The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 87–146
Review of the Dinematichthyini (Teleostei: Bythitidae) of the Indo-west Paciic.
Part IV. Dinematichthys and two new genera with descriptions of nine new species
PETER RASK MØLLER1 And WERnER SCHWARZHAnS2
1
Natural History Museum of Denmark, Zoological Museum, University of Copenhagen,
Universitetsparken 15, DK-2100 Copenhagen Ø, DENMARK
pdrmoller@snm.ku.dk
2
Ahrensburger Weg 103 D, 22359 Hamburg, GERMANY
wwschwarz@aol.com
ABSTRACT
A revision of the dinematichthyine ishes (Ophidiiformes: Bythitidae: Brosmophycinae) of the Indo-West Paciic based
on more than 6500 specimens is published in several parts. Part IV is the last part and includes 4719 identiied specimens
in the genera Alionematichthys (new genus with three described and eight new species), Dinematichthys Bleeker, 1855
(with one described and one new species) and Porocephalichthys (new genus with one described species). A neotype
of Dinematichthys iluocoeteoides Bleeker, 1855 is here designated. The genera reviewed here have in common a high
anterior nostril and are considered to be related to each other. When previously described, they were all included in
the genus Dinematichthys. The separating characters of the species are the pseudoclasper morphology, morphometric
characters, vertebrae and in ray counts, otolith morphology, head squamation, presence or absence of the upper
preopercular pore and development of cirri on the snout.
KEyWoRdS: Viviparous brotulas, Indo-west Paciic, Dinematichthys, new species, new genera.
CONTENTS
INTRODUCTION .......................................................................................................................88
MATERIAL AND METHODS ...................................................................................................88
COMPARATIVE MATERIAL....................................................................................................89
SYSTEMATICS ..........................................................................................................................89
Tribe Dinematichthyini Cohen and Nielsen, 1978 ................................................................90
Key to the genera and the species reviewed herein ...............................................................89
Alionematichthys gen. nov. ....................................................................................................90
Alionematichthys ceylonensis sp. nov...............................................................................92
Alionematichthys crassiceps sp. nov. ..............................................................................95
Alionematichthys minyomma (Sedor and Cohen, 1987) ..................................................99
Alionematichthys phuketensis sp. nov. ..........................................................................100
Alionematichthys piger (Alcock, 1890)..........................................................................102
Alionematichthys plicatosurculus sp. nov. ....................................................................109
Alionematichthys riukiuensis (Aoyagi, 1954) ................................................................113
Alionematichthys samoaensis sp. nov. ..........................................................................117
Alionematichthys shinoharai sp. nov. ............................................................................120
Alionematichthys suluensis sp. nov. ..............................................................................122
Alionematichthys winterbottomi sp. nov. ......................................................................125
Alionematichthys sp. 1 ...................................................................................................126
Alionematichthys sp. 2 ....................................................................................................127
Dinematichthys Bleeker, 1855 .............................................................................................128
Dinematichthys iluocoeteoides Bleeker, 1855 ...............................................................129
Dinematichthys trilobatus sp. nov. ................................................................................136
Porocephalichthys gen. nov. ...............................................................................................138
Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982).....................................138
DISCUSSION ............................................................................................................................141
REGIONAL CHECK LIST .......................................................................................................142
KEY TO THE GENERA OF THE DINEMATICHTHYINI ...................................................144
ACKNOWLEDGMENTS .........................................................................................................145
REFERENCES ..........................................................................................................................145
W. Schwarzhans and P. R. Møller
diverse in most instances. Other useful characters for
species differentiation are found to be head squamation,
vertebrae and in ray counts, morphometric measurements,
otolith morphology, presence or absence of the upper
preopercular pore and, new in this group, development of
cirri on the snout. Of these, meristic counts show a lower
degree of variation than in most other Dinematichthyini, and
although individual species show statistically signiicant
differences (average values), they are rarely separated in
absolute values to be of use in diagnostic keys.
Eight species have been described previously, as placed
in the genus Dinematichthys, of which ive are here regarded
as valid, and represent the three genera dealt with in this
part of the review. Three previously described species
are placed in synonymy. Dinematichthys Bleeker is the
earliest named genus of its tribe and hence, its deinition is
essential to achieving nomenclatural stability. The genus
was established by monotypy (D. iluocoeteoides) based
on a single type specimen, which according to extensive
research by Cohen and Nielsen (1978) must be regarded
as no longer extant. For the purpose of redeinition of the
genus and species, another original Bleeker specimen
mentioned by Günther (1862) (BMNH 1868.2.28.65),
probably the oldest extant, is here selected as neotype.
Although shrivelled, it represents a male specimen with
all the important diagnostic characters, and is in good
accordance with Bleeker’s original description except for
a slight variation in eye size, which is within the range of
variability observed in the species.
INTRODUCTION
The global review of the dinematichthyine ishes, a tribe
within the subfamily Brosmophycinae of the viviparous
family Bythitidae, is concluded with this sixth part. The irst
two parts dealt with the American Dinematichthyini (Møller
et al. 2004a, 2005) and were followed by four publications
revising the Dinematichthyini from the Indo-west Paciic
(Schwarzhans et al. 2005; Møller and Schwarzhans 2006;
Schwarzhans and Møller 2007; this paper). The latest
comprehensive account of ophidiiform ishes (Nielsen et
al. 1999) contained 12 genera now regarded as valid in the
Dinematichthyini, and 25 valid species. With the completion
of the world-wide review of the Dinematichthyini, the
current status of recognised taxa within the tribe has risen
to 26 genera with 110 species.
The ishes described in this paper were previously
referred to the genus Dinematichthys. Following Cohen
and Nielsen (1978) and redeined by Cohen and Hutchins
(1982) and Sedor and Cohen (1987), the position of the
anterior nostril high above the upper lip, and about midway
between the lip and the posterior nostril, was regarded as
the main diagnostic character for the genus, distinguishing
it from all other than known dinematichthyine ishes. In this
review we conclude that the formal genus Dinematichthys
“sensu lato” in fact contains three well deined groups for
which generic rank applies: two more closely related ones –
Dinematichthys with two species and Alionematichthys gen.
nov. with 11 species – and one more distant monospeciic
genus containing Porocephalichthys gen. nov. dasyrhynchus
(Cohen and Hutchins). The combining character of the high
anterior nostril, reformulated to “positioned 1/3 or less of
the distance from upper lip to anterior margin of eye” in
Schwarzhans and Møller (2007), however, is not unique
to these three genera and this distinguishing character has
therefore become weakened. Similarly, high positioned
anterior nostrils between 1/3 and 1/3.5 the distance from
upper lip to anterior margin of eye have been observed in
a few species of the genus Diancistrus Ogilby (for instance
D. alleni, D. atollorum and D. katrineae), in Eusurculus
pistillum Schwarzhans and Møller and in Lapitaichthys
frickei Schwarzhans and Møller. Fishes of the three
genera Dinematichthys, Alionematichthys gen. nov. and
Porocephalichthys gen. nov. can be easily distinguished
from Diancistrus by the otolith sulcus showing distinctly
separated colliculi (vs fused, undivided), from Lapitaichthys
in the presence of two pairs of pseudoclaspers (vs single
pair of outer pseudoclaspers), and from Eusurculus
pistillum in the lap-like inner pseudoclasper (vs stalked
with sucker-disk tip) (see also key below). The diagnostic,
distinguishing characters in the species of the three genera
covered here are similar to those of the previous parts of
our review. Again, pseudoclasper morphology represents
the most useful character. However, pseudoclaspers in
the following three genera are smaller than in most other
Dinematichthyini, with few exceptions, and are also less
MATERIAL AND METHODS
88
The examination of ca. 6500 specimens of Indo-west
Paciic Dinematichthyini yielded 4719 specimens which
were identiied in the genera treated herein. Also included
are additional specimens identiied in the collections of AMS
and USNM but not borrowed for detailed investigations.
These are listed as additional specimens and are not referred
to as type specimens for any of the new species.
The material described herein belongs to the following
institutions: AMS (Australian Museum, Sydney), ANSP
(Academy of Natural Sciences, Philadelphia), BMNH
(Natural History Museum, London), BPBM (Bishop
Museum, Honolulu), CAS (California Academy of
Sciences, San Francisco), KAUM (The Kagoshima
University Museum), KSHS (Kochi Prefectural Kochi
Nishi Senior High School, Kamobe, Kochi, Japan), MNHN
(Muséum National d’Histoire Naturelle, Paris), NSMT
(National Science Museum, Tokyo), NTM (Museum and
Art Gallery of the Northern Territory (formerly Northern
Territory Museum), Darwin), ROM (Royal Ontario
Museum, Toronto), SAIAB (South African Institute for
Aquatic Biodiversity, formerly RUSI (JLB Smith Institute
of Ichthyology), Grahamstown), SMF (Senckenberg
Forschungsinstutut und Museum, Frankfurt/Main), SMNS
Dinematichthyine ishes of the Indo-west Paciic, Part IV
and on the species reviewed by Schwarzhans and Møller
(2007). The irst dichotomy in that key refers to the anterior
nostril position and immediately separates the three genera
dealt with herein from all other dinematichthyine genera.
Since, however, certain other dinematichthyine genera exist
with almost similarly high anterior nostrils we begin the
following key with a separation from those few genera.
(Staatliches Museum für Naturkunde, Stuttgart), TAU
(Tel Aviv University, Tel Aviv, Israel); USNM (National
Museum of Natural History, Smithsonian Institution,
Washington D.C.), WAM (Western Australian Museum,
Perth), YCM (Yokosuka City Museum), and ZMUC
(Zoological Museum, University of Copenhagen).
For methodology used in analysing dinematichthyine
ishes, reference is made to Møller et al. (2004a) and
Schwarzhans et al. (2005). Abbreviations used in meristic
counts are: D/V = anterior dorsal in ray above vertebra
number; D/A = anterior anal fin ray below dorsal fin
ray number; V/A = anterior anal in ray below vertebrae
number; D-A = number of dorsal in rays minus number
of anal in rays; V in D = number of dorsal in rays per raybearing vertebra. The abbreviations ‘i.p.’ and ‘o.p.’ used in
the igures relating to male copulatory organ structures are
explained in Schwarzhans et al. (2005).
The ecology of most of the species is poorly known.
From available station data we have gathered some
information about habitat and depth range, but we have very
little data about behaviour, live colouration and feeding. A
number of females were examined for reproductive data,
e.g. number and size of embryos.
The distribution maps were created using Microsoft
Encarta 2001 digital world atlas.
1a. Anterior nostril positioned high (equal or less than
1/3 the distance from upper lip to anterior margin of
eye) ......................................................................... 2
1b. Anterior nostril positioned low (1/4 to 1/6 the distance
from upper lip to anterior margin of eye) ................
all other Dinematichthyini (see Schwarzhans and
Møller (2007))
2a. Single pair of (outer) pseudoclaspers .......................
............................................................. Lapitaichthys
2b. Pair of inner and outer pseudoclaspers ................... 3
3a. Sulcus of otolith with undivided colliculi ................
................................................................ Diancistrus
[note: only a few species of the genus exhibit an
elevated anterior nostril]
3b. Sulcus of otolith with separated colliculi (not known
for Alionematichthys samoensis) ............................ 4
COMPARATIVE MATERIAL
4a. Inner pseudoclasper stalked with sucker-disk shaped
tip .............................................Eusurculus pistillum
[note: the only species in Eusurculus with an elevated
anterior nostril]
4b. Inner pseudoclasper variable, lap or stick-shaped or
folded, but not stalked and without sucker-disk shaped
tip ............................................................................ 5
Indo-west Paciic Dinematichthyini: see Schwarzhans
et al. (2005), Møller and Schwarzhans (2006) and
Schwarzhans and Møller (2007).
American Dinematichthyini: see Møller et al. (2004a)
and Møller et al. (2005).
Brosmophycinae and Bythitinae: see Møller et al.
(2004b).
