirmed. Sazonov and Shcherbachev (1982:11) provided a good
diagnosis and discussion of relationships of the genus (as Paracetonurus); they listed four species
as belonging to the genus, including P. flagellicauda (Koefoed, 1927), P. parvipes (Smith and Rad-
cliffe, 1912), and P. cetonuropsis (Gilbert and Hubbs, 1916), and their new one P. pusillus, but not
P. japonica Okamura, 1970. Of the five species, only one is known from Taiwan.
Pseudonezumia pusilla (Sazonov and Shcherbachev, 1982)
Figures 22A–B.
Paracetonurus pusillus Sazonov and Shcherbachev, 1982:12–14, fig. 4 (holotype, ZMMU P.15306, Indian
Ocean, Ninety-East Ridge, 11°24.3ʹS, 88°50ʹE, 1500–1600 m; 17 paratypes, Indian Ocean and w. Pacif-
ic off New Guinea; 1380–2000 m)
Paracetonurus cetonuropsis (non Gilbert and Hubbs, 1916): Shao et al., 2008: table 2 (3 spec., Taiwan [SCS],
1098 m).
Pseudonezumia pusilla: Iwamoto and Williams, 1999:208–209, fig. 45 (1 spec., off Western Australia, w. of
NW Cape; 1460–1500 m).
MATERIAL EXAMINED (5 spec.).— SCS: ASIZP 66945 (3, 137–163 TL) and ASIZP66424 (1,
234 TL), CD 322, 1098 m. Other material: CAS 224176 (1, 28.6 HL; 175+ TL), SCS, off Viet-
nam, 15°38ʹN, 111°54ʹE; MV Stranger, NAGA Exped., sta. 60–67, GVF Reg. no. 2083, 28 Feb.
1960.
DISTINGUISHING FEATURES.— 1D II,8–9; P 17–18; V 6 or rarely 7; total GR-I (outer/inner)
10–12/10–11; GR-II 10–12/10–11; pyl. caeca 7–12. Snout length 29–32% HL; preoral length
21–22%; internasal width 23–26%; interorbital 30–33%; orbit 29–31%; posterior nostril 12–16%,
2.2–2.7 times into orbit; suborbital 14–18%; postorbital 41–46%; orbit to preop. 35–42%; upper
jaw 34–40%; barbel 12–20%; outer gill-slit 12–20%; pre-A 122–138%; isth. to A 57–72%; outer
V to A 35–52%; 1D-2D 22–46%; length base 1D 21–29%. Head relatively compressed, deeper than
wide; snout pointed, rather broad, internasal width about 1⁄4HL; interorbital 1⁄3HL, slightly greater
than orbit diameter; suborbital about half orbit; barbel relatively long, slender. Margin of preoper-
cle broadly rounded, almost vertical posteriorly, preopercle ridge rounded at angle. Origin of V
under posterior margin of preopercle; vent slightly removed from origin of A, under anterior 1⁄4of
1D. Posterior end of upper jaw below mid-orbit. Scale spinules 2–9, awl-shaped, erect to slightly
reclined. Periproct small, slightly removed from A origin. Paired fins small, weak; 1D spine with
weak, widely spaced denticles. (In part from Sazonov and Shcherbachev 1982:12–13.)
DISTRIBUTION.— Known from the holotype and 17 paratypes taken in the Indian Ocean and
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w. Pacific off New Guinea (Bismark Sea); one specimen from off the NW Cape of Australia; and
the current Taiwan specimens from the South China Sea, which extend the range of the species into
the w. North Pacific. Depth range 1098–2000 m; the Taiwan specimens represent the shallowest
recorded at 1098 m.
REMARKS.— Shao et al. (2008: table 2) first recorded the Taiwan specimens as Pseudonezu-
mia cetonuropsis, a species known only from the holotype and small paratype taken off central
Japan. That species differs from P. pusilla in having a much shorter barbel (6% HL), longer snout
(35%), and smaller orbit (27%).
Genus Sphagemacrurus Fowler, 1925
DISTINGUISHING FEATURES.— BR 7; ventral region of short head and short trunk shifted for-
ward, gill membranes united under orbit, V usually under opercle, anus under origin of 1D, A ori-
gin under hind margin of 1D; snout blunt, high, tip of snout with small, spiny tubercle, on level
with upper margin of pupil, naked areas, if present, confined to lower surfaces; jaws oblique; 1D
spine with saw-tooth serrations on leading edge, 1D base somewhat elevated; naked periproct
region broad, occupying 1⁄3or more of space between V and A, usually immediately before A ori-
gin; inner GR-I 15 or fewer; length barbel 1.7 to > 2.0 into orbit diameter. A well-developed sub-
orbital shelf composed of two parallel rows of stout, spiny, modified scales. Body scales with 5–9
subparallel rows of short, awl-shaped, slightly reclined spinules.
REMARKS.— A small genus of six species, two of which are recorded from Taiwan. Iwamoto
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 99
FIGURE 22. Pseudonezumia pusilla (Sazonov, and Shcherbachev, 1982). ASIZP66424, 234 mm TL. A. fresh. B. lateral
view of left head, preserved.
A
B

and Williams (1999:211–212) noted differences in recorded values and features of specimens iden-
tified as S. pumiliceps from the Indian and Pacific oceans and stated that there are unresolved prob-
lems with the species in the genus. There is overlap in many characters between the Sphagemacru-
rus specimens we have examined, making identification difficult. The genus is in need of further
study.
Key to the Species of Sphagemacrurus in Taiwan
1a. Rays of V 11–14; inner GR-I 8–11; barbel length 15–24% HL; interorbital width 23–28%. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pumiliceps
1b. Rays of V 9–11; inner GR-I 11–13; barbel length 13–16% HL; interorbital width 28–32% . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. richardi
Sphagemacrurus pumiliceps (Alcock 1894)
Figures 23A–B.
Macrurus pumiliceps Alcock 1894:125 (Laccadive Sea; 1,315 m).
Lionurus pumiliceps: Gilbert and Hubbs, 1920:559, 560 (Philippine and East Indies, 732–1646 m).
Sphagemacrurus pumiliceps: Iwamoto, 1990:288 (in key).— Iwamoto and Merrett, 1997:549 (28 spec., sw.
Pacific).— Iwamoto and Williams, 1999:210 (descr., 4 spec., Western Australia, 882–1880 m).— Merrett
and Iwamoto, 2000:773 (24 spec., sw. Pacific).— Iwamoto et al., 2004:199 (1 spec., Walters Shoals, sw.
Indian Ocean, 1310–1265 m).— Shao et al., 2008: table 2 (listed; Taiwan [ET, SCS], 736–1188 m).
MATERIAL EXAMINED (6 spec.).— NET: ASIZP 65551 (1, 36.9 HL, 265+ TL), CD 199,
1134–1188 m. SCS: ASIZP 65558 (1, 26.6 HL, 140+ TL), CD 134, 736–1040 m; ASIZP 66905 (3,
23.8–29.0 HL, 112+-202+ TL), CD 322, 1098 m.
DISTINGUISHING FEATURES.— 1D II,10–11; P i20–i23; V 11–14; inner GR-I 8–11 total; scale
rows below 2D 7–10; pyl. caeca 10–12. Snout length 29–37% HL; interorbital 23–27%; orbit
31–37%; suborbital 18–21%; postorbital 40–44%; orbit-preopercle 36–43; upper jaw 36–39%; bar-
bel 17–20%. Body relatively deep, about equal to HL; head about 7.5 in TL [all our specimens have
a broken tail]; mouth rather small, upper jaw extends posteriorly to under midorbit; barbel well
developed, about half diameter of orbit; underside of head mostly scaled except ventrally on snout
and suborbital margin; body scales with short fine spinules arranged in about 5–8 parallel rows.
Outer ray of V slightly prolonged, its tip extending to A origin or substantially beyond (to as far as
10th A ray). Periproct broad, abutting A origin posteriorly, spanning about 2⁄3distance between V and
A. Attains about 27 cm TL. (Data from Taiwan specimens only.)
DISTRIBUTION.— Widely distributed in the w. Pacific from se. Australia to Taiwan, in
732–1880 m; also found through most of Indian Ocean, where species was originally described.
The ASIZ specimens, captured in 736–1188 m off Taiwan, were the first recorded from the area
(Shao et al., 2008) and extend the range to n. of the Philippines.
REMARKS.— The reportedly wide distribution of this species and the variation in certain fea-
tures found by Iwamoto and Williams (2001:211–212) suggest that more than one species may be
currently recognized under this name.
Sphagemacrurus richardi (Weber, 1913)
Figures 24A–B.
Macrurus richardi Weber, 1913:154 (28 syntypes: FMNH 52442 ex cm 700 [1 spec.] Flores Sea; MOM
0091–1766 [1, disintegrated]; ZMA 110456 [1], 110447–49 [18, 1, 3], 110464 [1]; Makassar Strait;
Celebes [Sulawesi] Sea; Ceram Sea; Flores Sea; Siboga sta. 85, 122, 170, 314, and 316; 538–1260 m).
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Sphagemacrurus richardi: Iwamoto, 1990:288, fig. 658 (in key).— Iwamoto and Williams, 1999:212 (men-
tioned).— Iwamoto and Graham, 2001:493 (56 spec., se. Australia; 880–1100m).
MATERIAL EXAMINED (6 spec.).— SCS: ASIZP 65621 (1, 29.3 HL, 176 TL), CD 203; ASIZP
66087 (1, 32.7 HL, 165+ TL), CD 320; ASIZP 66184 (1, 35.5 HL, 186+ TL), CD 320; ASIZP
66192 (1, 32.9 HL, 176 TL), CD 320; ASIZP 66910 (1, 34.5 HL, 190+ TL), CD 321; ASIZP 66944
(1, 127 TL), CD 321.
DISTINGUISHING FEATURES.— 1D II,9–10; P i19–i21; V 10–11(12); inner GR-I 11–14 total;
scale rows below 2D 8–9; pyl. caeca about 10. Snout length 35–37% HL; interorbital 30–34%;
orbit 28–34%; suborbital 19–21%; postorbital 40–43%; orbit-preopercle 40–41%; upper jaw
37–39%; barbel 13–16%. Body relatively deep, about equal to HL; head about 5.3–6.0 in TL;
mouth rather small, upper jaw extends posteriorly to under midorbit; barbel well developed, about
half orbit diameter; underside of head mostly scaled except ventrally on snout and suborbital mar-
gin; body scales with short fine spinules arranged in about 7–9 parallel rows. Outer ray of V slight-
ly prolonged, its tip extending to or somewhat beyond A origin. Periproct broad, abutting A origin
posteriorly, spanning about 2⁄3distance between V and A; anus closer to A than to outer base of V.
Attains about 23 cm TL. (Data from Taiwan specimens only.)
DISTRIBUTION.— The ASIZ specimens captured in 634–954 m off Taiwan were the first
recorded from the area (Shao et al. 2008) and extend the range to northward from Indonesia where
it was first described.
REMARKS.— Our specimens represent the first record for Taiwan. In his original description
of the species, Weber (1913) gave 8 as the count of V rays, but one of the current author’s (TI)
examination of what was probably the illustrated syntype (ZMA 110.456) had 10 rays in each fin.
No other specimen of the species that we know of had such a low count, and we suspect it to be
erroneous. Sphagemacrurus decimalis (Gilbert and Hubbs, 1920) has a low V count of 10, similar
to S. richardi, but it differs in a number of morphometric features (as enumerated in the original
description), including a somewhat lower count of rakers on the lower limb of GR-I (7 cf. 9–12);
shorter snout (26–28% HL cf. 35–37%); shorter upper jaw (28–29% cf. 40–41%; shorter orbit-pre-
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 101
FIGURE 23. Sphagemacrurus pumiliceps (Alcock, 1894). A. ASIZP 66905, 198 mm TL, fresh. B. ASIZP 65551, 265+
mm TL, preserved.
A
B