5a. Head with continuous squamation on cheeks, opercle
and occiput .............................................................. 6
[note: juveniles of Dinematichthys may have a
scaleless gap between cheeks and opercle]
5b. Head with scales on cheeks and in some species with
scale patch on opercle above opercular spine, rarely
also below opercular spine (Alionematichthys) ..... 8
SYSTEMATICS
Family Bythitidae Gill, 1861
Subfamily Brosmophycinae Gill, 1862
Tribe Dinematichthyini Cohen and Nielsen, 1978
Diagnosis. Male copulatory organ with penis and 1–2
(rarely 3) pairs of pseudoclaspers in cavity of ventral body
wall covered by leshy hood. First anal in pterygiophore
slightly to strongly elongate. Head pore system generally not
reduced, 6 mandibular, 2–4 preopercular, 5–7 infraorbital
and 3–4 supraorbital pores, including supraorbital pore
above opercular spine. Posteriormost supraorbital headpore tubular.
Table 1 summarises meristic characters used in
distinguishing the species of the genera described here.
Key to the genera and the species reviewed herein
The following key to the genera Dinematichthys,
Alionematichthys gen. nov. and Porocephalichthys gen. nov.
is based on the key to the genera of the Dinematichthyini,
89
6a. Upper preopercular pore present, a pair of additional
mandibular pores, three additional pairs of pores on
occiput, one behind eye, two above preopercular pore;
sulcus of otolith with nearly equally long ostium and
cauda; precaudal vertebrae 14, total vertebrae 47–49,
dorsal in rays 96–104, D/A 37–43; canine teeth
absent ...................Porocephalichthys dasyrhynchus
6b. Upper preopercular pore absent (see Schwarzhans
et al. 2005, ig. 1A), no additional mandibular pores,
no pores on occiput; sulcus of otolith with ostium at
least two times as long as cauda; precaudal vertebrae
11–12, total vertebrae 41–45, dorsal in rays 75–92,
D/A < 30; canine teeth present (Dinematichthys)... 7
W. Schwarzhans and P. R. Møller
7a. Inner pseudoclasper with 2 lobes; head squamation
not extending beyond tip of maxilla .........................
................................. Dinematichthys iluocoeteoides
7b. Inner pseudoclasper with 3 lobes; head squamation
extending beyond tip of maxilla forward of 3rd
posterior mandibular pore .........................................
............................Dinematichthys trilobatus sp. nov.
14b. Outer pseudoclasper thin, without knob; posterior
nostril with anterior flap, not funnel-shaped
..............................................Alionematichthys piger
15a. Scales above opercular spine 2; sulcus of otolith
with extremely small cauda (ostium length to cauda
length 5.5–7, ostium height to cauda height 2.2–2.7)
....................... Alionematichthys shinoharai sp. nov.
15b. Scales above opercular spine absent; sulcus of otolith
with moderately small cauda (ostium length to cauda
length 3–4, ostium height to cauda height 1.5–2.0) .
............................................................................... 16
8a. Upper preopercular pore present on both head sides;
cirri on snout present or absent ............................... 9
8b. Upper preopercular pore absent (rarely present on one
head side only); cirri on snout absent ................... 15
9a. Cirri on snout absent, eye small (usually < 2% SL) .
............................................................................... 10
9b. Cirri on snout present (except absent in Alionematichthys
suluensis sp. nov.), eye relatively large (usually > 2%
SL)......................................................................... 11
16a. Inner pseudoclasper thick, with posterior lobe
folded over anterior thorn; outer pseudoclasper with
thickened tip ..............................................................
................ Alionematichthys plicatosurculus sp. nov.
16b. Inner pseudoclasper thin, unfolded; outer pseudoclasper
with thin or with knob on inner face..................... 17
10a. Scales above opercular spine 3–10; inner pseudoclasper
with anterior thorn as long as posterior lobe; otolith
length to otolith height 1.8–2.0, sulcus without
ventral indentation at junction of ostium and cauda
........................Alionematichthys crassiceps sp. nov.
10b. Scales above opercular spine absent; inner
pseudoclasper with anterior thorn shorter than
posterior lobe; otolith length to otolith height 2.2–2.3,
sulcus with ventral indentation at junction of ostium
and cauda ................... Alionematichthys minyomma
17a. Inner pseudoclasper with a symmetrically developed
anterior thorn and posterior lobe resembling in shape
a socket itting to a small knob on the inner face of the
outer pseudoclasper; snout rounded; otolith length to
height 2.3–2.4, with shallow dorsal rim ...................
...................... Alionematichthys ceylonensis sp. nov.
17b. Inner pseudoclasper a simple lap without signiicant
indentation; snout pointed; otolith length to height
2.1–2.3, with distinct postdorsal projection..............
......................Alionematichthys phuketensis sp. nov.
11a. Scales above opercular spine 5–17; pseudoclaspers
not extruding beyond hood in resting position ..... 12
[note: scales 2–6 in few tentatively assigned specimens
– Alionematichthys aff. riukiuensis]
11b. Scales above opercular spine 0–2; pseudoclaspers
usually extruding beyond hood in resting position
............................................................................... 13
Alionematichthys gen. nov.
(Tables 1–12)
Type species: Dinematichthys riukiuensis Aoyagi, 1954
(type locality: Ishigaki Island, Ryukyu Islands, Japan).
Gender masculine.
Dinematichthys (non Bleeker, 1855) in part. — Alcock
1890: 432; Aoyagi 1954: 237; Cohen and Nielsen 1978: 57;
Sedor and Cohen 1987: 6; Machida 1994: 451; Nielsen et
al. 1999: 129; Møller et al. 2004a: 148.
Diagnosis. A genus of Dinematichthyini with the
following combination of characters: anterior nostril
placed high on snout at about one-third or less the distance
from lip to anterior rim of eye; head with a scale patch
on cheeks, in some species also above opercular spine,
rarely below opercular spine; male copulatory organ
with two pairs of mostly small pseudoclaspers, the outer
up to twice as large as inner; inner pseudoclasper with
anteriorly oriented pointed lobe containing supporter and
leshy appendix posteriorly; large size, up to 150 mm SL;
precaudal vertebrae 10–12, total vertebrae 39–45, dorsal
in rays 71–92, anal in rays 55–72, V in D 2.0–2.4; upper
preopercular pore present or, less commonly, absent; sulcus
of otolith with ostium at least twice as long as cauda; maxilla
expanded posteroventrally.
12a. Inner pseudoclasper thick, large, anterior thorn
entirely fused to anterior rim of outer pseudoclasper;
scale patch on opercle below opercular spine in large
specimens ................... Alionematichthys riukiuensis
12b. Inner pseudoclasper thin, small, bifurcate,
anterior thorn free from outer pseudoclasper;
no scale patch on opercle below opercular spine
................. Alionematichthys winterbottomi sp. nov.
13a. Cirri on snout absent; anterior thorn of inner
pseudoclasper very long, twice as long as posterior
lobe ...................Alionematichthys suluensis sp. nov.
13b. Cirri on snout present; anterior thorn of inner
pseudoclasper about as long as posterior lobe ...... 14
14a. Outer pseudoclasper with thick knob on inner face
distally; posterior nostril funnel-shaped ...................
...................... Alionematichthys samoaensis sp. nov.
90
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Table 1. Selected meristic characters of the species of Alionematichthys, Dinematichthys and Porocephalichthys: A, frequency distribution
of precaudal vertebrae, total vertebrae and anal in ray counts; B, frequency distribution of dorsal in ray and pectoral in rays counts.
A
precaudal
total vertebrae
anal in rays
vertebrae
10 11 12 13 14 39 40 41 42 43 44 45 46 47 48 49 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72
Alionematichthys
2 64
ceylonensis
Alionematichthys
1 41
crassiceps
Alionematichthys
4 15
minyomma
Alionematichthys
9
phuketensis
Alionematichthys
309 25
piger
Alionematichthys
1 27
plicatosurculus
Alionematichthys
80 41
riukiuensis
Alionematichthys
25 8
samoaensis
Alionematichthys
2
shinoharai
Alionematichthys
15 14
suluensis
Alionematichthys
25 1
winterbottomi
Alionematichthys
2
sp. 1
Alionematichthys
1
sp. 2
Dinematichthys
2 147 5
iluocoeteoides
Dinematichthys
15
trilobatus
Porocephalichthys
dasyrhynchus
B
8 51 7
4 30 8
3 16
1
2
1
1
4 12 16 22 5
4
-
1
7
2
5 10 9
6
1
6
6
3
1
1
1
1
1
2
7
5 42 206 73 9
2
2 21 4
1
2
-
2
2 29 1
1
3
1
1
1
8
3
5 14 16 6 14 12 10 13 4
4
2
5
1
8 10 3
2
3
1
3
6
-
2
-
1
2
1
2
1
2
3
4
3
3 17 5
1
5
6
1
1 24 4
1
2
5 10 23 41 57 51 46 38 19 8
1 39 62 18
1
2
2
7
6
2
1
1
3
6
1 13
1
1
1
1
1
12 73 46 14 8
1
5 10
9
2
4
3
6
9 14 21 23 23 14 10 9
1
2
1
3
1
2
1
dorsal in rays
3
4
1
3
1
3
1
pectoral in rays
71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 19 20 21 22 23 24 25 26 27 28
Alionematichthys
4 - 10 15 15 9 8 3 2
ceylonensis
Alionematichthys
1 4 7 5 8 5 5 1 4 1
crassiceps
Alionematichthys
1 - 1 9 2 5 - - 1
minyomma
Alionematichthys
2 2 3 - - 1
phuketensis
Alionematichthys
1 - 2 9 21 28 34 50 42 42 41 18 7 6 1
piger
Alionematichthys
1 2 1 4 4 8 5 2 1
plicatosurculus
Alionematichthys
1 3 7 4 12 8 13 16 10 12
riukiuensis
Alionematichthys
2 3 5 6 7 3 4 3
samoaensis
Alionematichthys
1 - 1
shinoharai
Alionematichthys
1 - 1 2 3 8 5 2 5 1 1
suluensis
Alionematichthys
2 7 4 2 6 2 1
winterbottomi
Alionematichthys
1 - sp. 1
Alionematichthys
1
sp. 2
Dinematichthys
1 - 1 5 5 10 15 23 14 24 12 13
iluocoeteoides
Dinematichthys
5 1 2 1 3 1 trilobatus
Porocephalichthys
dasyrhynchus
1 4 5 4
2 1 4 2
5 7 1
- - 1
2 4 1
1 1
2 1 5 1
1 7 3
4 7 7 2 - 1
5 - 3 2 1
1 3 8
2
3 5 3 1
2 4 5
1
1 - 1
1
6 5 - 1 1 1
2 3 8 2
2
1 7 5 1
2 - 2 1 1
91
1
-
1
1
3 4 3 3
W. Schwarzhans and P. R. Møller
Comparison. Alionematichthys is clearly related to
Dinematichthys, from which it mainly differs by lacking
a uniform and continuous head squamation, including the
cheeks, opercle and occiput (vs scale patches on the cheeks
and, in some species, on the opercle). The head squamation
of Dinematichthys (and Porocephalichthys gen. nov.) is
unique within the Dinematichthyini and it supports the
stability of generic diagnoses in the tribe. Dinematichthys
cannot easily be separated from Alionematichthys without
that character.
Species. Alionematichthys comprises 11 species; 10
from the Indo-west Pacific – A. ceylonensis sp. nov.,
A. crassiceps sp. nov., A. phuketensis sp. nov., A. piger
(Alcock, 1890) (formerly placed in Dinematichthys),
A. plicatosurculus sp. nov., A. riukiuensis (Aoyagi, 1954)
(formerly placed in Dinematichthys with Dinematichthys
megasoma Machida, 1994, as a junior synonym),
A. samoaensis sp. nov., A. shinoharai sp. nov., A. suluensis
sp. nov., A. winterbottomi n. sp., and one species from the
tropical West Atlantic – A. minyomma (Sedor and Cohen,
1987) (formerly placed in Dinematichthys).
The species of the genus Alionematichthys can be
arranged in three species groups according to the presence
or absence of the upper preopercular pore, the presence
or absence of scales on the opercle, differences in the eye
size, and the presence or absence of cirri on the snout as
follows:
1) Alionematichthys plicatosurculus sp. nov. species
group: upper preopercular pore absent, no cirri on snout,
large eyes and no scales on the opercle, except for
A. shinoharai sp. nov. (two scales above opercular spine).
This group contains four species (A. ceylonensis sp. nov.,
A. phuketensis sp. nov., A. plicatosurculus sp. nov. and
A. shinoharai sp. nov.). However, strong differences in
the pseudoclasper and otolith morphology of some species
indicate that these species may not be closely related.