opercle distance 33–34% cf. 40–41%; and shorter barbel (9–11% cf. 13–16%). Sphagemacrurus
richardi differs from S. pumiliceps in having slightly fewer V rays (10–11, rarely 12, vs. 11–14),
shorter barbel (13–16% vs. 16–21%), and somewhat wider interorbital (30–34% vs. 24–28%) (S.
pumiliceps data from Iwamoto and Graham 2001:493).
Genus Spicomacrurus Okamura, 1970
DISTINGUISHING FEATURES.— Snout low, pointed and slightly protruding, median and lateral
processes of nasal bone forming broad horizontal plates; head about as broad as high; body and
head cylindrical; head mucous canals moderately developed; head covering mostly transparent;
light organ long, small lens on chest anterior to pelvic-fin bases connected by a black streak to elon-
gated posterior lens immediately before anus; ventral striae well developed; inner GR-I, lower
limb, 10–12; spinous ray of 1D completely smooth; chin barbel present; gular region with netlike
epidermal cover; outer pelvic ray with narrow membranous flange that is somewhat expanded dis-
tally near tip.
REMARKS.— Spicomacrurus was originally treated as a subgenus of Hymenogadus, but
Iwamoto et al. (2011) elevated the subgenus to full generic status. Although obviously related to
members of the genera Hymenocephalus and Hymenogadus, the four species of Spicomacrurus are
readily differentiated from them by the broad horizontal plates of their nasal bones, elongated pos-
terior lens of the light organ, netlike gular covering, and membranous flange of the outer pelvic fin
ray. Spicomacrurus kuronumai is the only member of the genus found in Taiwan.
102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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A
B
FIGURE 24. Sphagemacrurus richardi (Weber, 1913). A. ASIZP 66910, 190+ mm TL, fresh. B. ASIZP 65621, 176 mm
TL, preserved, photo reversed laterally.

Spicomacrurus kuronumai (Kamohara, 1938 )
Hymenocephalus kuronumai Kamohara, 1938:70, fig. 40 (Mimase, Kochi Pref., Japan; neotype: BSKU 4333,
invalid according to ICZN).— Shao et al., 2008: table 2 (1 spec., NET, 100–650 m; first record from Tai-
wan).
Hymenogadus (Spicomacrurus) kuronumai: Okamura, 1970:64–67, pl. III, text-figs. 25, 26–28 (79 spec.,
143–200 mm TL; Pacific coast of s. Japan, in about 350–450 m).
Hymenogadus kuronumai: Okamura in Masuda et al., 1984:94, pl. 80–I (compiled). Okamura in Okamura and
Kitajima, 1984:203, 359, fig. 144 (p.202) (3 spec., 135–194 mm TL, Okinawa Trough [East China Sea],
400–510 m.
Spicomacrurus kuronumai: Iwamoto et al., 2011:513–530, fig. 1A–C (subgenus elevated to genus).
MATERIAL EXAMINED (2 spec.).— NET: ASIZP 65232 (1, 165 TL), Da-xi. ASIZP 70247 (1,
97 TL), Da-xi.
DISTINGUISHING FEATURES.— 1D II 9–12; P i18–i22; V 8; inner GR-I 12–15 total; pyl. caeca
11–18. Snout depressed, nasal bones developed into three horizontal platelike processes; body sub-
cylindrical; outer V ray produced and expanded distally. Attains 240 mm TL.
DISTRIBUTION.— Pacific coast of s. Japan, and ne. Taiwan. in 350–650 m.
REMARKS.— Shao et al., (2008: table 2) first recorded the species from Taiwan based on
ASIZP 65232. The species is apparently relatively abundant off Japan, as suggested by the 95 spec-
imens (143–211 mm TL) recorded by Okamura (1970b:4, table 1), who treated it as a subgenus of
Hymenogadus. Kamohara (1961) designated a neotype for the species but not in a revisory work,
which renders the designation invalid (fide Eschmeyer 1998:852).
Genus Trachonurus Günther, 1887
DISTINGUISHING FEATURES.— BR 7; origin of V well behind P base and under posterior one-
quarter of 1D base; V-A distance short, less than orbit diameter; broad periproct region spanning
most of distance between V and A; second spinous ray of 1D smooth, rounded in cross-section,
slightly longer than adjacent segmented ray; head smoothly rounded, without sharp ridges, almost
fully scaled; scale patches usually present on branchiostegals; chin barbel short; body scales
strongly adherent, with short, conical, relatively erect spinules; V 6 or 7; color uniformly gray to
dark brownish black, fins dark, usually black.
REMARKS.— At least six species, two of which are found off Taiwan. The species in this genus
are closely similar and differences between them are difficult to find. The widespread distributions
reported for certain species (especially T. villosus and T. sentipellis) beg closer scrutiny.
Key to the Species of Trachonurus in Taiwan
1a. Body scales relatively large and coarsely spinulated; 8 or 9 scale rows between V base and gill
cover. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. sentipellis
1b. Body scales smaller, finer spinulated; about 10–14 scale rows between V base and gill cover
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. villosus
Trachonurus sentipellis Gilbert and Cramer, 1897
Figures 25A–C.
Trachonurus sentipellis Gilbert and Cramer, 1897:429 (syntypes SU 3140 and USNM 51689; Hawaiian Is.,
21°12ʹ00ʺN, 157°38ʹ30ʺW, Albatross sta. 3474, 375 fm [686 m]).— Iwamoto, 1997:945.— Iwamoto and
Merrett, 1997:551.— Iwamoto and Williams, 1999:213 (descr., w. and se. Australia).— Merrett and
Iwamoto, 2000:775 (6 spec., New Caledonia, 764–1,050 m).— Iwamoto and Graham, 2001:495 (3 spec.,
se Australia, 940–1130 m).— Shao et al., 2008: table 2 (7 spec., Taiwan [SWT, SCS], 441–1040 m).
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 103

MATERIAL EXAMINED (10 spec.).— SWT: ASIZP 64232 (2, 54.6–61.0 HL, 265+-295+ TL),
CD 193, 821 m; ASIZP 65539 (1, 59.9 HL, 305 TL), CD 134, 736–1040 m; ASIZP 65543 (1, 150
TL), CD 138, 441 m; ASIZP 65544 (1, 175 TL) and ASIZP 65548 (1, 135 TL), CD 203, 634–866
m; ASIZP 65637 (1, 310+ TL), CD 229, 880–1062 m. SCS: ASIZP 66150 (1, 49 HL, 250+ TL),
CD 321, 954 m; ASIZP 66743 (2, 21–32 HL, 105+-170+ TL), CD 320, 731 m.
DISTINGUISHING FEATURES.— 1D II,6–8; P 15–16 (rarely 18); V 7; total GR-I (inner) 11–13;
scale rows below origin 2D usually 6–7, below midbase 1D 5–7; lateral line scales over distance
equal to pre-1D length 26–34; scale rows between V base and gill cover 8–9; pyl. caeca 9–13,
short, thick. Snout length 23–26% HL; interorbital width 26–33%; orbit diameter 27–37%; subor-
bital width 8–12; postorbital length 38–53%; orbit-preopercle 26–33; upper jaw 28–37%; barbel
8–11%; 1D-2D interspace 11–29%. Grooved lateral line present; body scales relatively large,
coarsely covered with stout, erect spinules. Jaw teeth all small. Attains more than 31 cm TL.
(Adapted from Iwamoto 1997.)
DISTRIBUTION.— Hawaii, Australia, New Caledonia, and Taiwan in 441–1130 m. Specimens
from Taiwan were all taken from the South China Sea in 441–1040 m; they represent the first
record of the species from the w. North Pacific.
REMARKS.— Trachonurus sentipellis is closely similar to T. robinsi Iwamoto, 1997 from the
Philippines, but that species shows differences in some meristic values: P rays 10–14 vs. 15–16;
inner GR-I 9–11 vs. 11–13; scales below 2D origin 4–6 vs. 5–8 (usually 6–7). Also, where T. sen-
tipellis almost invariably has V 7, in T. robinsi the count is more often 6 than 7, and the maximum
size in T. robinsi probably does not exceed 25 cm, whereas T. sentipellis exceeds 31 cm.
Trachonurus villosus (Günther, 1877)
Figure 26.
Coryphaenoides villosus Günther, 1877:441 (holotype BMNH 1887.12.7.105; s. of Tokyo, Japan).— Günther,
1887:142 (second specimen BMNH 1887.12.7.106 [non-type] from s. of the Philippines added).
Trachonurus villosus: Okamura in Masuda et al., 1984:95 (compiled).— Okamura in Amaoka et al.,
1990:193.— Iwamoto, 1997:944–947 (comparison with T. robinsi).— Iwamoto and Williams, 1999:212
(mentioned).— Shao et al., 2008: table 2 (5 spec.; Taiwan [SCS], 634–954 m).
MATERIAL EXAMINED (4 spec.).— SWT: ASIZP 65626 (2, 175–255+ TL), CD 203, 634–866
m. SCS: ASIZP 66094 (1, 60 HL, 345+ TL) and ASIZP 66909 (1, 155+ TL), CD 321, 954 m.
DISTINGUISHING FEATURES.— 1D II,8; P 14–17; V 7; total GR-I (inner) 9–13; scale rows
below origin 2D 5–9, usually 6–7, below midbase 1D 5–7; lateral line scales over distance equal
to pre-1D length 26–34; scale rows between base of V and gill cover 10–14; pyl. caeca 9–13, short,
thick. Grooved lateral line present; body scales relatively small, finely covered with erect spinules.
Jaw teeth all small. Snout length 23–27% HL; interorbital width 31–34%; orbit diameter 29–30%;
suborbital width 11–12%; postorbital length 38–42%; orbit-preopercle 29–33%; upper jaw
30–33%; barbel 6+–11%; 1D-2D interspace 20–22%. (Adapted from Iwamoto 1997.)
DISTRIBUTION.— Known from Japan to Taiwan and the Philippines. In Taiwan, specimens
were taken from the South China Sea in 634–954 m.
REMARKS.— More specimens of this species must be examined from near the type locality off
Japan to properly circumscribe the species and determine its difference from other described
species.
104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3

Genus Ventrifossa Gilbert and Hubbs, 1920
DISTINGUISHING FEATURES.— BR 7. Head and body relatively compressed; gill membranes
narrowly united over isthmus and under midorbit; snout blunt to pointed and moderately protrud-
ing beyond mouth; tubercular scale lacking on snout tip except in V. misakia; no thickened modi-
fied scales on suborbital region; head uniformly and fully scaled except gular and BR membranes;
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 105
FIGURE 25. Trachonurus sentipellis Gilbert and Cramer, 1897. A. ASIZP 64232, 295+ mm TL. B. ASIZP 65539, 305
mm TL. C. ASIZP 65539, lateral view of right head, photo reversed laterally.
A
C
B

upper jaws generally more than 40% HL (35–42% in V. misakia), beset with long band of small
teeth, outer series slightly enlarged; teeth on lower jaw all small, in one or two series to long nar-
row band; chin barbel usually well developed. Second spinous 1D ray smooth or finely serrated;
V 8–10; no fin with greatly prolonged rays. Inner series GR-I 14–20 total. Periproct oval to tear-
drop shaped, connected anteriorly to small dermal window of light organ lying between V bases;
anus much closer to V bases than to A origin. Pyloric caeca more than 30. Color often silvery along
sides; lips usually black; leading edge of snout, suborbital shelf, dorsal snout ridges, gill and gular
membranes usually black or blackish. (Adapted from Iwamoto and Graham, 2001:496.)
REMARKS.— There are more than 25 species in this genus; nine are here recorded from Tai-
wan. There are likely to be other species still undiscovered, especially in Indonesia and the Indian
Ocean. Because of the often fragile nature of their integument and bones, smaller individuals of the
genus are often severely damaged in capture. Features distinguishing species of Ventrifossa are
often subtle, requiring relatively intact specimens, which add to the difficulties in their identifica-
tion.
106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 26. Trachonurus villosus (Günther, 1877). ASIZP 66909, 155+ mm TL. A. fresh. B. lateral view, preserved.
C. dorsal view of head.
A
C
B