2) Alionematichthys riukiuensis species group: upper
preopercular pore present, numerous cirri on snout (except
cirri lacking in A. suluensis sp. nov.) and large eyes.
Presence or absence of scales on the opercle could be used to
separate two subgroups, one with ive or more scales above
the opercular spine (A. riukiuensis and A. winterbottomi sp.
nov.), and the other usually without scales on the opercle
(rarely up to two scales above the opercular spine) (A. piger,
A. samoaensis sp. nov. and A. suluensis sp. nov.).
3) Alionematichthys minyomma species group: upper
preopercular pore present, no cirri on snout, small eyes and
scales on opercle present or absent (A. crassiceps sp. nov.
and A. minyomma). The small eyes and the big head readily
distinguish this small group, which combines a western
Paciic species and the only Atlantic species of the genus.
Remarks. Some of the more widespread species, namely
A. crassiceps sp. nov., A. piger and A. riukiuensis, show a
remarkable regionalisation usually based on variations of
a single character.
Alionematichthys crassiceps sp. nov. and A. riukiuensis
contain few specimens, which can only be tentatively
assigned to the respective species. In the case of A. piger
the pseudoclaspers exhibit a wider than usual variation
which, however, does not seem to follow regional trends
in this particular case. The distinction of these (regional)
variations is not suficient at this stage, but it is possible that
future genetic investigations may result in distinguishing
more species than we currently recognise.
Etymology. Combined from alius (Latin = the other,
different) and nematichthys, the stem of the genus name
Dinematichthys, to which Alionematichthys is most similar.
The gender is masculine.
92
Alionematichthys ceylonensis sp. nov.
(Figs 1–3; Tables 1, 2)
Material examined. (84 specimens, 19–72 mm SL).
HOLOTYPE – USNM 263707, male, 59 mm SL, Sri Lanka,
Weligama, 500 yards south of rest house, 05°57’25’’N,
80°26’12’’E, 5–6 m, C. Koenig, 15 Feb. 1970. PARATyPES
– USNM 263731, 4 males, 42–65 mm SL, 6 females, 47–63
mm SL, Sri Lanka, Weligama; USNM 366532, 4 specimens,
37–72 mm SL, 08°33’24’’N, 81°14’42’’E, Trincomalee, Sri
Lanka, 2–5 m, T. Iwamoto, 25 Sept. 2001; USNM 394975,
25 males, 37–69 mm SL and 40 females, 37–60 mm SL,
same data as holotype.
Additional material. USNM 366531, 7 specimens,
25–65 mm SL, Sri Lanka, Triconmalee; USNM 366533, 4
specimens, 40–59 mm SL, Sri Lanka, Battocalao; USNM
366534, 6 specimens, 34–60 mm SL, 07°56’N, 81°34’E,
Sri Lanka.
Diagnosis. Vertebrae 11–12+29–31=41–43, dorsal
in rays 75–83 (average 79), anal in rays 57–65 (average
60); eyes moderately large (2.0–2.8% SL); snout rounded,
without cirri; scales only on cheeks; no upper preopercular
pore; outer pseudoclasper broad-based, triangular, with
distal knob on inner side; inner pseudoclasper with
symmetrically developed anterior pointed lobe and
posterior lobe resembling in shape a socket itting to small
knob on inner face of outer pseudoclasper; otolith length
to height 2.3–2.4, with shallow dorsal rim, otolith length
to sulcus length 1.5–1.6, ostium length to cauda length
3.3–4.0.
Description (Figs 2–3).The principal meristic and
morphometric characters are shown in Table 2. Mature at
about 45 mm SL. Body slender, with rounded head proile.
No cirri on snout. Eye size 2.0–2.8 (2.6)% SL. Head with
scale patch on cheek containing 6–8 (8) vertical rows of
scales on upper part and 3 vertical rows on lower part.
Horizontal diameter of scales on body about 1.5% SL,
in 29 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, with small knob. Anterior
nostril positioned high, 1/2.5 distance from upper lip to
anterior margin of eye. Posterior nostril small, about oneifth the size of eye. Opercular spine with free tip, thin and
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 1. Sample sites of Alionematichthys ceylonensis sp. nov., A. phuketensis sp. nov., A. shinoharai sp. nov., A. plicatosurculus
sp. nov., A. crassiceps sp. nov., and A. aff. crassiceps. One symbol may represent several samples.
Fig. 2. Alionematichthys ceylonensis sp. nov., USNM 263707, holotype, male, 59 mm SL.
sharply pointed. Anterior gill arch with 14–18 (17) rakers,
3 elongate. A single short raker is placed between the two
lower elongate rakers in holotype and several paratypes.
Pseudobranchial ilaments 2.
Head sensory pores (Fig. 3A, B). Supraorbital pores
2 to 3: irst pore in front of second anterior infraorbital
pore, second pore above and behind eye usually absent,
third pore tubular, at upper termination of gill opening
above opercular spine. Infraorbital pores 6 (3 anterior and
3 posterior): irst anterior pore behind anterior nostril,
second and third anterior pores covered by dermal lap of
upper lip, three posterior pores on rear part of upper lip,
about 1/3 the size of three anterior pores. Mandibular pores
6 (3 anterior and 3 posterior): irst anterior pore large and
tubular, second anterior pore positioned in lateral skin fold,
93
small, third anterior pore at anterior termination of jugular
isthmus, three posterior pores on rear part of lower jaw.
Preopercular pores: 3 lower, irst and second with joined
opening, covered by dermal lap in lateral view; third nontubular; no upper preopercular pore. [This description of
the position of head sensory pores is used as reference for
all subsequent descriptions.]
Dentition (of holotype). Premaxilla with 6 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.
Vomer horseshoe-shaped, with 3 outer rows of small teeth
and 1 inner row of large teeth up to 1/5 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of long teeth up to 1/4 diameter of pupil. Dentary with 5
W. Schwarzhans and P. R. Møller
Fig. 3. Alionematichthys ceylonensis sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male
copulatory organ, holotype; D, view of left pseudoclaspers from inside, holotype; E, inclined lateral view of male copulatory organ, holotype;
F, view of left pseudoclaspers from inside, USNM 394975, 68 mm SL; G, view of left pseudoclaspers from inside, USNM 394975, 48 mm
SL; H, median view of right otolith, USNM 394975, male, 58 mm SL.
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/3 diameter of pupil.
Otolith (Fig. 3H). Elongate in shape, length to height
2.3–2.4 (40–70 mm SL) and thin (otolith height to otolith
thickness about 2.5). Anterior and posterior tips moderately
pointed, the latter expanded and irregularly ornamented.
Dorsal rim shallow; ventral rim likewise shallow, deepest
anterior to middle. Inner face moderately convex, outer
face slightly concave, both smooth. Otolith length to sulcus
length 1.5–1.6. Sulcus slightly supramedialy positioned,
not inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.3–4.0. Ventral furrow distinct, very
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to
12 (11). Pleural ribs on vertebrae 2 to 11 (10). First anal in
94
pterygiophore not or only slightly longer than subsequent
pterygiophore.
Male copulatory organ (Fig. 3C–G). Two pairs of
moderately large pseudoclaspers, outer pair about 50%
larger than inner; outer pseudoclasper with broad base,
triangular with pointed tip and distinct knob at inner face
near tip; inner pseudoclasper with symmetrically developed
anterior thorn and posterior lobe resembling in shape a
socket itting to small knob on the inner face of the outer
pseudoclasper. Isthmus narrow; penis curved, about as long
as outer pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly dark brown.
Comparison. Alionematichthys ceylonensis belongs
to the species group without an upper preopercular pore
together with A. phuketensis sp. nov., A. plicatosurculus
sp. nov. and A. shinoharai sp. nov., from which it differs
in the peculiar shape of the inner pseudoclasper and the
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Etymology. Named after the type locality, Sri Lanka,
previously known as Ceylon. It is an adjective.
Table 2. Meristic and morphometric characters of Alionematichthys
ceylonensis sp. nov.
Holotype
USNM263707
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal
in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
59
Holotype +
83 paratypes
n
Mean (range)
53.9 (19-72)
84
79
15
61
19
12
29
41
17
6
21
13
SL
27.8
15.3
16.0
5.5
14.8
2.6
1.1
7.8
4.7
19.9
47.9
32.4
18.7
78.9 (75-83)
15.9 (15-17)
60.5 (57-65)
20.9 (19-22)
12.0 (11-12)
30.0 (29-31)
42.0 (41-43)
16.6 (14-18)
2
5.8 (5-6)
22.6 (20-26)
13.3 (13-15)
66
40
66
14
66
66
66
13
13
66
66
66
26.8 (25.8-27.8)
14.1 (12.1-16.1)
16.4 (14.8-17.9)
6.0 (5.4-6.8)
11.8-15.1 (14.0)
2.5 (2.0-2.8)
1.5 (1.1-1.7)
7.3 (6.3-8.0)
4.4 (3.4-4.9)
19.4 (18.5-20.7)
47.5 (45.8-51.7)
31.0 (30.0-32.5)
18.3 (14.5-20.1)
11
11
11
11
11
11
11
11
11
11
11
11
11
12.6
6.5
23.3
28.3
12.6 (11.7-13.2)
6.2 (5.2-7.2)
22.7 (19.2-24.5)
28.3 (26.7-32.8)
11
11
12
11
knob at the inner face of the outer pseudoclasper and the
slender otoliths (otolith length to otolith height 2.3–2.4 vs
2.1–2.3). From A. phuketensis sp. nov., it differs further
in the rounded snout proile (vs pointed) and the lack
of a distinct postdorsal projection of the otolith. From
A. shinoharai sp. nov., it further differs in the lack of scales
above the opercular spine (vs 2), the long sulcus (otolith
length to sulcus length 1.5–1.6 vs 1.9–2.0) and the notch at
the ventral sulcus margin at the joint of ostium and cauda
(vs no notch).
Distribution. Alionematichthys ceylonensis occurs only
along the shores of the island of Sri Lanka (Fig. 1).
Biology. A 53 mm SL gravid female (USNM 394975)
contains 80 embryos, length 4.8 mm TL, and 60 orange
eggs, diameter 0.7 mm.
95
Alionematichthys crassiceps sp. nov.
(Figs 1, 4–6; Tables 1, 3)
Material examined. 53 specimens, 34–63 mm SL.
HOLOTYPE – BPBM 9310, male, 63 mm SL, Truk,
Caroline Islands, Southfield beach, 08°0’N, 147°0’E,
inshore, 0–4 m, rock and Halimeda algae, J. E. Randall, 10
July 1969. PARATYPES – BPBM 9217, 1 male, 40 mm
SL, 1 female, 42 mm SL, Palau Islands, NE Arakabesan
Island, 0.5–1 m, corals, J. E. Randall and E. S. Helfman,
5 June 1968; BPBM 40932, 3 females, 46, 46 and 47 mm
SL, 3 males, 34, 36, 43 mm SL, same data as for holotype;
AMS I. 20547-042, 1 male, 40 mm SL, 2 females, 45 and
48 mm SL, Panasesa reef, Louisiade Archipelago, Papua
New Guinea, 5–8 m, B. Goldman, 24 March 1969; CAS
227282, 1 male, 49 mm SL, 1°4’28”N, 154°42’28”E,
0–15 m, lagoon coral heads on inner margin of reef lat at
Tewawaelal, extreme W end of atoll; low tide; water clear,
sand & coral, Kapingamarangi Atoll, Pohnpei, Caroline
Islands, Micronesia, Atta, Kindaro and H. H. Rofen, 14
July 1954; CAS 222543, 2 males, 50 and 52 mm SL, Vanua
Levu, Fiji; CAS 222563, 2 males, 40 and 47 mm SL, 1
female, 37 mm SL, Suva, Fiji; CAS 227285, 1 female,
52 mm SL, Vanua Levu, Fiji; CAS 227303, 1 female, 30
mm SL, 1 male, 40 mm SL, Suva, Fiji; USNM 223247,
1 male, 55 mm SL, 07°00’30’’N, 158°11’55’’E, Caroline
Islands, Pohnpei, off east side of Jokaj Passage, 0–4 m, V. G.