Key to the Species of Ventrifossa in Taiwan
1a. Second spinous ray of 1D smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. Second spinous ray of 1D finely serrated along leading edge . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Pectoral fin long, 1.3–1.5 into HL; no enlarged spinules on scales along 2D base . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. macroptera
2b. Pectoral fin moderate to short, about 1.7 into HL; scales along 2D base with enlarged spinules
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. sazonovi
3a. Scales extremely small and finely spinulated, scaled surfaces almost velvety when stroked fore
and aft; scale rows below 2D origin 9–10 a small tubercular scale at snout tip; suborbital shelf
greatly constricted below anterior quarter of orbit . . . . . . . . . . . . . . . . . . . . . . . . . . V. misakia
3b. Scales moderate in size with spinules of variable size, skin surface not velvety smooth; scale
rows below 2D origin 5–8; no terminal snout scute; suborbital shelf not greatly constricted
anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Spinules on body scales broad, short, and triangular (V 8–9, usually 8; 1D dusky to clear, with-
out a distinct black blotch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. garmani
4b. Spinules on body scales narrowly lanceolate to conical (V 8–10; 1D dusky to blackish, some
species with a black blotch). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Body color uniformly brown except swarthy over abdomen (underlain by bluish) and no silvery
reflections, suborbital uniformly dark brown, no black shelf; snout rather blunt in lateral view,
scarcely protruding beyond mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. saikaiensis
5b. Body color light grayish-brown dorsally, silvery along sides, suborbital with black shelf; snout
moderately pointed to rather blunt and prominently to scarcely protruding beyond mouth. . 6
6a. Barbel less than 24% of HL; lateral-line scale rows 43 or less; 1D with prominent black blotch
or broad midlateral streak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 b. Barbel 25% or more of HL; lateral-line scale rows 45 or more; 1D uniformly dusky or with
midlateral black streak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7a. Rays of V 9 or 10; mesial GR-I 15–17 total, lateral GR-II 14–17 total; length upper jaw usual-
ly 43–46% HL; black distal margin anteriorly on A fin . . . . . . . . . . . . . . . . V. rhipidodorsalis
7b. Rays of V 8 or 9; mesial GR-I 13–15 total, lateral GR-II 12–14 total; length upper jaw 40–43%
HL; no black margin on A fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. nigrodorsalis
8a. 1D usually uniformly dusky or somewhat darker proximally. . . . . . . . . . . . . . . . . V. divergens
8b. 1D dark or black midlaterally, distally pale or dusky . . . . . . . . . . . . . . . . . . . . V. longibarbata
Ventrifossa divergens Gilbert and Hubbs, 1920
Figure 27.
Ventrifossa divergens Gilbert and Hubbs, 1920:549 (holotype, USNM 78230, Celebes Sea near Sibuko Bay,
Borneo, 4°12ʹ44ʺN, 118°27ʹ44ʺE, Albatross sta. 5592, 305 fm [558 m]; 59 paratypes).— Iwamoto, 1990:
299–300, fig. 676–677 (descr.).
MATERIAL EXAMINED (21 spec.).— SWT: ASIZP 66928 (1, 62 HL, 320+ TL), PCP 339, 846
m; ASIZP 66255 (1, 67 TL), OCP 303, 807 m. SCS: ASIZP 66781 (4, 59–64 HL, 297+-330+ TL),
and ASIZP 66782 (2, 58–59 HL, 275+-310+ TL), CD 320, 731 m; ASIZP 66784 (2, 41–47 HL,
205+-217+ TL), OCP 317, 515 m. Other specimens (all from Albatross Philippines Expedition of
1907–1909): Indonesia: USNM 78230 (holotype, 48.4 mm HL, 271 mm TL). USNM 122917
(paratypes, 3, 34.2–40.3 HL), Molucca Sea, 545 m, sta. 5621. Philippines: USNM 122916
(paratypes, 2, 142+-469+ TL), off ne. tip of Luzon, 410 m, sta. 5325; USNM 122919 (41.5 HL,
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 107

222 TL), off ne. Mindoro, 623 m. USNM 148988 (paratype, 53.0 HL, 281 TL), off Balayan Bay,
Batangas, sw. Luzon, sta. 5116 or 5115. USNM 149361 (paratype, 24.7 HL, 140 TL), off sw.
Luzon, 315 m, sta. 5289. USNM 149363 (paratype, 53.2 HL, 250 TL) and USNM 150400
(paratype, 19.5 HL, 105 TL), off sw. Luzon, 391 m, sta. 5290.
DISTINGUISHING FEATURES (from 11 USNM specimens, holotype data with asterisk*).— 1D
II,9–11(10*); P i18–i25 (21–22*); V 8*–9; total GR-I (mesial) (12*)15–18, GR-II (lat.)
14*–15(17); scale rows below 1D 10–12, below midbase 1D (10*), below 2D 8–9* (11); lateral
line scales over distance equal to predorsal length (48*). Snout length (29*) 26–31% HL; preoral
length (15*) 14–19%; internasal 19*–25%; interorbital (24*) 30–37%; orbit (31*) 30–38%; subor-
bital (11*) 13–15%; postorbital (44*) (38) 41–47%; orbit to preopercle (45*) 40–46%; upper jaw
43*–49%; barbel (28*) 26–35%; outer gill slit (19*) 24–28%. Color in alcohol silvery on sides,
brown dorsally but darker anteriorly, blackish ventrally on head and trunk; lips, lower jaw and gill
membranes black; leading edge of snout blackish; mouth and gill cavity as in others of genus; 1D
dusky proximally, paler distally; V dusky to black; base and axil of P black. Attains more than 47
cm TL.
DISTRIBUTION.— From South China Sea off Taiwan and Hong Kong; Philippines (e., n. and
w. Luzon, off n. Mindoro, Sulu Sea off Panay); Celebes Sea off e. Borneo; Molucca Sea off Halma-
hera. Depth range about 183 to 807 m. Records of the species from s. Africa were misidentifica-
tions of a species that was subsequently described as V. mystax Iwamoto and Anderson, 1994.
REMARKS.— Chiou et al., (2004b:46, fig. 19) recorded this species from Taiwan based on two
specimens, ASIZP 61309 and 61310, which we re-identified as V. saikaiensis. However, new spec-
imens were collected off Taiwan at depths of 515–807 m. Based on examination (by TI) of many
108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 27. Valtrifossa divergens Gilbert and Hubbs, 1920. ASIZP 66928, 320+ mm TL, fresh. B. ASIZP 66781, 292+
mm TL, preserved, photo reserved laterally.
A
B

USNM specimens, there may actually be more than one species represented in the type series. The
holotype (USNM 78230) had several characters that appeared to be outside the normal range of
other specimens. These characters include a low GR-I count of 12 total (mesial side) cf. 15–18 in
others examined, narrower interorbital (24% cf. 26–32%), narrower suborbital (11% cf. 13–15%);
shorter upper jaw (43% cf. 43–49%); and shorter outer gill slit (19% cf. 24–28%). Other measure-
ments appear to be at one or another extreme in the range. The relatively sharply pointed snout and
high anterior dorsal profile also differ in degree with other specimens. Among other Ventrifossa
species from Taiwan, the one most similar is V. longibarbata, and so far as our limited study mate-
rial allow, the two species overlap or are essentially identical in their counts and measurements.
The snout in V. divergens appears to be slightly more pointed and protruding, and the broad black
stripe across the 1D in V. longibarbata is generally prominent, compared to the difuse duskiness of
that fin in V. divergens. A more in-depth study may reveal other distinguishing characters. The
overall brownish color of V. saikaiensis, its longer postorbital length (44–49% HL), and fewer scale
rows below the 2D (6.0–6.5 cf. 8–9 in V. divergens), among other characters, distinguish that
species from V. divergens. Compared with V. rhipidodorsalis, V. divergens has a longer preoral
(15–19% cf. 11–15% of HL), somewhat broader interorbital (26–32% cf. 21–26%), somewhat larg-
er orbits (with overlap, 29–38% cf. 27–32%), and somewhat shorter postorbital (with overlap,
38–44% cf. 43–48%); V. rhipidodorsalis also has a somewhat higher average number of V rays
(9–10) and more prominent black blotches on 1D.
Ventrifossa garmani (Jordan and Gilbert, 1904)
Figures 28A–B.
Coryphaenoides garmani Jordan and Gilbert in Jordan and Starks, 1904:610 (holotype USNM 50933, Saga-
mi Bay, Japan, 110–259 fm [201–474 m]; paratypes ANSP 114108 [1 spec.], CAS-SU 8548 [5 spec.],
USNM 51415 [9 spec.]).
Ventrifossa garmani: Matsubara, 1955:1315.— Kamokara, 1964:96.— Okamura, 1970:74.— Iwamoto, 1979:
152.— Okamura in Okamura et al., 1982:145, 348.— Okamura in Masuda et al., 1984:94.— Okamura in
Okamura and Kitajima, 1984:213, 360.— Iwamoto, 1990:300–301, fig. 678–679 (descr.).— Shen et al.,
1993:172 (desc.).— Chiou et al., 2004b:37, 47 (in key, list).— Shao et al., 2008: table 2 (2 spec., SWT).
MATERIAL EXAMINED (8 spec.).— SWT: ASIZP 61306 (1, 260+ TL), Dong-gang. ASIZP
65586 (1, 235 TL), CD 140, 280–452 m. Other materials: Japan: USNM 50933 (holotype, 52.8
HL, 292 TL), Albatross sta. 3695, Sagami Bay, 201–474 m. USNM 51415 (5 of 9 paratypes,
32–46.1 HL, 180–250+ TL), Suruga Bay, Albatross sta. 3738, 305 m.
DISTINGUISHING FEATURES.— 1D II,10–11; P i19–i22; V 8, rarely 9; inner GR-I 16–17 total,
outer GR-II 16–17; scale rows below midbase 1D 7.5–9.0, below 2D 4.5–5.0; lateral line scales
over distance equal to predorsal length 38–42. Snout length 25–27% HL; preoral length 18–19%;
interorbital 29–32%; orbit 30–35%; suborbital 11–15%; orbit to preopercle 41–45%; upper jaw
42–47%; barbel 23–29%; outer gill slit 24–27%. Head and body relatively compressed; snout rel-
atively short, high, forming obtuse angle viewed dorsally. Body scales covered with broad-based,
triangular spinules in quincunx arrangement. Spinous ray of 1D usually less than HL, leading edge
finely sawtoothed. Periproct occupying about half anterior space between V and A, close behind V
bases, dermal window of light organ extends forward between V bases. Gill membranes narrowly
united over isthmus and under midorbit. Color in preservative light to medium gray-brown over-
all, silvery over most of head, trunk, and anteriorly on sides of tail, but not over dorsum; jaws, gill
membranes, and below suborbital ridge blackish; mouth pale, gill cavities blackish posteriorly;
leading edge of snout blackish; 1D dusky, somewhat darker midlaterally; V blackish; P and A clear
to dusky. Attains about 30 cm TL.
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 109