Springer and party, 5 Sept. 1980; USNM 394976, 3 females,
42, 43 and 56 mm SL, 4 males 40–57 mm SL, 2 juveniles
12 and 22 mm SL, 18°44’S, 174°06’W, Vava’u Islands,
Tonga, 0–11 m, J.T. Williams et al., 17 Nov. 1993; USNM
352743, 7 males, 48–60 mm SL, 5 females, 47–67 mm
SL, 18°44’25’’S, 169°12’41’’E, Vanuatu, Erromango, Port
Narevin, 0–5 m, J. T. Williams et al., 28 May 1996; USNM
374196, 1 male, 57 mm SL, 01°31’12’’S, 145°01’30’’E,
Papua New Guinea, Hermit Islands, Jalun Island, V. G.
Springer and party, 1 Nov. 1978.
Tentatively assigned specimens. USNM 366687, 3
males, 47, 50 and 53 mm SL, Bay SE of K’en Ting, Taiwan,
0–3 m, V. G. Springer and party, 22 April 1968.
Additional material. USNM 334123, 1 male, 49 mm
SL, Tonga; USNM 374210, 1 male, 45 mm SL, 4 females,
34–50 mm SL, Tongatapu, Tonga.
Diagnosis. Vertebrae 10–11+29–31=40–42, dorsal in
rays 72–81, anal in rays 57–64; eyes small (1.3–2.5% SL);
snout blunt, without cirri; scales on cheeks and patch above
opercular spine with 3–10 scales; upper preopercular pore
present; outer pseudoclasper broad-based, simple lap like;
inner pseudoclasper with free, forward-pointing anterior
lobe and broad posterior lobe; otolith length to height
1.9–2.0, with rounded dorsal rim, otolith length to sulcus
length 1.7–1.8, cauda very short, ostium length to cauda
length 4.5–5.0.
W. Schwarzhans and P. R. Møller
Fig. 4. Alionematichthys crassiceps sp. nov., BPBM 9310, holotype, male, 63 mm SL.
Description (Figs 4–6).The principal meristic and
morphometric characters are shown in Table 3. Mature at
about 45 mm SL. Body stout, with rounded, robust head
proile (interorbital width 6.6–8.9 (7.4)% SL). No cirri on
snout. Eye size 1.3–2.5 (1.3)% SL. Head with scale patch
on cheek containing 6–10 (10) vertical rows of scales on
upper part and 3–5 vertical rows on lower part. Horizontal
diameter of scales on body of holotype 2.1% SL, in about
26 horizontal rows. Maxillary ending far behind eye, dorsal
margin of maxillary covered by upper lip dermal lobe,
posterior end strongly expanded. Anterior nostril positioned
high, one-third distance from upper lip to anterior margin of
eye. Posterior nostril small, about one-sixth the size of eye
or less. Opercular spine with free tip, pointed. Anterior gill
arch with 16–19 (16) rakers, 3 elongate. Pseudobranchial
ilaments 1–2 (1).
Head sensory pores (Figs 5A–E, 6A,B). Supraorbital
pores 2 to 3. Infraorbital pores 6 (3 anterior and 3 posterior),
posterior pores about one-third the size of anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): irst anterior
pore large, tubular, with cirrus. Preopercular pores: 3 lower,
irst and second with joined opening, covered by dermal lap
in lateral view; third nontubular; small upper preopercular
pore. [See description of Alionematichthys ceylonensis for
position of pores.]
Dentition (of holotype). Premaxilla with 6 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 3/4 diameter of pupil.
Vomer horseshoe-shaped, with 4 outer rows of small teeth
and 1 inner row of large teeth up to 1/2 diameter of pupil.
Palatine with 4 rows of equally long teeth up to 1/2 diameter
of pupil. Dentary with 2 outer rows of granular teeth and
2 inner rows of larger teeth anteriorly, up to about size of
diameter of pupil.
Otolith (Figs 5M–P, 6E). Compact in shape, length to
height 1.9–2.0, moderately thin (otolith height to otolith
thickness about 2.5). Anterior tips moderately pointed,
posterior tip much expanded postero-dorsally. Dorsal rim
evenly curved; ventral rim deep, deepest at its middle or
slightly anteriorly. Inner face moderately convex, outer face
lat, both smooth. Otolith length to sulcus length 1.7–1.8.
Sulcus medianly positioned, not inclined, with separated
colliculi but no notch at ventral rim of sulcus at joint of
96
ostium with cauda. Cauda very small, ostium length to
cauda length 4.5–5.0. Ventral furrow distinct, very close
to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to
11 (10). Pleural ribs on vertebrae 2 to 10 (9). First anal in
pterygiophore not or only slightly longer than subsequent
pterygiophore.
Male copulatory organ (Figs 5F–L, 6C, D). Two pairs
of moderately large pseudoclaspers, outer pair about 50%
larger than inner; outer pseudoclasper with widened base
and rounded tip, lat, lap-like; inner pseudoclasper with
free, forward-pointing anterior thorn, connected at base
to anterior rim of outer pseudoclasper and broad posterior
lobe with deep inclined furrow on its inner margin. Isthmus
moderately wide; penis curved, somewhat longer than outer
pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly light brown.
Remarks. Alionematichthys crassiceps shows a high
degree of regional differentiation.
Specimens from Vanuatu (Fig. 5C) and Fiji have a
generally higher number of scales (7–10) above the opercular
spine than those from Tonga (Fig. 5D) or Micronesia (Fig.
5A, B) (5–7) on either side of the distribution range. Those
from Tonga show a peculiar pattern in the way that 3–4
larger scales form a straight row with 2–3 small scales
above (Fig. 5D). The ones from western Micronesia (Truk
and Palau) (Fig. 5A–B) have the highest number of vertical
scale rows on the cheeks (mostly 8 vs 6–7). Those from
Pohnpei and Kapingamarangi Atoll (Fig. 5E) appear to
have slightly larger eyes than those from other locations.
All these differences, however, are very subtle and are not
supported by other characters so as to warrant a separation
into different species.
Three specimens from Taiwan (Fig. 6A–E) are
tentatively placed in the species (A. aff. crassiceps) due to
some folded, cirri-like dermal laps on the chin, a generally
lower number of scales above the opercular spine (3–6), a
rather short anterior thorn of the inner pseudoclasper and its
remote location from the other specimens of the species.
Comparison. Alionematichthys crassiceps is readily
recognised by its big head and the small eyes, both characters
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 5 (part). Alionematichthys crassiceps sp. nov., A, lateral view of head, holotype; B, ventral view of head, holotype; C, lateral view
of head, holotype; D, lateral view of head, USNM 394976, male, 49 mm SL; E, lateral view of head, USNM 223247, male, 55 mm SL; F,
inclined lateral view of male copulatory organ, holotype; G, ventral view of male copulatory organ, USNM 374196, 58 mm SL; H, view
of left pseudoclaspers from inside, holotype; I, view of left pseudoclaspers from inside, USNM 394976, male, 49 mm SL; J, view of left
pseudoclaspers from inside, USNM 394976, 57 mm SL; K, view of left pseudoclaspers from inside, CAS 222543, 50 mm SL; L, view of left
pseudoclaspers from inside, USNM 223247, 55 mm SL.
97
W. Schwarzhans and P. R. Møller
Fig. 5 (continued). M, median view of right otolith, CAS 222543, male, 50 mm SL; N, median view of right otolith, USNM 352743, female, 67
mm; O, median view of right otolith, USNM 374196, male, 58 mm SL; P, median view of right otolith, USNM 394976, female, 56 mm SL.
that connects it with the West Atlantic A. minyomma, from
which it is distinguished by the presence of scales above
the opercular spine (vs absent) and the compact otolith
with an otolith length to otolith height ratio of 1.9–2.0 (vs
2.2–2.3). The only co-occurring Dinematichthyini with big
heads and small eyes are those of the genus Diancistrus
Ogilby of the Diancistrus erythraeus species group which,
however, shows a completely different pseudoclasper
pattern (see Schwarzhans et al. 2005), no scales above the
opercular spine and an otolith with an undivided sulcus
Fig. 6. Alionematichthys aff. crassiceps, USNM 366687, A, lateral view of head; B, ventral view of head; C, inclined lateral view of male
copulatory organ; D, view of left pseudoclaspers from inside; E, median view of right otolith.
98
Dinematichthyine ishes of the Indo-west Paciic, Part IV
the Kiriwina (Trobriand) Islands of New Guinea, Vanuatu,
Fiji to Tonga (Fig. 1). There are a few tentative records
from Taiwan.
Biology. A 56 mm SL female (USNM 394976)
contained 95 embryos, length 4.1 mm TL, and ca. 50 orange
eggs, diameter 0.6 mm.
Etymology. Named after the big head characteristic
for the species; crassus (Latin = thick) and adjective
ending derived from caput (Latin = head). It is a noun in
apposition.
and fused colliculi. From other co-occurring species of
Alionematichthys, it is distinguished by the following
additional characters: from A. piger, by the small outer
pseudoclasper (vs long), the lack of cirri on the snout (vs
present) and the presence of scales above the opercular
spine (vs absent); from A. plicatosurculus sp. nov., by the
different pseudclasper pattern, the presence of an upper
preopercular pore (vs absent), the presence of scales above
the opercular spine (vs absent); from A. riukiuensis, through
the anteriorly free inner pseudoclasper (vs connected), the
compressed otolith (vs elongate) and the lack of cirri on the
snout (vs present); and from A. winterbottomi sp. nov., in
the shape of the inner pseudoclasper and the lack of cirri
on the snout (vs present).
Distribution. Alionematichthys crassiceps occurs along
oceanic islands of the West Paciic from Micronesia (Palau,
Chuuk, Pohnpei, Kapingamarangi Atoll), Hermit Island and
Alionematichthys minyomma (Sedor and Cohen, 1987)
(Figs 7, 8; Tables 1, 4)
Dinematichthys minyomma Sedor and Cohen, 1987:
6; Nielsen et al. 1999: 130; Møller et al. 2004: 149 (as
D. minyomma).
Species 2. — Sedor 1985.
Table 4. Meristic and morphometric characters of Alionematichthys
minyomma (Sedor and Cohen, 1987)
Table 3. Meristic and morphometric characters of Alionematichthys
crassiceps sp. nov.
Holotype
BPBM
9310
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
63
73
16
59
22
11
30
41
16
1
6
20
13
SL
28.1
17.6
19.9
7.2
14.7
1.3
1.0
7.4
4.7
21.0
49.1
33.1
20.6
15.5
6.6
21.8
25.7
Holotype +
42 paratypes
n
Mean (range)
46.6 (12-67)
43
76.0 (72-81)
16
60.7 (57-64)
21.7 (20-23)
11.0 (10-11)
30.2 (29-31)
41.2 (40-42)
17.4 (16-19)
1.9 (1-2)
5.9 (5-7)
20.6 (18-23)
12.7 (12-13)
38
10
38
9
39
39
39
9
8
39
39
39
30.0 (28.1-34.3)
15.6 (13.7-18.1)
18.6 (17.3-20.8)
6.7 (5.7-8.0)
14.5 (13.6-16.9)
1.9 (1.3-2.5)
1.2 (0.9-1.8)
7.8 (6.6-8.9)
4.9 (4.6-5.7)
22.3 (21.0-26.3)
49.1 (44.0-57.9)
33.1 (31.6-36.6)
19.4 (17.9-23.1)
15.5 (12.6-18.8)
6.7 (6.0-8.2)
26.3 (21.8-29.8)
27.7 (24.3-32.1)
9
9
9
9
9
9
9
9
8
9
9
9
9
9
9
7
9
Holotype
USNM
280122
99
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
66
75
16
58
23
10
29
39
19
2
6
22
13
SL
29.0
17.0
23.5
15.5
1.5
8.5
6.0
22.0
49.5
32.5
13.5
28.0
32.0
Holotype +
10 paratypes
and
28 non-types
Mean (range)
53.3 (26-78)
n
39
74.7 (71-79)
16
57.8 (55-61)
21.7 (21-23)
10.8 (10-11)
29.1 (29-30)
39.8 (39-40)
17.5 (15-19)
2
5.5 (5-6)
22.4 (21-25)
13.1 (12-14)
19
19
19
19
19
19
19
19
18
19
19
19
28.0 (26.6-29.8)
15.5 (13.8-17.5)
20.3 (18.0-23.7)
7.0 (6.3-7.9)
14.8 (13.8-15.9)
2.0 (1.6-2.3)
1.2 (0.9-1.6)
8.1 (7.0-9.1)
5.0 (4.8-5.4)
20.3 (19.3-21.5)
49.9 (45.4-56.0)
32.1 (30.7-34.0)
20.9 (18.9-23.4)
14.8 (13.3-16.8)
7.1 (6.6-8.1)
25.6 (20.7-28.5)
29.1 (25.9-32.1)
19
18
18
15
19
19
16
19
18
18
19
19
19
18
15
19
18
W. Schwarzhans and P. R. Møller
Fig. 7. Sample sites of Alionematichthys minyomma (Sedor and
Cohen, 1987). One symbol may represent several samples.