DISTRIBUTION.— From s. Japan to sw Taiwan [SCS], in 280–452 m.
REMARKS.— This is one of the most abundant species of macrourid in Japan, but it is not par-
ticularly common off Taiwan. The broadly triangular scale spinules on body scales are unique
among the Taiwan members of this genus.
Ventrifossa longibarbata Okamura, 1982
Ventrifossa longibarbata Okamura in Okamura et al., 1982:157–159, pl. 94 (holotype BSKU 29494; Tosa
Bay, Japan, 32°58.0ʹN, 133°32.0ʹE, 605 m; 4 paratypes, Tosa Bay, Suruga Bay, and Okinawa Trough, in
382–620 m).— Okamura in Masuda et al., 1984:94.— Okamura in Okamura and Kitajima, 1984:215,
361.— Iwamoto and Williams, 1999:228 (comparisons with V. nigrodorsalis et al.).— Chiou et al.,
2004b:48, fig.20 (2 spec.; sw. Taiwan).
MATERIAL EXAMINED (45 spec.).— NET: ASIZP 65609 (1, 301 TL), CD 209, 508–522 m;
ASIZP 70242 (1, 133 TL), Da-xi. SWT: ASIZP 61242 (1, 272 TL), Dong-gang; ASIZP 61243 (2,
182–268 TL), Dong-gang and CAS 224884 [ex ASIZP 61243] (1, 238+ TL), Dong-gang; ASIZP
64114 (5, 185–225+ TL) and ASIZP 66950 (5, 53–55 TL), CD 194, 507 m; ASIZP 65532 (1, 290
TL) and ASIZP 65556 (1, 180 TL), CD 137, 316–477 m; ASIZP 65602 (1, 185 TL), CD 138, 441
m; ASIZP 65629 (1, 155 TL), CD 233, 448–526 m; ASIZP 65537 (1, 260 TL), CD 133, 690–748
m. SCS: ASIZP 66102 (1, 58 HL, 285+ TL), CP 315, 509 m; ASIZP 66257 (1, 35.5 HL, 155+ TL),
ASIZP 66267 (1, 31.6 HL, 140+ TL), and ASIZP 66779 (7, 23–36 HL, 110+-180+ TL), CD 311,
516 m; ASIZP 66948 (2, 150–210 TL), CD 136, 998–1211 m; ASIZP 66263 (1, 37 HL, 200+ TL),
CD 310, 364 m; ASIZP 66749 (3, 25–31 HL, 110+-170+ TL),) and ASIZP 66778 (2, 30–37 HL,
150+-160+ TL), CP 314, 506 m; ASIZP 66777 (2, 145–145+ TL, 24–30 HL; CP 313, 513 m;
110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
FIGURE 28. Ventrifossa garmani Jordan and Gilbert, 1904. ASIZP 61306, 260+ mm TL, preserved. A. lateral view of
whole fish. B. lateral view of right head, photo reversed laterally.
A
B

ASIZP 66812 (2, 24–25.5 HL, 120+-130+ TL), OCP 317, 515 m; ASIZP 66829 (2, 100+-130+ TL),
CP 315, 509 m.
DISTINGUISHING FEATURES (data from six Taiwan specimens).— 1D II,10–11; P i18–i21; V
8–9; total GR-I (lat/mesial) 10–12/15–17, GR-II 14–17/14–16; scale rows below midbase 1D
4.5–7.5, below 2D 5.0–6.5(9.0); lateral line scales over distance equal to predorsal length 37–41.
Snout length 26–28% HL; preoral length 16–17%; interorbital 25–31%; orbit 29–36%; suborbital
12–14%; postorbital 42–48%; orbit to preopercle 41–46%; upper jaw 40–47%; barbel 28–39%;
outer gill slit 26–28%. Color in life (from Okamura in Okamura et al., 1982:159) dark gray or
swarthy dorsally, silvery on sides of head and body, black over gular region, lips, lower jaw, gill
membranes, chest and ventrally on trunk; abdomen bluish under silvery reflections, suborbital
region and ventral and posterior margin of gill cover blackish; mouth pale, gill cavities blackish
posteriorly; leading edge of snout blackish; 1D with wide dusky to blackish midlateral section, but
pale distally and along base; V black; P blackish; base of P silvery but with black distal edge and
mesial lunate area. Attains at least 31 cm TL.
DISTRIBUTION.— From s. Japan to Taiwan (SWT) and South China Sea in 382–1211 m.
REMARKS.— This species closely resembles V. garmani in general physiognomy but differs in
having a slightly longer barbel (28–39% HL vs. 21–29%), a darker midlateral region of 1D, more
pyloric caeca (about 70 vs. <53 fide Okamura 1970:77), slender scale spinules (vs. triangular), and
possibly darker V (black vs. dusky or blackish) and P (which appears to be blackish in Okamura’s
[in Okamura et al., 1982:156] photograph of the holotype vs. “light” in V. garmani [Okamura,
1970:77]). The six Taiwan specimens from which the above measurements and counts were taken
agree closely with the original description of V. longibarbata except for slight differences in
interorbital width (25–31% HL cf. 21–26.5%), postorbital length (42–48% cf. 39–44%), P ray
count (i18–i21 cf. i21–i26”), and scale rows (below 1D origin 7–12 cf. 12–14, below mid-base 1D
4.5–7.5 cf. 8.5–11.5, and below 2D origin 5.0–9.5 cf. 9.0–10.0). The 1D was also almost entirely
dusky, with only faint trace of a midlateral stripe. Four additional examples taken off Kochi, Japan
(BSKU 12919, 12920, 12922, and 12923) were examined by TI many years ago; they agree well
with the original description of the species.
Ventrifossa macroptera Okamura, 1982
Ventrifossa macroptera Okamura in Okamura et al., 1982:149 (holotype BSKU 32185, Kyushu-Palau Ridge,
27°55.1ʹN, 134°44.8ʹE, 685–710 m; paratypes BSKU 30432 [1 spec.], 30435–36 [2], 30467–69 [3],
29429 [1], 32153–84 [31], 32186–89 [4], 32192–200 [8]).— Okamura in Masuda et al., 1984:94 (com-
piled).— Iwamoto, 1990:303, figs. 682–683 (descr.).— Chiou et al., 2004:48, fig. 21 (2 spec.; Tungkang,
Taiwan [SWT]).
Ventrifossa atherodon (not of Gilbert and Cramer, 1897): Chiou et al., 2004b:46, fig. 18 (ASIZP 61311 (1
spec., Da-xi, Taiwan [NET]).
MATERIAL EXAMINED (103 Taiwan spec.).— NET: ASIZP 60253 (1, 286 TL), Da-xi; ASIZP
65533 (1, 190 TL), CD 210, 445–1185 m; ASIZP 61311 (1, 245 TL), Da-xi; ASIZP 70682 (1, 47
HL), Da-xi; ASIZP 70729 (1, 254 TL), Da-xi. SWT: ASIZP 61244 (1, 199 TL), Dong-gang; ASIZP
61245 (1, 202 TL), Dong-gang; ASIZP 65525 (3, 170–190 TL0, CD 142, 227–335 m; ASIZP
66947 (2, 235–250 TL), CD 140, 280–452 m; ASIZP 67596 (3, 118–143 TL), CP 348, 395 m;
ASIZP 70254 (1, 180 TL), Dong-gang. SCS: ASIZP 65522 (18, 90–185 TL), ASIZP 65590 (41,
143–221 TL), ASIZP 65612 (5, 175–225 TL), and ASIZP 65614 (4, 111–132 TL), CD 136,
998–1211 m; ASIZP 65534 (1, 280 TL), CD 137, 316–477 m; ASZIP 65623 (1, 208 TL), CD 233,
448–526 m; ASZIP 65640 (1, 218 TL), CD 233, 448–562 m; ASIZP 65679 (1, 21 HL, 97+ TL), CP
315, 509 m; ASIZP 66751(1, 125 TL), OCP 312, 517 m; ASIZP 66764 (2, 35–38 HL, 130+-160+
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 111