Material examined. (40 specimens, 26–80 mm SL):
see Møller et al., 2004.
Diagnosis. Vertebrae 10–11+29–30=39–40, dorsal in
rays 71–79, anal in rays 55–61; eyes small (1.5–2.3% SL);
snout rounded, without cirri; scales only on cheeks; upper
preopercular pore present; outer pseudoclasper broadbased, simple lap-shaped, about twice as large as inner
pseudoclasper; inner pseudoclasper very short, its margin
almost straight from short anterior pointed lobe to posterior
lobe; isthmus between pseudoclaspers very wide; otolith
length to height 2.2–2.3, with regular dorsal rim, otolith
length to sulcus length 1.6, ostium length to cauda length
about 3.5–4.0.
Remarks. For a detailed description and additional
illustrations see Møller et al. (2004) (as Dinematichthys
minyomma).
Comparison. Alionematichthys minyomma belongs to
the small species group of the genus characterised by their
small eyes and big head, further characterised by the lack
of cirri and the presence of an upper preopercular pore. It
is distinguished from the only other species of the group
(A. crassiceps) by the lack of scales above the opercular
spine, differences in the shape of the inner pseudoclaspers
and the slender otoliths (otolih length to otolith height
2.2–2.3 vs 1.9–2.0).
Distribution. Alionematihthys minyomma represents
the only species of the genus in the western tropical Atlantic
(off Honduras, Atlantic Panama and Columbia, Haiti,
Puerto Rico and northern Antilles) (Fig. 7) i.e. outside of
the Indo-west Paciic.
Alionematichthys phuketensis sp. nov.
(Figs 1, 10, 11, Tables 1, 5)
Material examined. (9 specimens, 25–41 mm SL).
HOLOTYPE – ZMUC P771659, male 40 mm SL, Rawai
beach, Phuket, Thailand, 07°47’0”N, 98°19’0”E, J. Nielsen,
24 Nov. 1972. PARATYPES – ZMUC P771660-67, 1 male,
36 mm SL, 6 females, 28–41 mm SL, 1 juvenile, 25 mm
SL, same data as holotype.
Diagnosis. Vertebrae 12+30–31=42–43, dorsal in rays
81–89, anal in rays 60–65; eyes moderately large (2.3–
2.9% SL); body slender, snout pointed, without cirri; scales
only on cheeks; no upper preopercular pore (occasionally
single pore on one side); outer pseudoclasper broad-based,
small; inner pseudoclasper small, anteriorly inclined lap;
otolith length to height 2.1–2.3, with distinct postdorsal
Fig. 8. Alionematichthys minyomma (Sedor and Cohen, 1987), A, lateral view of head, USNM 280123, male, 66 mm SL; B, ventral view of
head, USNM 280123, female, 61 mm SL; C, view of left pseudoclaspers from inside, USNM 280123, male, 71 mm SL; D, inclined lateral
view of male copulatory organ of same specimen; E, median view of right otolith, USNM 280123, male 71 mm SL.
100
Dinematichthyine ishes of the Indo-west Paciic, Part IV
lower two elongate rakers interrupted by one plate-like
raker. In rest of paratypes, three elongate rakers in single
row. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 11A, B). Supraorbital pores
2. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about 1/3 the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): irst anterior
pore large, tubular, without cirri. Preopercular pores: 3
lower, irst and second with joined opening, covered by
dermal lap in lateral view; third non-tubular; no upper
preopercular pore (rarely single pore on one side). [See
description of Alionematichthys ceylonensis for position
of pores.]
Dentition (of holotype). Premaxilla with 5 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.
Vomer horseshoe-shaped, with 1 outer row of small teeth
and 1 inner row of large teeth up to 1/5 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of large teeth up to 1/5 diameter of pupil. Dentary with 5
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/3 diameter of pupil.
Otolith (Fig. 11E–G). Elongate in shape, length to height
2.1–2.3 (36–41 mm SL) and moderately thin (otolith height
to otolith thickness about 1.6). Anterior tip moderately
pointed, posterior tip expanded and irregularly ornamented.
Dorsal rim shallow, with distinct postdorsal projection and
furrow thereafter. Inner face moderately convex, outer face
almost lat, both smooth. Otolith length to sulcus length
1.6–1.7. Sulcus slightly supramedially positioned, slightly
inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.0–3.3. Ventral furrow very close
to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined
and 5–8 depressed. Parapophyses present from vertebrae
6 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygiophore not or only slightly longer than subsequent
pterygiophore.
Male copulatory organ (Fig. 11C, D). Two pairs of
small pseudoclaspers, which may not be fully mature in
all specimens investigated. Outer pseudoclasper a simple
flap with broad base; inner pseudoclasper a forward
inclined small lap without signiicant indentation. Isthmus
moderately narrow; penis curved, longer than outer
pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly light brown.
Comparison. Alionematichthys phuketensis belongs
to the species group without an upper preopercular pore
together with A. ceylonensis, A. plicatosurculus sp. nov.
and A. shinoharai sp. nov. It differs from all of them in
the combination of a slender body shape with a pointed
snout and the peculiar postdorsal projection of the otolith
rim. From A. plicatosurculus sp. nov., it differs further
in the simple inner pseudoclasper morphology and from
Table 5. Meristic and morphometric characters of Alionematichthys
phuketensis sp. nov.
Holotype +
n
Holotype 8 paratypes
ZMUC
P771659
Mean (range)
Standard length in mm
40
33.0 (25-41)
9
Meristic characters
Dorsal in rays
83
83.3 (81-89)
9
Caudal in rays
15
15.2 (14-16)
6
Anal in rays
62
62.9 (60-65)
9
Pectoral in rays
20
19.9 (19-21)
7
Precaudal vertebrae
12
12
9
Caudal vertebrae
31
30.8 (30-31)
9
Total vertebrae
43
42.8 (42-43)
9
Rakers on anterior gill arch
18
16.0 (15-18)
6
Pseudobranchial ilaments
2
2
4
D/V
6
6
9
D/A
24
23.1 (21-25)
9
V/A
14
13.3 (13-14)
9
Morphometric characters in % of SL
Head length
26.8
26.7 (25.1-27.5) 9
Head width
12.5
11.7 (11.0-12.5) 9
Head height
15.0
15.1 (14.0-16.1) 9
Snout length
6.2
6.2 (5.6-6.5)
9
Upper jaw length
13.1
12.9 (12.5-13.7) 9
Diameter of pigmented eye
2.9
2.6 (2.3-2.9)
9
Diameter of pupil
1.8
1.6 (1.4-1.8)
9
Interorbital width
7.3
7.3 (6.9-7.9)
9
Posterior maxilla height
4.4
3.7 (3.3-4.4)
9
Postorbital length
18.4
18.6 (17.9-19.4) 8
Preanal length
44.9
43.4 (41.0-45.6) 9
Predorsal length
31.0
30.7 (29.2-32.2) 9
Body depth at origin of anal in 16.9
15.3 (13.7-16.9) 9
Pectoral in length
12.4
13.2 (11.9-15.0) 9
Pectoral in base height
5.2
5.8 (5.2-6.3)
9
Ventral in length
19.8
23.5 (19.8-24.8) 9
Base ventral in - anal in origin 25.2
23.9 (22.4-25.2) 9
projection, otolith length to sulcus length 1.6–1.7, ostium
length to cauda length 3.0–3.3.
Description (Figs 10, 11).The principal meristic and
morphometric characters are shown in Table 5. Mature
at about 40 mm SL (largest male at 40 mm SL probably
subadult). Body slender, with pointed head proile. No cirri
on snout. Eye size 2.3–2.9 (2.9)% SL. Head with scale patch
on cheek containing 7 vertical rows of scales on upper part
and 4 vertical rows on lower part. Horizontal diameter
of scales on body about 1.5% SL, in about 30 horizontal
rows. Maxillary ending far behind eye, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded. Anterior nostril positioned high, 1/3 distance
from upper lip to anterior margin of eye. Posterior nostril
small, about 1/8 the size of eye. Opercular spine with free
tip, thin and sharply pointed. Anterior gill arch with 15–18
(18) rakers, 3 elongate. In holotype and two paratypes,
101
W. Schwarzhans and P. R. Møller
Fig. 9. Sample sites of Alionematichthys suluensis sp. nov. and A. piger (Alcock, 1890). One symbol may represent several samples.
A. shinoharai sp. nov., in the lack of scales above the
opercular spine (vs two), the longer sulcus (otolith length
to sulcus length 1.6–1.7 vs 1.9–2.0) and the notch at the
ventral sulcus margin at the joint of ostium and cauda (vs
no notch). Alionematichthys phuketensis co-occurs with
A. piger, A. riukiuensis and Dinematichthys iluocoeteoides.
From the latter two it differs readily in the absence of scales
above the opercular spine (vs present or entire opercle with
scales in D. iluocoeteoides). From A. piger it differs in the
lack of cirri on the snout, the lack of an upper preopercular
pore and the peculiar dorsal projection of the otolith rim.
Distribution. Alionematichthys phuketensis so far
is only known from the type locality, Ravai Beach near
Phuket, Thailand (Fig. 1).
Etymology. Named after the type locality, the town of
Phuket, Thailand. It is an adjective.
Alionematichthys piger (Alcock, 1890)
(Figs 9, 12, 13, Tables 1, 6)
Dinematichthys piger Alcock, 1890: 432 (Great Coco
Island, Andaman Islands); Alcock, 1905: plate 37, ig. 3;
?Brotulina piger (Alcock, 1890). — Nielsen et al.
1999: 126.
Material examined. 2288 specimens, 16–84 mm SL.