TL), CP 314, 506 m; ASIZP 66798 (6, 20–38 HL, 110–180+ TL), CD 311, 516 m; ASIZP 66804
(2, 125–255 TL), OCP 317, 515 m; ASIZP 66808 (1, 180 TL), CP 314, 506 m; ASIZP 66832 (1,
143 TL), CP 315, 509 m; ASIZP 66949 (2, 130+-144 TL), CP 234, 547 m.
Other specimens: Japan (all from Kyushu-Palau Ridge). HUMZ 75049 (330+ TL); 75050
(325+ TL); 75051 (328+ TL); 75052 (332+ TL); 75053 (250+ TL); 75054 (338+ TL); 75055 (290+
TL); 75056 (356+ TL); 75057 (295 TL). CAS 52971 (3 paratypes, ex. BSKU spec.; 60.0–73.0 HL,
290+-376+ TL); CAS 52972 (7 paratypes, ex. BSKU; 58.2–70.3 HL, 235+-360+ TL).
DISTINGUISHING FEATURES.— 1D II,9–10(11); P i19–i23; V 9–10; GR-I (outer/inner) (10)
13–15 / 16–18 total, GR-II (outer/inner) 16–18 / 16–18; scale rows below 1D origin 9.5–11.0
(12.0), below midbase 1D 6.0.0–7.5, below 2D 7.5–10.0; lateral line scales over distance equal to
predorsal length 48–57. Snout length 27–32% HL; preoral length 12–17%; internasal 23–27%;
interorbital 29–32%; orbit 26–31%; suborbital 12–15%; postorbital 42–48%; orbit to preopercle
45–52%; upper jaw 47–51%; barbel 20–27%; outer gill slit 26–33%. Second spinous ray of 1D
smooth. Snout rather blunt, little protruding beyond mouth; suborbital shelf narrowly constricted
anteriorly below posterior nostril. Premaxillary tooth band broad; outer series of recurved, canine-
like teeth with arrowhead-shaped tips; lower jaw teeth in two series. Color overall dark, head
swarthy to black, somewhat silvery on sides of head and body, blackish over gular region and chest,
de-scaled areas on trunk and anterior part of tail bluish; mouth pale, gill cavities blackish posteri-
orly; gill arches dark; leading edge of snout blackish; fins blackish; broad lunate area mesial to P
base black. Attains about 40 cm TL.
DISTRIBUTION.— Kyushu-Palau Ridge to Taiwan and the South China Sea. In Taiwan, speci-
mens were collected off NET, SWT, and SCS, mostly at depths between 280 and 516 m, but one
collection (CD 136) from 998–1211 m.
REMARKS.— Chiou et al. (2004b:46, fig. 18) recorded a specimen referred to V. atherodon that
is now re-identified as V. macroptera. Okamura (in Okamura et al. 1982:151) considered this
species to be the most dominant of the grenadiers off the Kyushu-Palau Ridge. It appears to be sim-
ilarly abundant in the South China Sea. In respect to the smooth spinous 1D ray, the low blunt
snout, and dentition, V. macroptera resembles V. atherodon (Gilbert and Cramer, 1897),
V. macrodon Sazonov and Iwamoto, 1992 from the se. Pacific, and V. sazonovi Iwamoto and
Williams, 1999 from w., nw., and ne. Australia and the South China Sea. The last species is notable
in having enlarged spinules on scales of the dorsum below the anterior portion of the 2D.
Ventrifossa misakia Jordan and Gilbert, 1904
Figure 29.
Coryphaenoides misakius Jordan and Gilbert, 1904:611–612 (holotype, CAS-SU 8107, 340 mm TL; Misaki,
Japan).
Macrourus asper (not of Günther): Jordan and Thompson, 1914:306, pl. 38, fig. 2 (“Misaki in deep water;”
no description.)
Lionurus misakius: Gilbert and Hubbs, 1916:194–195 (1 spec., 113+ mm TL).
Ventrifossa misakia: Okamura, 1970:78–81, pl. VI, text-fig. 35 (47 spec., 150–260 mm TL; s. Japan).—
Iwamoto, 1990:304–305, fig. 684–685 (descr.).— Sazonov and Shcherbachev, 1997:529–533 (compar-
isons; synonymized V. fusca with V. misakia).— Nakabo, 2002:423 (compiled).
Ventrifossa fusca Okamura in Okamura et al., 1982:153–155, fig. 93, A-D (p. 152) (holotype, BSKU 26067,
female, 612 mm TL; Kyushu-Palau Ridge, 27°55ʹN, 134°39ʹE, 700 m; paratypes [same locality], BSKU
26068 [female, 526 TL], 26069 [male, 434+ TL], 26115 [male, 468 TL]).— Okamura in Masuda et al.,
1984:94, pl. 81–B (compiled).— Nakabo, 2002:422 (compiled).— Shao et al., 2008 (6 spec., SCS; first
record for Taiwan).
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MATERIAL EXAMINED (6 spec. from Taiwan).— SCS: ASIZP 66783 (6, 157–380+ TL), CD
320, 731 m. Other specimens: Japan: CAS-SU 8107 (holotype, 60 HL, 340 TL), Sagami Bay
near Misaki, Japan. USNM 51421 (3 paratypes, 54–76 HL, 165+-380 TL), Misaki. BSKU 12002
(1, 50 HL, 240 TL); Tosa Bay. BSKU 12953 (1, 46 HL, 225+ TL), Tosa Bay. BSKU 12955 (1, 28
HL, 157 TL). NSMT P49952 (1, 42.7 HL, 198+ TL), Suruga Bay, 440 m. NSMT P78960 (1, 44.3
HL, 230+ TL) and P78961 (1, 43.0 HL, 230+ TL), Suruga Bay, 376 m. FAKU 5270 (1, 36.8 HL,
197+ TL), M7634 (1, 42 HL, 221 TL), M7651 (1, 38 HL, 190+ TL), and M7655 (39.5 HL, 194
TL), Chiba Pref., off Choshi.
DISTINGUISHING FEATURES.— 1D II,10–12; P i19–i23; V 8 (rarely 9); total GR-I
(lateral/mesial) 11–12 / 13–16 total, (lateral/mesial) GR-II 14–15 / 12–15; scale rows below mid-
base 1D 8.5–10.5, below 2D 9–10; lateral line scales over distance equal to predorsal length 57–76.
Snout length 27–33% HL; preoral length 20–27%; internasal 22–29%; interorbital 29–35%; orbit
32–38%; postorbital 36–44%; orbit to preopercle 40–45%; suborbital 12–15%; upper jaw 37–42%;
barbel 4–9%; outer gill slit 18–24%; 1D-2D interspace 40–56%. Head broad, body relatively com-
pressed; snout conical, protruding well beyond mouth, with a small tubercle at tip; suborbital area
gently rounded, ridge defined but lacking stoutly modified scales, suborbital shelf abruptly con-
stricted anteriorly; head fully scaled, but lacking large pores; body scales covered with slender,
needle-like spinules in quincunx pattern. Color in alcohol light to medium gray-brown with silvery
reflections along sides; bluish black over abdomen and chest; gill cover and gill membranes black-
ish; mouth and gill cavities pale, but blackish posteriorly; tip of snout (and in young along leading
edge) blackish; 1D dusky, V black, P and A dusky. Attains about 40 cm TL.
DISTRIBUTION.— From se. coast Japan to the East China Sea and South China Sea off sw. Tai-
wan, in 200 to 731 m.
REMARKS.— We have been informed by Dr. Hiromitsu Endo of Kochi University that speci-
men BSKU 28596 recorded by Okamura (in Okamura and Kitajima 1984:213) as V. misakia is
actially a specimen of V. johnboborum. We follow Sazonov and Shcherbachev (1997:529) in syn-
onymizing V. fusca Okamura with V. misakia. It is rather peculiar that Okamura (in Okamura et al.
1982) failed to recognize specimens of his new species as representing V. misakia or a species very
similar, knowing that he had examined many specimens of V. misakia for his work on the Japan-
ese macrourids in the Fauna Japonica book series (Okamura 1970:78) and treating both species in
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 113
FIGURE 29. Ventrifossa misakia Jordan and Gilbert, 1904. ASIZP66783, 1 of 6, 380+ mm TL, preserved.

Masuda et al. (1984). The first author (TI) has examined two paratypes of V. fusca (BSKU 26068,
99 mm HL, 526 mm TL, and 26069, 106 HL, 434+ TL [large pseudocaudal]), both of which are
much larger than any other we have seen. They are nonetheless identical in every feature with our
smaller Tawian representatives of V. misakia, so far as we can tell. Sazonov and Shcherbachev
(1997:532) pointed out that Okamura compared his species with Kuronezumia macronema (Smith
and Radcliffe, 1912), a very different species that has been classified in another genus. The close
similarity of V. misakia and V. johnboborum Iwamoto, 1982 was also recognized by Iwamoto
(1982:59–60; 1990:305) and Sazonov and Iwamoto (1992:80). Sazonov and Shcherbachev
(1997:529) compared specimens of V. johnboborum from the w. Indian Ocean, e. and w. Australia,
the South China Sea, the Sala y Gomez Ridge (se. Pacific), and the holotype taken in the Bismar-
ck Sea. They found these populations “differing in a few morphometric indices while retaining sev-
eral common characters not peculiar to V. misakia (including V. fusca).” Among the different pop-
ulations, that from the South China Sea was most similar to V. misakia, but they were unable to
arrive at any conclusions as to “the independence of V. johnboborum.” Accordingly, we continue
recognition of V. johnboborum while awaiting a more detailed analysis, perhaps using DNA infor-
mation. Our specimens represent the first record of V. misakia from Taiwan and the South China
Sea and also the deepest record at 731 m.
Ventrifossa nigrodorsalis Gilbert and Hubbs, 1920
Ventrifossa nigrodorsalis Gilbert and Hubbs, 1920:546 (holotype, USNM 83627; n. coast Mindanao, Philip-
pines, 391 m; 168 paratypes from Formosa [Taiwan], Philippines, and East Indies, 290–868 m). Iwamoto,
1990:307–309, fig. 690 (descr.).— Iwamoto and Merrett, 1997:559 (New Caledonia and vicinity).—
Iwamoto and Williams, 1999:224–225, 228, fig. 54 (descr., e. and w. Australia).— Iwamoto and Graham,
2001:497–498, fig. 112 (descr., se. Australia, 300–790 m).— Chiou et al., 2004b:47 (listed, Taiwan,
Table 1).
MATERIAL EXAMINED (3 Taiwan spec.).— NET: ASIZP 66903 (1, 197 TL), CP 248, 536 m;
ASIZP 70715 (1, 188 TL), Da-xi. SCS: ASIZP 57974 (1, 183+ TL), Tong-sa Islands. Other spec-
imens: USNM 8362 (holotype, 41.5 HL, 214 TL), Philippines, Mindanao Sea; 8˚37’37ʺN,
124˚35ʹE, 391 m, Albatross sta. 5502, 4 Aug. 1909. USNM 149302 (paratypes, 4, 32.3–36.0 HL,
125+-195+ TL), same data as for holotype; CAS 64574 (5, 30–9–36.6 HL, 499+-193+ TL), SCS
off Vietnam; 15˚40’00ʺN, 109˚47ʹE, 479 m. CAS 221057 (38.5 HL, 210+ TL), Philippines, e. coast
Luzon, 14˚41’04ʺN, 123˚24’07ʺE, 435–451 m, Fishery Researcher I, field no. TI95–12, 27 Sept.
1995.
DISTINGUISHING FEATURES.— 1D II,9–10; P i19–i23; V 8–9; total GR-I (lateral/mesial)
9–13/13–15, GR-II 13–14 /13–14; scale rows below 1D 8–9, below 2D 7–9; lateral line scales over
distance equal to predorsal length 36–43. Snout length 29–30% HL; preoral length 19–21%; inter-
nasal 22–23%; interorbital 23–28%; orbit 28–34%; postorbital 42–44%; orbit to preopercle
39–42%; suborbital 13–16%; upper jaw 37–41%; barbel 17–26%; outer gill slit 21–25%; 1D-2D
interspace 39–66%. Body and head moderately deep and compressed, nape somewhat elevated;
snout slightly protruding beyond mouth, acutely pointed in lateral view, broadly obtuse in dorsal
view. Body scales covered with small, needle-like spinules in wide chevron-like rows; small area
of spinuleless scales along and behind 1D base. Color in alcohol grayish-brown dorsally on trunk,
becoming paler along 2D base to form a dorsolateral stripe on tail; top of head and snout pale,
integument translucent; faint silvery reflections along sides of head and ventrally on trunk and tail;
underlying silvery pigmentation abdomen and chest bluish-black, tail pale with light peppering;
gill cover and gill membranes mostly black; mouth and gill cavities pale, but dark in gullet; black
margins on suborbital shelf, leading edge of snout, and supranarial ridges; 1D with black blotch or
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blackish over middle third or so of fin but pale distally and along base; V black, sometimes paler
distally and near base; P and A dusky, but base of P black with lateral surface mostly silvery. Lips,
jaws, and gill cavity as in others of genus. Attains about 25 cm TL in Taiwan region, possibly more
elsewhere.
DISTRIBUTION.— Known from off Australia, New Caledonia region, and Indonesia n. to the
Philippines, Taiwan (NET), and the South China Sea.
REMARKS.— Okamura (in Okamura et al. 1982:147, 348, fig. 90) initially recorded this
species from two specimens collected in Tosa Bay. He later (Okamura et al. 1984) treated those
specimens as representatives of his new species V. rhipidodorsalis, and used the photograph of one
of the specimens (fig. 146C; BSKU 29495, 172 mm TL) to illustrate another example of the new
species. Ventrifossa nigrodorsalis is an apparently broadly distributed species abundant in the
Philippines, Indonesia, and the warm-water regions of Australia, as well as the sw. Pacific off New
Caledonia, Vanuatu, and near regions, but it was scarce off Taiwan. Characters that have been used
to distinguish species of Ventrifossa are often subtle and dependent on well-preserved specimens
to observe. The existence of seemingly widely scattered populations of V. nigrodorsalis heighten
the possibility that other species lie hidden under the cloak of that species name as currently cir-
cumscribed, and they may be unveiled after closer study of more specimens. The description pro-
vided above is from specimens collected off Taiwan, the South China Sea, and the Philippines in
the Mindanao Sea between Mindanao and Cebu.
Ventrifossa rhipidodorsalis Okamura, 1984
Ventrifossa rhipidodorsalis Okamura in Okamura and Kitajima, 1984:205 (Okinawa Trough, 28°42.0ʹN,
127°09.0ʹE, 500–535 m (holotype, BSKU 27695; 23 paratypes, Okinawa Trough and Tosa Bay, 500–650
m [but 1 spec. at 220 m]).— Okamura in Masuda et al., 1984:94 (compiled).— Iwamoto and Williams,
1999:228 (mentioned).— Chiou et al., 2004b:48, fig.22 (13 spec.; Nanfangao, Taiwan [NET]).
? Ventrifossa nigrodorsalis: Okamura in Okamura et al., 1982:147, 348, fig. 90 (2 spec.; BSKU 29495, later
identified as V. rhipidodorsalis by Okamura in Okamura and Kitajima, 1984: fig. 146C).
MATERIAL EXAMINED (70 Taiwan spec.).— NET: ASIZP 61246 (1, 170 TL), Nan-fang-ao;
ASIZP 61247 (12, 142–168 TL), Nan-fang-ao; ASIZP 62331 (2, 126+-162+ TL), Fong-kang, 200
m; ASIZP 64236 (3, 110+-215 TL) and CAS 224885 (ex. ASIZP 64236) (2, 121+~161 TL), CP
234, 547 m; ASIZP 65549 (1, 160+ TL), CP 124, 1129–1165 m; ASIZP 70231 (2, 109–113 TL),
Da-xi; ASIZP 70248 (9, 98–180 TL), Nan-fang-ao; ASIZP 70250 (1, 125 TL), Da-xi; ASIZP 70685
(1, 40 HL), Da-xi; ASIZP 70733 (1, 196 TL), Da-xi. SWT: ASIZP 62388 (1, 181 TL), Dong-gang,
300 m; ASIZP 66322 (1, 153 TL), Dong-gang. SCS: ASIZP 65521 (11, 100–142 TL) and ASIZP
65592 (18, 105–165 TL), CD 136, 998–1211 m; ASIZP 65588 (3, 140–155 TL), CD 141, 985–1110
m. SET: ASIZP 66104 (1, 65 HL, 285+ TL), CP 299, 799 m. Other specimens: BSKU 27695
(holotype) and six paratypes, BSKU 27123, 27191, 27660, 27696, 27861, 27865, Okinawa Trough.
CAS 88668 (5, 51.6–58.7 HL, 233+-312+ TL), Philippines, off se coast Luzon; 14˚50.46ʹN,
123˚17.30ʹE, 760–770 m; R/V Fishery Researcher I sta. TFRI-Ph1–12–95, 27 Sep. 1995.
DISTINGUISHING FEATURES.— 1D II,10–12; P i19–i23; V 9–10; total GR-I (lateral/mesial)
10–13/15–17 total, (lateral/mesial) GR-II 14–17/(13)15–17; scale rows below midbase 1D 5.5–7.5,
below 2D 7–8.5; lateral line scales over distance equal to predorsal length 36–43. Snout length
25–32% HL; preoral length 11–15%; internasal 18–23%; interorbital 21–27%; orbit 26–34%; pos-
torbital 42–48%; orbit to preopercle (38) 41–44%; suborbital 11–14%; upper jaw (39) 42–47%;
barbel (16) 19–26%; outer gill slit 22–28%; 1D-2D interspace 45–62%. Body moderately deep and
compressed; snout low, blunt in adults, somewhat more pointed and protruding in young; subor-
bital area gently rounded. Body scales covered with small, slender, needle-like spinules in wide
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 115

chevron-like rows; an area of spinuleless scales along and behind 1D base. Color in alcohol over-
all dark, with faint silvery reflections along sides of head and trunk; bluish-black over abdomen
and chest; gill cover and gill membranes mostly black; mouth and gill cavities pale, but dark in gul-
let; black margins on suborbital shelf, leading edge of snout, suparnarial ridges, and a V-shaped
stripe joined at apex to black nape stripe before 1D; 1D black except distal third or less white, base
dusky to pale; V black, P dusky but base black with bulk of lateral face of base silvery, A dark
dusky overall but with narrow black distal margin anteriorly. Lips, mouth, jaws, and gill cavity as
in others of genus. Attains about 33 cm TL.
DISTRIBUTION.— From s. Japan to sw. Taiwan (SCS) in 400–1211 m.
REMARKS.— Among the Taiwanese grenadiers, V. rhipidodorsalis is most similar to V. nigro-
dorsalis, but that species has 8 or 9 V rays (cf. 9 or 10), overall paler coloration; paler, more sil-
very sides; slightly sharper, more protruding snout; generally narrower black streak on 1D; and
shorter upper jaw (37–41% HL). In its overall dark color and V count of 9–10, it resembles
V. macroptera, but that species has a smooth 1D spinous ray and differences in dentition. Another
species with a black streak across the 1D is V. longibarbata, but that species has slightly lower
average number of V rays (8–9), much longer barbel (28–39% HL cf. 19–26%), slightly longer pre-
oral length (16–17% cf. 11–15%), and somewhat fewer scale rows below 2D (5.0–6.5 cf. 7.0–8.5).
Ventrifossa saikaiensis Okamura, 1984
Ventrifossa saikaiensis Okamura in Okamura and Kitajima, 1984:209 (holotype, BSKU 28004; Okinawa
Trough, 28°50.0ʹN, 127°14.0ʹE, 700–740 m; 11 paratypes, Okinawa Trough).— Chiou et al., 2004b:
48–49, fig. 23 (19 spec.; Taiwan [NET]).
Ventrifossa divergens (not of Gilbert and Hubbs, 1920): Chiou et al., 2004b:46, fig.19 (2 spec.; Da-xi, Taiwan
[NET]).
MATERIAL EXAMINED (58 Taiwan spec.).— NET: ASIZP 61248 (1, 250 TL), Da-xi; ASIZP
61249 (14, 102–168 TL), Da-xi; ASIZP 61307 (1, 224 TL), Da-xi; ASIZP 61310 (1, 182 TL), Da-
xi; ASIZP 64094 (1, 53 HL, 260+ TL), CP 235, 764 m); ASIZP 63271 (1, 144 TL), Da-xi; ASIZP
64094 (1, 260+ TL), CP 235, 764 m; ASIZP 65608 (1, 145+ TL), CP 120, 520 m;ASIZP 65602 (1,
295+ TL), CD 138 m; ASIZP 70223 (1, 60 HL), Da-xi; ASIZP 70225 (2, 270+-395 TL), Da-xi;
ASIZP 70730 (1, 170 TL), Da-xi. SWT: ASIZP 61242 (1, 272 TL), Dong-gang; ASIZP 61309 (1,
220 TL), Dong-gang; ASIZP 64119 (1, 278+ TL) and ASIZP 64253 (1, 200+ TL), CD 193, 821 m;
ASIZP 65601 (2, 285 TL) and CAS 224887 (ex. ASIZP 65601, 1, 299 TL), CD 139, 718–852 m;
ASIZP 65608 (1, 145+ TL), CP 120, 520–640 m; ASIZP 65616 (1, 185 TL), CD 138, 441 m;
ASIZP 70273 (1, 295 TL), CD 138, 441 m; ASIZP 71146 (1, 267 TL), CD 271, 700–800 m. SCS:
ASIZP 64562 (1, 275 TL) and ASIZP 65562 (1, 275 TL), CD 140, 280–452 m; ASIZP 65522 (18,
90–185 TL), CD 136, 998–1211 m; ASIZP 61242 (1, 272 TL), Dong-gang. Other specimens (all
from Okinawa Trough, East China Sea): BSKU 28004 (holotype, 305+ TL); BSKU 27576
(paratype, 285+ TL); BSKU 28005–28008 (4, 180+–290 TL).
DISTINGUISHING FEATURES.— 1D II,9–11; P i18–i22; V 8–9; total GR-I (lateral/mesial) 12–16
/ 16–18, GR-II 15–18 /15–18; scale rows below midbase 1D 5.5–7.0, below 2D 6.0–7.5; lateral line
scales over distance equal to predorsal length 33–44. Snout length 28–31% HL; preoral length
14–19%; internasal 21–26%; interorbital 27–32%; orbit 27–32%; postorbital 44–48%; orbit to pre-
opercle 43–46%; suborbital 14–16%; upper jaw 45–50%; barbel 27–35%; outer gill slit 27–30%;
1D–2D interspace 43–68%. Body and head moderately deep and compressed; snout low, rather
blunt and protruding little beyond large mouth; barbel long, slender. Body scales densely covered
with slender, reclined, needle-like spinules in wide chevron-like rows; no spinuleless scales around
1D base. Color in alcohol medium brownish, lacking silvery reflections; blackish over gill cover;
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bluish-black over abdomen and chest; gill and gular membranes black; mouth and gill cavities pale,
but dark in gullet; margins on suborbital shelf and leading edge of snout faintly blackish or not dis-
tinguished; barbel pale, but base dark; 1D uniformly dusky; V blackish; P and A dark dusky, but
base of P black along distal margin. Coloration of lips, mouth, jaws, teeth, and gill cavities as in
others of genus. Attains more than 31 cm TL.
DISTRIBUTION.— Known only from the Okinawa Trough (in East Sea), Taiwan (NET, SWT),
and the South China Sea in 280–1211 m.
REMARKS.— Chiou et al. (2004b:46, fig.19) recorded two specimens (ASIZP 61309 and
61310) as V. divergens, but we re-identified them as V. saikaiensis. Among the Taiwan grenadiers,
V. saikaiensis is most similar to V. divergens in having a uniformly dusky 1D, but it is easily dis-
tinguished from that species (and all other Taiwan members of Ventrifossa) in having a uniformly
brownish overall coloration, with little or no silvery reflections on the sides, and dark brown scale
pockets. In these color features and in its general appearance, the species resembles certain mem-
bers of the genus Nezumia, especially N. atlantica (Parr, 1946) and N. africana (Iwamoto, 1970).
However, the dentition, squamation, luminescent organ, gill rakers and arches, and other features
clearly support its position within Ventrifossa.
Ventrifossa sazonovi Iwamoto and Williams, 1999
Figure 30.
Ventrifossa sazonovi Iwamoto and Williams, 1999:231, fig. 56 (holotype: CAS 13564, South China Sea, off
Vietnam, 15°48ʹN, 109°47ʹE, depth 479 meters; 15 paratypes, n. and w. Australia).
MATERIAL EXAMINED.— SWT: BSKU 98976 (1, 38.5 mm HL, 202+ mm TL), Dong-gang
Fish Market, ca. 200–300 m depth, bottom trawl, coll. by H.-C. Ho, 16 Nov. 2007.
DISTINGUISHING FEATURES.— 1D II,10; P i21–22; V 9; GR-I (outer/inner) 13/17; GR-II
(outer/inner) 18/17. Snout 27% HL; preoral 10% HL; interorbital 25% HL; orbit 33% HL; subor-
bital 11% HL; orbit to preopercle 45% HL; upper jaw 47% HL; barbel 21% HL; outer gill slit 27%
HL. Head and body moderately compressed; snout relatively short, barely protruding beyond upper
jaw; suborbital shelf somewhat constricted anteriorly; chin barbel fairly developed; pectoral fin
rather short, 1.7 in HL. Branchiostegal membranes narrowly united over isthmus; gill opening
extending forward to below hind 1⁄3of orbit. Periproct situated just behind V insertions, occupying
about 1⁄3of V –A interspace; dermal window of light organ small, reaching on a line passing through
outer V bases. Premaxillary teeth in broad tapered band; outer series distinctly enlarged; dentary
teeth aligned in irregular 2 rows. Spinules on body scales needle-like, arranged in quincunx order;
those on scales along 2D base prominently enlarged. No modified scutes on snout. 1D smooth
along its leading edge. Color in preservative light brown overall, but dark over ventral parts of head
and abdomen; prominent blackish streaks along leading edge of snout, upper suborbital shelf, and
IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN 117
FIGURE 30. Ventrifossa sazonovi Iwamoto and Williams, 1999. BSKU 98976, 202+ mm TL, preserved.