AMS I.17094-070, 3 males, 5 females, 11 juveniles,
08°30’S, 151°05’E, Kiriwina Islands (Trobriand), Papua
New Guinea; AMS I.17102-004, 2 specimens, 08°30’S
151°05’E, Kiriwina Islands (Trobriand), Papua New
Guinea; AMS I.17424-012, 1 female, 31°32’S, 159°04’E,
Lord Howe Island; AMS I.18052-040, 2 females, 28–30
mm SL, 01°44’N, 172°59’E, Abaiang Atoll, Gilbert Islands,
Kiribati; AMS I.20629-001, 3 males, 10 females, 12°N,
102
165°E, Bikini Atoll, Marshall Islands; AMS I.27134-040,
1 female, 29°27’2’’S, 159°06’8’’E, Lord Howe Island, 6–9
m; AMS I.27138-047, 1 male, 46 mm SL, 2 females, 34–67
mm SL, 29°27’8’’S, 159°05’E, Middleton Reef, Lord Howe
Rise, Tasman Sea, 4–9 m; AMS I.27148-029, 1 female,
29°57’S, 159°01’2’’E, Elizabeth Reef, Tasman Sea, 0–10
m; AMS I.27891-040, 1 specimen, 29°56’S, 159°01’E,
Elizabeth Reef, Tasman Sea, 0–5 m; AMS I.28950-045, 2
specimens, 17°29’1’’S, 149°51’3’’W, Moorea, French
Polynesia, 8–12 m; AMS I.33740-026, 3 males, 49–59 mm
SL and 6 females, 10°09.58’’S, 144°34.94’’E, Ashmore
Reef, Coral Sea; AMS I.34510-020, 42 specimens,
15°44.8’S, 144°38.9’W, Taiaro Atoll, Tuamotu Islands, 0–2
m; AMS I.37315-032, 2 males, 65–75 mm SL, 4 females,
40–54 mm SL, 18°44’45’’S, 169°12’68’’E, Erromango,
Vanuatu, 0–6 m; AMS I.39387-001, 12 specimens, Pelagai
Island, Guam, 1–2 m; BPBM 7553, 1 male, 4 females,
Fanning Island, Line Islands; BPBM 8009, 2 males, 5
females, Marshall Islands; BPBM 8242, 1 male, 2 females,
Enewetak Atoll, Marshall Islands; BPBM 8651, 1 male, 5
females, Tahiti; BPBM 8658, 1 female, Tahiti; BPBM 9311,
1 female, Chuuk, Micronesia; BPBM 10240, 1 specimen,
Rangiroa, Tuamotu Islands; BPBM 10307, 2 specimens,
Rangiroa, Tuamotu Islands; BPBM 10658, 1 male and 9
females, Gilbert Island, Kiribati; BPBM 10830, 1 specimen,
Aitutaki, Cook Islands; BPBM 11588, 1 female, Tahiti;
BPBM 12254, 1 male, 2 juveniles, Ducie, Pitcairn; BPBM
13574, 1 male and 1 female, Mangareva, Gambier Islands,
French Polynesia; BPBM 13915, 2 females, Rarotonga,
Cook Islands; BPBM 13929, 1 male, Rarotonga, Cook
Islands; BPBM 14161, 1 female, Palmyra Atoll, Line
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Islands; BPBM 14788, 1 male, 3 females, Lord Howe
Island; BPBM 15169, 1 male, 1 female, Wake Island, USA;
BPBM 15199, 1 male, 58 mm SL, 6 females, 26–60 mm
SL, Kiritimati, Line Islands; BPBM 15323, 1 male, 4
females, Betio, Tarawa Atoll, Kiribati; BPBM 15616, 1
female, Guadalcanal, Solomon Islands; BPBM 16495, 5
males, 7 females, Pitcairn, Polynesia; BPBM 16534, 2
males, 1 female, Oeno, Pitcairn; BPBM 16607, 1 female,
Oeno, Pitcairn; BPBM 16912, 2 females, Pitcairn; BPBM
16970, 2 males, 31 and 59 mm SL, 5 females, 28–55 mm
SL, Pitcairn; BPBM 16993, 1 female, Pitcairn; BPBM
17042, 3 females, Pitcairn; BPBM 17223, 1 male, 1 female,
Mangareva, Gambier Islands, French Polynesia; BPBM
17690, 1 female, Ryukyu Islands, Japan; BPBM 22357, 2
females, Marshall Islands; BPBM 22834, 2 males, 1 female,
Similan Island, Thailand; BPBM 23262, 1 male and 3
females, Taiwan; BPBM 23338, 2 males and 2 females,
Taiwan; BPBM 33514, 1 male, Chesterield Bank, Coral
Sea; BPBM 37451, 1 female, Kiritimati; BPBM 38268, 1
female, Tonga; BPBM 38773, 2 males, Tahiti; CAS 227288,
43 specimens, Ifalik Atoll, Micronesia; CAS 57943, 3
specimens, Raroia, Tuamotu Islands; CAS 65671, 1 female,
Madang, Papua New Guinea; CAS 81437, 29 specimens,
Raroia, Tuamotu Islands; CAS 227280, 1 juv, 21 mm SL,
24 males 40–63 mm SL, 27 females 42–71 mm SL,
Kapingamarangi Atoll, Micronesia; CAS 81446, 16
specimens, Ifalik Atoll, Micronesia; CAS 81451, 4
specimens, Kapingamarangi Atoll, Micronesia; CAS 81468,
52 specimens, Raroia, Tuamotu Islands; CAS 81483, 96
specimens, Kapingamarangi Atoll, Micronesia; CAS
222525, 29 specimens, Fiji;CAS 222527, 2 specimens, Fiji;
CAS 222531, 1 female, Fiji; CAS 222537, 1 female, Fiji;
CAS 227283, 2 specimens, Fiji; CAS 222549, 2 specimens,
Fiji; CAS 222553, 2 specimens, Fiji; CAS 222554, 4
specimens, Yadua Island, Fiji; CAS 222556, 5 specimens,
Fiji; CAS 222569, 1 specimen, Bua Bay, Fiji; CAS 227286,
2 specimens, Vanua Levu, Fiji; CAS 222572, 1 specimen,
Vanua Levu, Fiji; CAS 222582, 5 specimens, Balvu
Harbour, Fiji; CAS 227293, 61 specimens, Fiji; CAS
222601, 45 specimens, Budd Reef, Fiji; CAS 222604, 9
specimens, Fiji; CAS 222606, 20 specimens, Suva, Fiji;
CAS 222607, 40 specimens, Suva, Fiji; CAS 227301, 1
specimen, Suva, Fiji; CAS 222625, 8 specimens, Suva, Fiji;
CAS 222626, 13 specimens, Suva, Fiji; CAS 222633, 19
specimens, Suva, Fiji; CAS 222640, 2 specimens, Suva,
Fiji; KAUM-I. 2042, 1 female, 77 mm SL, off Okinoerabu
Island, Kagoshima, Japan, July 1962; KAUM-I. 10652, 1
female, 76 mm SL, off Okinoerabu Island, Kagoshima,
Japan; KAUM-I. 10655, 1 female, 56 mm SL, off
Okinoerabu Island, Kagoshima, Japan; KAUM-I. 10658,
1 male, 49 mm SL, off Okinoerabu Island, Kagoshima,
Japan; KAUM-I. 10663, 1 male, 33 mm SL, off Okinoerabu
Island, Kagoshima, Japan; KAUM-I. 11482-83, 1 male, 58
mm SL and 1 female, 50 mm SL, 30°27’23’’N, 130°29’59’’E,
Isso, Yakushima Island, Kagoshima, Japan; MCZ 158557,
1 specimen, 04°27’S, 171°13’W, Manra Atoll, Kiribati;
MCZ 162572, 4 specimens, 04°27’S, 171°14’W, Manra
Atoll, Kiribati; MCZ 162573, 5 specimens, 04°27’S
171°15’W, Manra Atoll, Kiribati; MNHN 1976-0218, 3
females, 54–56 mm SL, Marshall Islands; MNHN 19800563, 2 specimens, New Caledonia; MNHN 1980-0696, 1
female, New Caledonia; MNHN 1980-0961, 1 male, 65
mm SL, 1 female, New Caledonia; NMNZ P.035811, 1
male, 44 mm SL, 1 female, 35 mm SL, Niue, Polynesia;
NSMT-P 55637, 5 specimens, Hainan Island, China;
NSMT-P 55823, 2 males, 3 females and 1 juvenile, Hainan
Island, China; ROM 82976, 10 specimens, 18°45’52’’S,
178°31’13’’E, Fiji; ROM 47600, 5 specimens, 18°46’30’’S,
178°30’28’’E, Fiji; ROM 47601, 10 specimens, 18°46’13’’S,
178°28’05’’E, Fiji; ROM 61176, 7 specimens, 17°31’00’’S,
149°55’30’’W, Society Islands; ROM 61180, males, 30 and
47 mm SL, female 35 mm SL, 17°36’31’’S, 149°48’42’’W,
Society Islands; ROM 82979, 1 female, 22°21’00’’S,
166°21’00’’E, New Caledonia; ROM 82978, 1 female, 68
mm SL, 20°48’51’’N, 107°05’13’’E, Vietnam; RUSI 40747,
7 specimens, Kiritimati, Kiribati; SMNS 17837, 1 specimen,
41 mm SL, Aitutaki Island, Cook Islands, 18°50’34’’S,
159°47’28’’W, 0.2–3.5 m depth. SMNS 19762, 11
specimens, 21°38’57’’S, 166°18’31’’E, New Caledonia,
0–8.5 m; SMNS 25171, 4 specimens, 21°35’45’’S,
167°50’06’’E, Loyalty Islands, New Caledonia, 2–3.8 m;
SMNS 21646, 13 specimens, 20°53’35’’S, 167°08’02’’E,
Loyalty Islands, New Caledonia, 0–2.5 m; SMNS 22732,
1 male, 1 female, 20°36’30’’S, 164°51’50’’E, 0–6 m; SMNS
22782, 1 male, 20°31’49’’S, 164°47’01’’E, New Caledonia,
0.2–3.5 m; SMNS 22918, 5 specimens, 20°52’10’’S,
167°08’02’’E, Loyalty Islands, New Caledonia, 0–3 m;
SMNS 22967, 5 specimens, 20°51’45’’S, 167°08’10’’E,
Loyalty Islands, New Caledonia, 0–3.5 m; SMNS 23747,
1 male, 1 female, 20°46’54’’S, 167°08’19’’E, Loyalty
Islands, New Caledonia, 0–3 m; SMNS 23923, 8 specimens,
20°47’11’’S, 167°07’20’’E, Loyalty Islands, New
Caledonia, 0–3 m; SMNS 24849, 18 specimens, 21°56’13’’N,
120°47’53’’E, Taiwan; SMNS 24864, 2 specimens,
21°56’13’’N, 120°47’53’’E, Taiwan; SMNS 24866, 1
specimen, 21°56’13’’N, 120°47’53’’E, Taiwan; UF 42494,
2 males, 1 female, Enewetak Atoll, Marshall Islands;
USNM 376214, 2 males, 04°01’S, 101°01’E, Sumatra,
Mentawai; USNM 384607, 24 specimens, 21°N, 120°E,
Taiwan; USNM 224327, 34 specimens, 05°51’N, 158°20’E,
Pohnpei, Micronesia, 0–2 m; USNM 297347, 59 specimens,
20°24’45’’N, 121°55’02’’E, Batanes, Philippines, 0–6 m;
USNM 384608, 10 specimens, 18°51’N 121°22’E, Fuga
Island, Philippines; USNM 384604, 17 specimens,
19°51’36’’S, 174°25’06’’W, Tonga, 0–2 m; USNM 338463,
18 specimens, 18°40’55’’S, 174°06’09’’W, Vava’u Group,
Luamoko Island, Tonga, 0–3 m; USNM 384605, 26
specimens, 18°44’31’’S, 174°06’36’’W, Vava’u Group,
Tonga, 0–11 m; USNM 384593, 6 specimens, 20°29’12’’S,
166°19’18’’E, Loyalty Islands, New Caledonia, 3–10 m;
USNM 384601, 38 specimens, 19°31’33’’S, 169°29’50’’E,
Tanna, Vanuatu, 1–5 m; USNM 394979, 3 females,
103
W. Schwarzhans and P. R. Møller
Fig. 10. Alionematichthys phuketensis sp. nov., ZMUC P771659, holotype, male, 40 mm SL.
Fig. 11. Alionematichthys phuketensis sp. nov., A, lateral view of head, ZMUC ZMUC P771661, male, 37 mm SL; B, ventral view of head,
ZMUC P771661, male, 36 mm SL; C, view of left pseudoclaspers from inside, holotype; D, inclined lateral view of male copulatory organ,
holotype; E, median view of right otolith, holotype; F, ventral view of right otolith, holotype; G, median view of right otolith, ZMUC P771660,
female, 41 mm SL.
13°52’25’’S, 167°33’20’’E, Banks Islands, Vanuatu, 9–13
m; USNM 366488, 3 females, 28–43 mm SL, Phuket,
Thailand; USNM 366507, 23 males, 40–84 mm SL, 35
females, 39–71 mm SL, Kiriwina Islands (Trobriand),
Papua New Guinea; USNM 366510, 3 males and 19
females, 39–64 mm SL, 05°33’N, 95°09’E, Pulau Boenta,
Sumatra; USNM 366569, 2 females, 62–84 mm SL,
23°25’S, 151°55’E, Heron Island, Queensland, Australia;
USNM 394980, 1 female, 10°35’05’’N, 122°08’30’’E;
Iloilo, Philippines, 0–7 m; USNM 374192, 1 female, 44
mm SL, 09°06’N, 122°55’E, Negros Island, Philippines;
USNM 376177, 1 male, 45 mm SL, Taiwan; USNM
104
376178, 1 female, 18°56’34’’S, 168°59’34’’E, Erromango,
Vanuatu, 0–6 m; USNM 376180, 3 males and 7 females,
46–62 mm SL, 08°29’N, 97°39’E, Thailand; USNM
376187, 1 male, 52 mm SL, Sabah; WAM P.29081-016, 3
females, 15°31’S, 123°09’E, Adele Island, Western
Australia, 0.1–2.0 m; WAM P.30633-017, 1 female,
05°09’S, 145°50’E; WAM P.31646-002, 1 male, 13°S
121°E; YCM-P 30026, 1 female, Ishigaki Island, South
Ryukyu Islands, Japan; YCM-P 34215, 1 female, Kakeroma
Island, Amami Island, North Ryukyu Islands, Japan.