median nasal ridge; lips and chin barbel pale; oral cavity immaculate; 1D darker proximally, paler
distally; V blackish; other fins dusky overall. Attains at least 34 cm TL.
DISTRIBUTION.— Known from the South China Sea off Vietnam and Taiwan, and northern
coasts of Australia from Western Australia to Queensland, in about 200–610 m.
REMARKS.— This species is readily distinguished from other Taiwanese congeners except
V. macroptera in having a smooth leading edge of 1D. It further differs from V. macroptera in hav-
ing enlarged spinules on scales along 2D base. The Taiwanese specimen is the shallowest captures
(previously known only below 420 m) and the northernmost record of the species.
Family Macrouroididae
DISTINGUISHING FEATURES.— Head huge, rounded, with consistency of water-filled balloon;
eyes tiny, diameter about 10 times into head length, placed forward of upper jaws; dorsal fin sin-
gle, low; A long low; V absent (Macrouroides) or tiny with five short rays (Squalogadus); chin bar-
bel absent; outer GR-I somewhat lathlike; first (outer) gill slit not restricted dorsally and ventrally
by opercular membrane.
REMARKS.— Two genera, each with one widespread species, found in most tropical and tem-
perate oceans.
Genus Squalogadus Gilbert and Hubbs, 1916
DISTINGUISHING FEATURES.─V present but tiny, five rays, none prolonged; other features as
for family.
Squalogadus modificatus Gilbert and Hubbs, 1916
Figure 31.
Squalogadus modificatus Gilbert and Hubbs, 1916:156 (Bungo Channel, 32°32ʹN, 132°25ʹE, off Kyushu,
Japan, Albatross sta. 4956, 720 fm [1317 m]; holotype USNM 76864, paratypes SU 22928).— Marshall,
1973:517–518.— Okamura, 1970:16–18, pl. IX (Sagami Bay and off Choshi, Japan).— Shiobara,
1982:143–146, figs. 1–3 (2 spec., 261–397 mm TL; Suruga Bay, Japan).— Sawada in Amaoka et al.,
1983:105, 192, fig. 57 (HUMZ 78126, 1 spec., off Miyagi, Tohoku district, 1110 m).— Okamura in Masu-
da et al., 1984:93.— Endo et al., 1994:332 (HUMZ 121632, 1 spec., Sea of Okhotsk off ne. Hokkaido,
1393 m; n. record).— Okamura and Amaoka, 1997:129 (photograph of living individual).— Shinohara et
al., 1996:170 (7 spec., east-central Honshu, Japan; 967–4867 m).— Shinohara et al., 1997:290 (listed).—
Shinohara et al., 2001:306 (1 spec., Tosa Bay, 765–823 m).— Amaoka, 2009:177, fig. 304 (photo of fresh
specimen). See also Shao et al. (2008: table 1, 1 spec., SCS) first record for Taiwan.
Squalogadus intermedius Grey, 1959:330–333, fig. 53 (holotype USNM 185606, n. Gulf of Mexico, R/V Ore-
gon sta 1426, 1098 m; 5 paratypes, FMNH 64489).
MATERIAL EXAMINED.— SCS: ASIZP 64070 (1, 330+ TL), CP 178, 1241 m.
DISTINGUISHING FEATURES.— As for genus.
DISTRIBUTION.— Widespread in tropical and southern temperate seas, but absent in e. Pacific.
In nw. Pacific, known off Japan, from Hokkaido to Kyushu and Taiwan.
REMARKS.— A single specimen collected from SCS off sw. Taiwan in 1241 m represents the
first record from Taiwan.
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ACKNOWLEDGMENTS
Iwamoto owes thanks to many individuals who helped in a variety of ways, but perhaps most
notably his co-authors. He is particularly indebted to co-author Shao, who made possible three trips
to Taiwan, provided gracious hospitality, and the use of his facilities and student helpers—this pub-
lication would not have been possible without his support and encouragement. Co-author Ho, a
most amicable travel companion and host, was instrumental in pushing this project along and doing
much of the detailed work of checking specimen data and literature, photographing specimens and
touching up the images, and arranging visits to fish markets. Dr. H. Endo and co-author Nakaya-
ma were most helpful during a visit in 2010 to BSKU; Dr. Endo also provided a very thorough and
detailed review, catching some errors and adding important information, to improve the manu-
script. Thanks go to many others for various help: D.-M. Chen for collecting many of the speci-
mens for this study; T.-Y. Chan led the research team aboard the R/V Ocean Researcher I and the
F/V Fishery Researcher I; K. Matsuura and G. Shinohara (NMST); H. Imamura, K. Nakaya,
M. Yabe, K. Amaoka (HUMZ); C-W. Chang, S.-I. Wang and R.-R. Chen (NMMBA); M.-Y. Lee,
P.-L. Lin, Y.-C. Liao and P.-H. Kao (ASIZP); H. Endo, T. Yamakawa and R. Asaoka (BSKU),
Y. Kai (FAKU) for curatorial assistance and hospitality; J. E. McCosker, D. Catania, J. Fong, and
M. Hoang (CAS). This study is under the project “Study on the Deep-Sea Fish Biodiversity of Tai-
wan” supported by the National Science Council to K-T Shao.
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Station/Location Longitude and Latitude Region Date Depth (m)
CP 119 122°01ʹE, 24°56ʹN NET 7/31/01 123–140
CP 120 122°02ʹE, 24°51ʹN NET 7/31/01 520–640
CP 124 122°17ʹE, 24°58ʹN NET 8/1/01 1129–1165
CP 127 121°03ʹE, 22°08ʹN SET 8/21/01 1263–1268
CP 130 120°07ʹE, 22°19ʹN SCS 8/22/01 709–728
CD 133 120°08ʹE, 22°15ʹN SCS 11/21/01 690–748
CD 134 120°06ʹE, 22°16ʹN SCS 11/22/01 736–1040
CD 136 120°00ʹE, 22°07ʹN SCS 11/22/01 998–1211
CD 137 120°25ʹE, 22°12ʹN SWT 11/23/01 316–477
CD 138 120°20ʹE, 22°13ʹN SWT 11/23/01 441
CD 139 120°14ʹE, 22°10ʹN SWT 11/23/01 718–852
CD 140 120°22ʹE, 22°11ʹN SWT 11/23/01 280–452
CD 141 119°59ʹE, 22°12ʹN SCS 11/24/01 985–1110
CD 142 120°13ʹE, 22°21ʹN SWT 11/24/01 227–335
CP 178 119°55ʹE, 22°25ʹN SCS 8/25/02 1241
CD 191 118°22ʹE, 21°41ʹN SCS 8/28/02 1621–1630
CD 192 120°01ʹE, 22°17ʹN SWT 8/29/03 1305
CD 193 120°06ʹE, 22°22ʹN SWT 8/29/02 821
CD 194 120°24ʹE, 22°11ʹN SET 8/29/02 507
CP 195 122°03ʹE, 24°52ʹN NET 9/11/02 570
CP 196 122°03ʹE, 24°51ʹN NET 9/11/02 646–787
CP 197 122°17ʹE, 24°51ʹN NET 9/11/02 1040–1141
CD 199 122°12ʹE, 24°25ʹN ET 9/12/02 1134–1188
CD 203 120°28ʹE, 22°00ʹN SCS 5/29/03 634–866
CD 209 122°11ʹE, 24°40ʹN NET 5/30/03 508–522
CD 210 122°12ʹE, 24°28ʹN ET 5/31/03 445–1185
CD 211 122°11ʹE, 24°40ʹN NET 8/26/03 517–529
CD 214 122°12ʹE, 24°28ʹN ET 8/27/03 488–1027
CP 214 122°13ʹE, 24°29ʹN NET 8/27/03 488
CD 226 121°04ʹE, 22°18ʹN SET 8/29/03 1174–1212
CD 228 121°01ʹE, 22°08ʹN SET 8/30/03 1262–1290
CD 229 120°01ʹE, 22°13ʹN SWT 8/30/03 880–1062
CD 230 120°03ʹE, 22°19ʹN SWT 8/30/03 810–850
CD 233 120°19ʹE, 22°11ʹN SWT 8/31/03 448–526
CP 234 122°31ʹE, 25°22ʹN NET 7/22/04 547
CP 235 122°43ʹE, 25°23ʹN NET 7/23/04 764
TABLE 1. Collection data for station of research vessels and commercial trawls operating out of nine
fishing ports. Abbreviations: CD—otter trawl station; OCP—ORE-type beam trawl station and
CP—the French type beam trawl station.
Appendix

126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
Station/Location Longitude and Latitude Region Date Depth (m)
CP 242 122°29ʹE, 25°08ʹN NET 7/23/04 979
CP 247 122°02ʹE, 24°52ʹN NET 8/28/04 480
CP 248 122°02ʹE, 24°51ʹN NET 8/28/04 536
CP 250 122°04ʹE, 24°55ʹN NET 8/28/04 220
CD 271 120°.8ʹE, 22°19ʹN SWT 12/28/04 700–800
CP 299 122°03ʹE, 22°19ʹN SET 8/11/05 799
OCP 301 120°06ʹE, 22°20ʹN SET 8/11/05 687
OCP 302 120°06ʹE, 22°21ʹN SET 8/11/05 695
OCP 303 120°15ʹE, 22°10ʹN SWT 8/15/05 807
CD 307 118°14ʹE, 21°35ʹN SCS 8/16/05 1591
CD 310 117°17ʹE, 21°35ʹN SCS 8/17/05 364
CD 311 117°43ʹE, 21°40ʹN SCS 8/17/05 516
OCP 312 117°43ʹE, 21°40ʹN SCS 8/17/05 517
OCP 313 117°43ʹE, 21°40ʹN SCS 8/17/05 513
CP 314 117°43ʹE, 21°40ʹN SCS 8/17/05 506
CP 315 117°43ʹE, 21°40ʹN SCS 8/17/05 509
CP 316 117°43ʹE, 21°40ʹN SCS 8/17/05 514
OCP 317 117°43ʹE, 21°40ʹN SCS 8/17/05 515
CD 320 117°27ʹE, 20°50ʹN SCS 8/17/05 731
CD 321 117°33ʹE, 20°43ʹN SCS 8/19/05 954
CD 322 117°39ʹE, 20°44ʹN SCS 8/19/05 1098
CD 324 117°45ʹE, 20°40ʹN SCS 8/20/05 1293
CD 325 118°03ʹE, 20°40ʹN SCS 8/20/05 1982
CP 338 120°20ʹE, 22°10ʹN SCS 3/7/06 569
CP 339 120°15ʹE, 22°10ʹN SCS 3/7/06 846
CP 347 120°13ʹE, 22°25ʹN SCS 3/9/06 305
CP 348 120°12ʹE, 22°22ʹN SCS 3/9/06 395
CP 350 121°08ʹE, 22°21ʹN SET 6/2/06 1148
CP 353 121°04ʹE, 22°15ʹN SET 6/2/06 1205
CP 366 121°10ʹE, 22°01ʹN SET 8/24/06 1302
Da-xi 121°54ʹE, 24°56ʹN NET ca. 100–650
Diao-yu-tai 123°30ʹE, 25°43ʹN NET ca. 100–400
Hsiao-liou-chiou 120°22ʹE, 22°20ʹN SWT ca. 100–400
Fong-kang 120°41ʹE, 22°11ʹN SWT ca. 100–200
Jin-shan 121°55ʹE, 25°33ʹN NT ca. 100
Lyu-dao 121°29ʹE, 22°38ʹN ET ca. 400
Nan-fang-ao 121°52ʹE, 24°35ʹN NET ca. 100–600
Dong-gang 120°26ʹE, 22°27ʹN SWT ca. 100–400
Tong-sa Islands 116°51ʹE, 20°41ʹN SCS ca. 100–600
TABLE 1. Continued.

IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN
A
Abyssicola 61, 62
Asthenomacrurus 85
Australia 34, 38, 46, 50, 52, 62, 67, 68, 73, 80,
94, 95, 98, 99, 100, 101, 103, 104, 112, 114,
115, 117, 118
B
Bathygadidae 31, 34, 36, 37
Bathygadus 31, 36, 37, 38, 39, 40, 41
antrodes 31, 37, 38, 40, 41
bowersi 38
entomelas 41
furvescens 31, 37, 38, 39
garretti 31, 37, 38, 39, 40, 41
nipponicus 31, 37, 39, 40, 41
spongiceps 38, 41
C
Caelorinchus 50, 51, 52, 54, 57, 58, 59, 60, 63,
65, 66, 67
California 69
Cetonurus 31, 45, 46
crassiceps 46
globiceps 31, 45, 46, 47
Coelocephalus 87
acipenserineus 87
Coelorhynchus 50, 51, 52, 54, 56, 57, 58, 59, 60,
61, 63, 65, 66, 67, 68
Coelorinchus 31, 34, 36, 44, 47, 48, 50, 51, 52,
53, 54, 55, 56, 57, 58, 59, 60, 61, 62, 63, 64,
65, 66, 67, 68, 74
abbreviatus 54, 55
anatirostris 31, 34, 49, 50, 51, 62, 67
argus 64
asteroides 31, 49, 50, 51
brevirostris 31, 48, 51, 52, 56
cingulatus 31, 48, 52, 56
cylindricus 36
divergens 31, 33, 49, 53, 54, 107, 108, 109,
116, 117
dorsalis 64, 116
formosanus 31, 48, 54, 55, 58
fuscigulus 32, 36, 48, 52, 55
hexafasciatus 32, 34, 36, 49, 56
hubbsi 32, 48, 56, 57, 59
intermedius 54, 55, 118
japonicus 32, 49, 57, 58, 68, 80, 98
jordani 59, 64
kamoharai 32, 48, 58, 61
kermadecus 54, 65
kishinouyei 32, 49, 59, 62, 64
leptorhinus 32, 36, 49, 59, 60, 68
longissimus 32, 48, 58, 60
macrochir 32, 48, 61, 62
macrolepis 64
macrorhynchus 32, 34, 36, 49, 60, 62, 63,
68
maculatus 64
multispinulosus 32, 48, 58, 63, 64
notatus 32, 34, 36, 49, 64, 65
parallelus 32, 49, 54, 65, 66
productus 32, 34, 49, 50, 62, 66, 67
sexradiatus 64
sheni 32, 36, 49, 56, 67
smithi 32, 49, 62, 67, 68, 69
spinifer 32, 36, 49, 68
thurla 64
tokensis 60
triocellatus 64
velifer 64
Coelorincus 53
Coryphaenoides 32, 34, 36, 44, 69, 70, 71, 73,
74, 87, 88, 93, 104, 109, 112
acrolepis 69
armatus 69
asper 32, 36, 69, 112
marginatus 71, 72
microps 32, 69, 70, 71, 72
misakius 112
nasutus 32, 69, 71, 72, 93
rudis 32, 69, 73, 74
rupestris 34, 69
yaquinae 69
E
East China Sea 50, 51, 53, 54, 55, 56, 58, 61, 62,
64, 66, 72, 75, 76, 82, 87, 91, 93, 113, 116
Index

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
Diao-yu-tai Archipelago 55
Diaoyutai 55
Okinawa Trough 37, 39, 42, 50, 51, 52, 53,
57, 58, 59, 60, 61, 62, 63, 64, 65, 66, 67,
75, 77, 82, 86, 87, 91, 93, 103, 110, 116
Emperor Seamounts 37, 38, 78
G
Gadiformes 31, 34
Gadomus 31, 36, 37, 38, 41, 42, 43, 44
aoteanus 41
colletti 31, 37, 41
denticulatus 43
introniger 43
magnifilis 31, 37, 42, 43
melanopterus 41
multifilis 31, 36, 37, 43, 44
H
Hawaiian Islands 46
Hoplostethus atlanticus 34
Hymenocephalus 32, 34, 36, 45, 46, 73, 74, 75,
76, 77, 78, 102, 103
lethonemus 32, 75
longibarbis 76
longiceps 32, 75
papiraceus 76
papyraceus 32, 34, 36, 75, 76, 77
striatissimus 32, 75, 77, 78
aeger 77, 78
hachijoensis 77, 78
torvus 78
Hymenogadus 32, 34, 36, 45, 74, 78, 102, 103
gracilis 32, 36, 45, 78
tenuis 78
Hyomacrurus 88
Hyostomus 33, 44, 87, 88, 89, 90
I
Indian Ocean 73, 79, 85, 86, 88, 98, 100, 106,
114
Ninety-East Ridge 98
Indo-Australian Archipelago 66
Indo-Malaysian Archipelago 84
Indonesia 37, 38, 67, 68, 84, 90, 95, 101, 106,
107, 115
Borneo 92, 108
Celebes Sea 76, 92, 107, 108
Halmahera 92, 108
Java 67
Molucca Sea 92, 107, 108
Sulawesi 68
J
Japan 37, 40, 46, 51, 53, 56, 57, 58, 59, 61, 62,
64, 65, 66, 68, 72, 76, 80, 82, 87, 91, 93, 94,
95, 103, 104, 109, 110, 111, 112, 113, 118
Hokkaido 62, 72, 118
Kyushu 62, 75, 118
Japan Sea 62, 64, 71
K
Kermadec Islands 65
Korea 71
Kumba 32, 45, 79, 80, 81
gymnorhynchus 32, 79
japonica 32, 79, 80, 98, 113
punctulata 32, 79, 80, 81
Kuronezumia 32, 46, 82, 83, 114
bubonis 82, 83
dara 32, 46, 82, 83
darus 82
macronema 114
Kyushu-Palau Ridge 42, 56, 57, 67, 71, 75, 77,
78, 79, 80, 86, 87, 91, 93, 111, 112
L
Lepidorhynchus 74
Lionurus 80, 82, 83, 84, 91, 92, 93, 94, 100, 112
darus 82
misakius 112
nigromaculatus 84
proximus 93
spinosus 94
Lord Howe Rise 47, 65
Lucigadella 83, 84
Lucigadus 32, 46, 83, 84
lucifer 84
nigromarginatus 32, 46, 83, 84
M
Macrosmia 32, 45, 84, 85, 86
phalacra 32, 45, 84, 85, 86

IWAMOTO, NAKAYAMA, SHAO, & HO: GRENADIER FISHES OF TAIWAN
Macrouridae 31, 34, 36, 44, 47, 69
Macrouroididae 33, 34, 36, 118
Macrourus 34, 47, 70, 73, 83, 84, 86, 87, 88, 93,
98, 112
berglax 34
proximus 93
Macrurus 46, 57, 61, 65, 71, 100
Malacocephalus 32, 36, 45, 86, 87
hawaiiensis 87
laevis 36, 86, 87
nipponensis 32, 36, 45, 86, 87
paradoxus 73
Malay Archipelago 43
Mataeocephalus 32, 44, 45, 87, 88, 89, 90
accipenserinus 87, 88
adustus 87, 88
cristatus 32, 88, 89
Melanobranchus 37, 39, 40
N
Nematonurus macrocephalus 73
New Caledonia 50, 52, 55, 65, 73, 80, 81, 84, 95,
103, 104, 114, 115
New Guinea 80, 98, 99
Bismark Sea 80, 99
New Zealand 34, 46, 65
Nezumia 33, 36, 46, 80, 82, 84, 90, 91, 92, 93,
94, 95, 96, 117
africana 117
atlantica 117
coheni 33, 36, 90, 95, 96, 97
condylura 33, 90, 91, 92
darus 82
evides 33, 91, 92
loricata 36
propinqua 91, 92
proxima 33, 36, 90, 93, 94
proximus 93, 94
spinosa 33, 90, 93, 94, 95
Norfolk Ridge, 46, 65
North Atlantic 69, 85, 86
North Pacific 99, 104
O
Okinawa 52, 53
P
Papyrocephalus 76
Paracetonurus 36, 98
cetonuropsis 36, 98, 99
flagellicauda 98
parvipes 98
pusillus 98
Paramacrurus 59, 64
Pawnurus 86
Philippine Islands 43, 71, 87
Philippines 37, 38, 42, 43, 46, 50, 58, 60, 61, 62,
63, 64, 66, 67, 70, 71, 73, 75, 76, 77, 78, 84,
88, 90, 92, 93, 94, 95, 100, 104, 107, 108,
114, 115, 121
Bohol Sea 76
Luzon 63, 70, 71, 77, 94, 114, 115
Verde Island Passage 63
Mindanao 42, 43, 114, 115
Mindoro 108
Panay 108
Sulu Sea 76, 108
Pseudocetonurus 33, 36, 45, 97
septifer 33, 36, 45, 97
Pseudonezumia 33, 36, 45, 85, 98, 99
pusilla 33, 36, 45, 98, 99
R
Regania nipponica 40
Ryukyu Islands 55
Okinawa Trough 50, 51, 53, 58, 61, 75, 77,
78, 115, 116, 117
S
South Atlantic 47, 85
South China Sea 35, 37, 46, 51, 52, 54, 59, 61,
66, 68, 70, 72, 73, 75, 76, 77, 79, 81, 82, 83,
87, 88, 90, 91, 92, 93, 94, 96, 99, 104, 108,
111, 112, 113, 114, 115, 117, 118
Sphagemacrurus 33, 45, 99, 100, 101, 102
decimalis 101
pumiliceps 33, 100, 101, 102
richardi 33, 100, 101, 102
Spicomacrurus 33, 34, 36, 45, 74, 78, 102, 103
kuronumai 33, 36, 45, 78, 102, 103
Squalogadus 33, 118, 119
modificatus 33, 118, 119
Steindachneria 74

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 62, No. 3
T
Taiwan 31
Teleostei 31
Trachonurus 33, 45, 103, 104, 105, 106
robinsi 104
sentipellis 33, 103, 104, 105
villosus 33, 103, 104, 106
V
Vanuatu 46, 50, 52, 65, 81, 85, 86, 115
Ventrifossa 33, 34, 36, 46, 80, 83, 84, 97, 105,
106, 107, 109, 110, 111, 112, 113, 114, 115,
116, 117
atherodon 111, 112
fusca 112, 113, 114
garmani 33, 107, 109, 110, 111
johnboborum 113, 114
longibarbata 33, 107, 109, 110, 111, 116
macroptera 33, 107, 111, 112, 116, 118
misakia 33, 34, 36, 105, 106, 107, 112, 113,
114
nigrodorsalis 33, 107, 110, 114, 115
nigromarginata 84
rhipidodorsalis 33, 107, 109, 115, 116
saikaiensis 33, 107, 108, 109, 116, 117
sazonovi 33, 34, 36, 107, 112, 117
Vietnam 114, 117