Additional material. USNM 115392, 1 male, 43 mm
SL, Phoenix Islands, Kiribati; USNM 115395, 1 male, 52
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 12. Alionematichthys piger (Alcock, 1890), AMS I.ex. 37315-032, male, 65 mm SL.
mm SL, Phoenix Island, Kiribati; USNM 142010, 4 males
and 4 females, 28–42 mm SL, Rongerik Atoll, Marshall
Islands; USNM 374219, 5 males and 5 females, 25–45 mm
SL, Rongelap Atoll, Marshall Islands; USNM 142012, 1
male, 45 mm SL, Kwajalein Atoll, Marshall Islands; USNM
142013, 11 specimens, 40–62 mm SL, Enewetak Atoll,
Marshall Islands; USNM 142014, 9 specimens, 16–40
mm SL, Bikini Atoll, Marshall Islands; USNM 142017,
18 females, 27–58 mm SL, Bikini Atoll, Marshall Islands;
USNM 374220, 46 specimens, 32–57 mm SL, Bikini Atoll,
Marshall Islands; USNM 142020, 92 specimens, 28–56
mm SL, Bikini Atoll, Marshall Islands; USNM 374215, 2
males and 1 female, 45–58 mm SL, Arno Atoll, Marshall
Islands; USNM 167355, 3 males and 7 females, 40–55
mm SL, Onotoa, Gilbert Islands, Kiribati; USNM 210154,
1 male, 32 mm SL, Seram, Tandjungliang, Indonesia;
USNM 376161, 86 specimens, 30–84 mm SL, Taiwan;
USNM 263663, 5 males and females, 48–60 mm SL,
Tahiti, French Polynesia; USNM 263674, 19 males and
42 females, 25–56 mm SL, 16°00’S, 175°53’W, Tonga;
USNM 263690, 30 specimens, 29–57 mm SL, 10°55’N,
121°02’E, Palawan, Philippines; USNM 263711, 1 male,
86 mm SL, 21°55’N, 120°49’E, Taiwan; USNM 300090,
1 male and 2 females, 40–45 mm SL, 20°25’N, 121°57’E,
Batanes Islands, Philippines; USNM 300093, 7 males and
8 females, 38–75 mm SL, 20°20’N, 121°49’E, Batanes
Islands, Philippines; USNM 300098, 1 female, 47 mm SL,
20°28’N, 121°58’E, Batanes Islands, Philippines; USNM
300102, 1 male and 2 females, 21°07’N, 121°56’E, Batanes
Islands, Philippines; USNM 319899, 7 males and females,
45–73 mm SL, Ouvea, Loyalty Islands; USNM 329746, 17
males and 12 females, 31–57 mm SL, 21°18’S, 174°26’W,
Tonga; USNM 333251, 3 males and females, 29–41 mm SL,
Tongatapu, Tonga; USNM 336507, 2 females, 55–60 mm
SL, 21°25’S, 174°56’W, Tonga; USNM 374214, 6 males
and 3 females, 35–55 mm SL, 19°16’S, 174°22’W, Tonga;
USNM 338465, 35 specimens, 32–61 mm SL, Vavau,
Tonga; USNM 338966, 7 males and 2 females, 30–45 mm
SL, 18°42’S, 174°02’W, Tonga; USNM 338967, 9 males
and 14 females, 30–50 mm SL, 18°38’S, 174°04’W, Tonga;
USNM 352073, 1 female, 47 mm SL, 16°35’S, 168°06’E,
Lamen Island, Vanuatu; USNM 352742, 123 specimens,
38–80 mm SL, 18°44’S, 169°12’E, Erromango, Vanuatu;
USNM 358502, 18 males and females, 24–71 mm SL,
Tanna, Vanuatu; USNM 361266, 7 females, 35–80 mm
SL, 13°44’S, 167°24’E, Banks Islands, Vanuatu; USNM
376169, 1 female, 48 mm SL, 15°00’S, 168°03’E, Maewo,
Vanuatu; USNM 366200, 5 males and females, 17–65
mm SL, Tonga; USNM 366222, 5 males and females,
38–61 mm SL, Line Islands; USNM 366223, 1 male, 51
mm SL, Starbuck Atoll, Line Islands; USNM 366225,
1 male, 2 females, Bora Bora, Society Islands; USNM
366489, 2 females, 34–49 mm SL, Taka Atoll, Marshall
Islands; USNM 376218, 1 female, 44 mm SL, 04°14’S,
152°10’E, Bismarck Archipelago, Papua New Guinea;
USNM 366511, 2 females, 36 mm SL, Amami, Kikai
Island, Ryukyu Islands; USNM 376213, 2 females, 38–63
mm SL, Kiriwina Islands (Trobriand); USNM 366556, 1
female, 38 mm SL, Taiwan; USNM 366560, 1 female, 49
mm SL, 25°12’N, 121°41’E, Taiwan; USNM 366561, 12
specimens, 43–78 mm SL, 25°12’N, 121°41’E, Taiwan;
USNM 366574, 2 males and 8 females, 44–51 mm SL,
Kwajalein Atoll, Marshall Islands; USNM 366575, 1 male,
35 mm SL, Phoenix Atoll, McKean Island; USNM 374221,
1 female, 52 mm SL, Kwajalein Atoll, Marshall Islands;
USNM 366719, 1 male and 2 females, 51–67 mm SL, Jarvis
Island, Line Islands; USNM 366725, 2 males and 1 female,
40–51 mm SL, Bora Bora, Society Islands; USNM 366843,
1 male and 1 female, 54–55 mm SL, 20°37’S, 178°40’E,
Fiji; USNM 376211, 1 male and 4 females, 30–48 mm SL,
18°57’S, 178°17’E, Fiji; USNM 366847, 1 male, 49 mm
SL, 18°56’S, 178°21’W, Fiji; USNM 366848, 2 males and
4 females, 32–45 mm SL, 17°47’S, 177°13’E, Fiji.
Diagnosis. Vertebrae 11–12+29–33=40–45, dorsal in
rays 72–88, anal in rays 55–70; eyes moderately large (1.7–
2.8% SL); body slender, snout rounded, with many cirri;
scales only on cheeks (rarely 1–2 scales above opercular
spine); upper preopercular pore present; outer pseudoclasper
broad-based, large, often extending beyond hood in resting
position; inner pseudoclasper small, anterior and posterior
lobes of equal size; otolith length to height 1.9–2.1, with
gently curved dorsal rim, otolith length to sulcus length
1.5–1.6, ostium length to cauda length 3.5–4.5.
105
W. Schwarzhans and P. R. Møller
Fig. 13 (part). Alionematichthys piger (Alcock, 1890), A, lateral view of head, ROM 82976, male, 51 mm SL; B, lateral view of head, USNM
384707, male, 61 mm SL; C, lateral view of head, USNM 366507, male, 54 mm SL; D, ventral view of head, USNM 384707, male, 61 mm
SL; E, lateral view of head, CAS 227288, female, 51 mm SL; F, inclined lateral view of male copulatory organ, USNM 376180, male, 57 mm
SL; G, view of left pseudoclaspers from inside, USNM 376180, male, 57 mm SL; H, view of left pseudoclaspers from outside, USNM 376180,
male, 44 mm SL; I, ventral view of male copulatory organ, USNM 366507, male, 54 mm SL; J, view of left pseudoclaspers from inside,
SMNS 17837, 41 mm SL; K, view of left pseudoclaspers from inside, ROM 82976, 43 mm SL; L, view of left pseudoclaspers from inside,
MNHN 1980-0961, 65 mm SL; M, view of left pseudoclaspers from inside, CAS 227288, male, 59 mm SL; N, view of left pseudoclaspers
from inside, BPBM 8009, male, 43 mm SL; O, ventral view of female copulatory organ, SMNS 19762, 85 mm SL; P, inclined lateral view
of female copulatory organ, SMNS 19762, 85 mm SL.
106
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 13 (continued). Q, median view of right otolith, USNM 297347, male, 67 mm SL; R, ventral view of right otolith, USNM 297347, male,
67 mm SL; S, median view of right otolith, USNM 384605, male, 52 mm SL; T, median view of right otolith, AMS I.23381-001, isolated
otolith; U, median view of right otolith, MNHN 1976-0218, female, 56 mm SL.
Description (Figs 12, 13). The principal meristic and
morphometric characters are shown in Table 5. Mature at
about 40 mm SL. Body slender, with rounded head proile.
Many cirri on snout. Eye size 1.7–2.8 (2.6)% SL. Head
with scale patch on cheek containing 6–9 vertical rows of
scales on upper part and 3–4 vertical rows on lower part.
Horizontal diameter of scales on body of a 65 mm SL male
(AMS I.37315-032) about 1.8% SL, in about 27 horizontal
rows. Maxillary ending far behind eye, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded. Anterior nostril positioned high, 1/2.5 distance
from upper lip to anterior margin of eye. Posterior nostril
small, about one-ifth to one-sixth the size of eye. Opercular
spine with free tip, thin and sharply pointed. Anterior gill
arch with 13–19 rakers, 3 elongate, sometimes with short
plate-like raker between the two lower elongate rakers.
Pseudobranchial ilaments 2.
Head sensory pores (Fig. 13A–E). Supraorbital pores 2
to 3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about 1/3 the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): irst anterior
pore large, tubular, without cirri. Preopercular pores: 3 lower,
irst and second with joined opening, covered by dermal lap
in lateral view; third nontubular; upper preopercular pore
present. [See description of Alionematichthys ceylonensis
for position of pores.]
Dentition (of a 59 mm SL specimen – USNM 376180).
Premaxilla with 5 outer rows of granular teeth and 2 inner
107
row of larger teeth anteriorly. Anteriormost teeth in inner
row up to 1/2 diameter of pupil. Vomer horseshoe-shaped,
with 2 outer rows of small teeth and 1 inner row of larger
teeth up to 1/3 diameter of pupil. Palatine with 2 outer
rows of small teeth and 1 inner row of larger teeth up to 1/3
diameter of pupil. Dentary with 5 outer rows of granular
teeth and 1 inner row of larger teeth anteriorly, up to about
3/4 diameter of pupil.
Otolith (Fig. 13Q–U). Moderately elongate in shape,
length to height 1.9–2.1 and thin (otolith height to otolith
thickness about 2.3). Anterior tip broadly rounded, posterior
tip expanded and irregularly ornamented. Dorsal rim gently
curved, without distinct angles. Inner face moderately
convex, outer face almost lat, both smooth. Otolith length
to sulcus length 1.5–1.6. Sulcus slightly supramedially
positioned, slightly inclined, with separated colliculi and
marked notch at ventral rim of sulcus at joint of ostium
with cauda. Ostium length to cauda length 3.5–4.5. Ventral
furrow very distinct and close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to
11 (12). Pleural ribs on vertebrae 2 to 10 (11). First anal
in pterygiophore not or slightly longer than subsequent
pterygiophore.
Male copulatory organ (Fig. 13G–N). Two pairs of
pseudoclaspers, outer usually very large and extending
beyond hood in resting position. Outer pseudoclasper a
large, lat lap with broad base; inner pseudoclasper short,
W. Schwarzhans and P. R. Møller
sometimes compressed, with anterior thorn and posterior
lobe being equal in size. Isthmus narrow; penis curved,
about as long as outer pseudoclaspers.
Female copulatory organ (Fig. 13O, P). Few female
specimens of A. piger show soft dermal laps around the
vagina which may reach a quite considerable size, as in the
igured specimens but do not exhibit diagnostic valuable
morphology.
Colour. According to Alcock (1890: 433), the live colour
is uniform dark brown, almost black. The preserved colour
is similar.
Variability. Alionematichthys piger is remarkable for an
unusual wide variation in pseudoclasper morphology that
does not follow a geographic pattern. The majority of the
male specimens show a relatively large outer pseudoclasper
for a member of this genus, extending beyond the hood
in resting position and about twice as long as the inner
pseudoclasper. In extreme forms the inner pseudoclasper
is depressed (Fig. 13J, K). On the other extreme, the inner
pseudoclasper is rather stubby and the outer pseudoclasper
less than half the length of the inner pseudoclasper and not
extruding in resting position (Fig. 13M, N). The very large
variation is also relected in the meristic counts (Tables
1, 5) and variations in the amount of cirri on the snout
(Fig. 13A–E) which may represent geographic variances.
Otherwise, heads (Fig. 13A–E) and otoliths (Fig. 13Q–U)
of ishes with such diverse pseudoclaspers do not document
matching variation.
Remarks. Alcock (1890) described Dinematichthys
piger from Great Coco Island, Andaman Islands. The
unique holotype of D. piger (ZSI F12939) was not available
for review (see also Schwarzhans and Møller 2007: 66).
According to Alcock’s description it is a 61 mm long
specimen with 75 dorsal in rays, 55 anal in rays and 90
scales along the lateral line and his igure shows a ish with a
rather high position of the anterior nostril at 1/3 the distance
from upper lip to aggregate distance to anterior margin of
eye and no scales on the opercle.
The latter two observations agree with a species of
the genus Alionematichthys. There are two species of the
genus Alionematichthys observed along the shores of the
Andaman Sea, one without scales above the opercle, the
other with many scales above the opercle. The latter has
been attributed to a species described from the Ryukyu
Islands, namely Dinematichthys riukiuensis, now placed in
the genus Alionematichthys (see below). Another species
described above from the Thailand coast, A. phuketensis,
across the Andaman Sea from Alcock’s type locality, does
not seem to attain such size and has higher dorsal (81–89)
and anal (60–65) in ray counts. Finally, Alcock’s drawing
is detailed enough to reveal that the holotype must be a
female. With all these facts in mind it has to be considered
that the specimens attributed to A. piger in the following
and the diagnosis of the species and description are not fully
satisfactory until the holotype has been re-studied.
108
Comparison. Alionematichthys piger belongs to the
species group with an upper preopercular pore, large eyes
and no (rarely 1 or 2) scales above the opercular spine
comprising also A. samoaensis sp. nov. and A. suluensis
sp. nov., which both have a much more restricted distribution.
It differs from A. samoaensis sp. nov. by the lack of a knob
at the distal inner end of the outer pseudoclasper and the
simple lap at the anterior margin of the posterior nostril
(vs strongly elevated funnel shaped). From A. suluensis, it
differs in the anterior lobe of the inner pseudoclasper being
equally long as its posterior lobe (vs very long anterior
lobe, twice as long as posterior lobe) and many cirri on the
snout (vs no cirri).
Distribution. Alionematichthys piger is one of the most
common and most widely distributed Dinematichthyini
(Fig. 9). Its distribution ranges from the Andaman Islands
to Sumatra, Thailand, Vietnam, Taiwan and the Ryukyu
Table 6. Meristic and morphometric characters of Alionematichthys
piger (Alcock, 1890)
Holotype +
Holotype 340 non-types
*
Mean (range)
61
50.6 (26-86)
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
75
55
SL
25.1
16.7
2.6
17.8
50.4
31.2
16.6
13.9
5.7
-
n
99
79.7 (72-88)
16.0 (14-18)
61.8 (55-70)
20.6 (19-22)
11.1 (11-12)
31.1 (29-34)
42.1 (40-45)
15.8 (13-19)
2
5.7 (5-7)
22.7 (19-28)
13.0 (12-16)
305
82
305
9
340
337
337
31
2
338
336
337
25.7 (24.0-27.5)
14.0 (11.8-15.0)
15.9 (13.6-16.8)
5.8 (5.1-6.6)
13.5 (12.5-14.4)
2.3 (1.7-2.8)
1.4 (0.9-1.6)
7.1 (6.5-7.9)
4.5 (4.0-5.2)
18.6 (17.6-19.5)
48.0 (44.4-50.9)
30.1 (27.4-31.2)
17.8 (14.8-19.5)
13.5 (11.9-14.7)
6.1 (5.7-6.9)
24.0 (21.2-27.6)
29.7 (25.3-34.5)
10
9
10
9
9
10
9
9
9
10
10
10
10
10
10
9
9
* Data taken from Alcock (1890 and 1905).
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Islands in the north, to Australia and as far south as Lord
Howe Island and eastwards to Micronesia, the Line Islands,
the Tuamotu Archipelago and Pitcairn Group. Together with
Diancistrus katrineae Schwarzhans, Møller and Nielsen, it
represents the species farthest to the east in the West Paciic
at Ducie Atoll. The species is found in holes and crevices
of coralline and volcanic rock.
Alionematichthys plicatosurculus sp. nov.
(Figs 1, 14, 15, Tables 1, 7)
Material examined. (152 specimens, 28–79 mm
SL). HOLOTYPE – USNM 384198, male, 79 mm SL,
10°34’45’’N, 122°30’30’’E, 0–4 m, small cove lined with
volcanic rock and with rocky boulders in cove, volcanic
rock boulders and sandy channels, coral over part of
area; Guimaras Island, Pulang Duta, Sinabsapan, Panay,
Philippines, J. T. Williams, D. G. Smith, K. E. Carpenter,
C. Carpenter, N. Minsalan and T. Minsalan, 24 Sept.
1995. PARATYPES - USNM 209801, 1 male, 53 mm SL,
Indonesia, Ambon, off Tandjung Suli, shallow coral reef,
0–2 m, V. G. Springer and M. F. Gomon, 11 Jan. 1973;
CAS 227311, 1 female, 45 mm SL, 05°14’S, 145°47’E,
Papua New Guinea, Madang, Nagada Harbour, 1–7 m,
S. G. Poss, D. Catania et al., 5 December 1987; USNM
365822, 2 males, 57 and 62 mm SL, Papua New Guinea,
Papua, Harvey Bay, east of Oro Bay, 0–10 m, T. R. Roberts,
6–7 August 1975; USNM 365832, 1 male, 55 mm SL, 2
females, 46 and 58 mm SL, 05°11’S, 145°50’E, Papua
New Guinea, Kranket Island, mangrove right up to coral,
0–1 m, V. G. Springer et al., 7 Nov. 1978; USNM 366567,
1 male, 50 mm SL, Solomon Islands, Bougainville, Kieta
Harbor, coral reef off main market, 1–4 m, D. M. Cohen and
party, 10 Mar. 1965; USNM 366596, 1 female, 44 mm SL,
09°06’30’’N, 122°55’24’’E, Philippines, near Giligaon, N
of Maloh, Negros, 0–2 m, L. W. Knapp et al. 26 April 1979;
USNM 376183, 1 male, 47 mm SL, 04°14’S, 152°10’E,
Papua New Guinea, Bismarck Archipelago, Rabaul, New
Britain, W. Davis et al., 27 Feb. 1965; USNM 394981, 3
females, 40, 52 and 62 mm SL, 1 male 48 mm SL, same
data as holotype; WAM P.30635-001, 1 male, 45 mm SL,
Papua New Guinea, Madang, 0–4 m, G. R. Allen et al., 10
Feb. 1993; WAM P.29624-044, 5 females, 37–52 mm SL,
Table 7. Meristic and morphometric characters of Alionematichthys
plicatosurculus sp. nov.
Holotype
USNM
384198
79
Standard length in mm
Meristic characters
Dorsal in rays
77
Caudal in rays
16
Anal in rays
64
Pectoral in rays
20
Precaudal vertebrae
12
Caudal vertebrae
31
Total vertebrae
43
Rakers on anterior gill arch
15
Pseudobranchial ilaments
2
D/V
6
D/A
20
V/A
14
Morphometric characters in % of SL
Head length
24.7
Head width
14.5
Head height
16.6
Snout length
5.3
Upper jaw length
13.3
Diameter of pigmented eye
2.2
Diameter of pupil
1.2
Interorbital width
7.5
Posterior maxilla height
4.5
Postorbital length
17.8
Preanal length
45.9
Predorsal length
30.4
Body depth at origin of anal in
18.2
Pectoral in length
13.6
Pectoral in base height
6.0
Ventral in length
22.0
Base ventral in - anal in origin
22.3
Holotype +
29 paratypes
n
Mean (range)
48.9 (28-79)
30
80.3 (76-84)
16
62.6 (60-67)
20.2 (19-21)
12.0 (11-12)
31.1 (30-33)
43.1 (42-45)
16.3 (14-18)
2
5.9 (5-6)
22.3 (19.0-25.0)
13.4 (11.0-19.0)
28
7
28
11
27
27
27
11
11
27
27
27
26.0 (24.7-28.1)
13.6 (11.7-15.6)
16.0 (13.8-17.7)
5.9 (5.1-7.3)
13.4 (12.5-14.5)
2.6 (2.2-3.0)
1.6 (1.2-2.2)
7.2 (6.7-8.1)
4.4 (3.8-4.8)
18.5 (17.6-19.2)
46.1 (43.5-47.7)
31.3 (29.8-33.0)
18.4 (17.1-19.4)
13.9 (11.9-16.0)
5.9 (5.2-6.7)
23.5 (19.4-25.8)
26.6 (22.3-28.5)
11
11
11
11
11
11
11
11
11
11
10
11
11
11
11
14
11
9 males, 28–52 mm SL, 05°09’S, 145°48’E, Papua New
Guinea, Madang, G. R. Allen, 15 Oct. 1987.
Fig. 14. Alionematichthys plicatosurculus sp. nov., USNM 394198, holotype, male, 79 mm SL.
109
W. Schwarzhans and P. R. Møller
Fig. 15. Alionematichthys plicatosurculus sp. nov., A, lateral view of head, holotype; B, ventral view of head, holotype; C, lateral view of
head, paratype, USNM 394981, female, 52 mm SL; D, inclined lateral view of male copulatory organ, holotype; E, view of left pseudoclaspers
from inside, holotype; F, ventral view of left pseudoclaspers, holotype; G, ventral view of male copulatory organ, holotype; H, view of left
pseudoclaspers from inside, USNM 366567, 50 mm SL; I, median view of right otolith, holotype; J, ventral view of right otolith, holotype.
110
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Additional material. BMNH 1907.2.28.11, 1 specimen,
(bad condition), Neu-Lauenburg/ Duke of York Islands,
Bismarck Archipelago, Papua New Guinea; USNM
263681, 33 males and 52 females, 30–70 mm SL, 09°06’N,
122°55’E, Philippines, Negros; USNM 263683, 4 males and
9 females, 33–71 mm SL, 09°03’N, 122°59’E, Philippines,
Negros; USNM 377193, 1 male and 3 females, 68–72
mm SL, 09°10’N, 123°26’E, Philippines, Siquijor Island;
USNM 377202, 8 specimens, 48–71 mm SL, 10°35’N,
122°08’E, Philippines, Panay Island; USNM 345618, 5
specimens, 11°25’N, 123°15’E, Panay, Philippines; USNM
365818, 20 specimens, Papua New Guinea, Bagabag Island;
USNM 366480, 1 female, 52 mm SL, 05°14’S, 145°47’E,
Papua New Guinea, Madang; USNM 3766222, 1 male, 66
mm SL, 05°14’S, 145°47’E, Papua New Guinea, Madang,
USNM 366508, 3 females, 40–60 mm SL, 03°23’S,
143°40’E, Papua New Guinea, Mushu Island.
Diagnosis. Vertebrae 11–12+31–33=42–45, dorsal in
rays 76–84, anal in rays 60–67; eyes moderately large
(2.2–3.0% SL); body slender, snout rounded, without cirri;
scales only on cheeks; no upper preopercular pore; outer
pseudoclasper broad-based, large, sometimes extending
beyond hood in resting position, with thickened tip,
particularly on the inner face; inner pseudoclasper large,
thick, with posterior lobe folded over anterior pointed lobe;
otolith length to height 2.1–2.3, with gently curved dorsal
rim, otolith length to sulcus length 1.6–1.7, ostium length
to cauda length 3.5–4.5.
Description (Figs 14–15).The principal meristic and
morphometric characters are shown in Table 7. Mature at
about 40 mm SL. Body slender, with rounded head proile.
No cirri on snout.
Eye size 2.2–3.0 (2.2)% SL. Head with scale patch
on cheek containing 5–6 (6) vertical rows of scales on
upper part and 2–3 vertical rows on lower part. Horizontal
diameter of scales on body of holotype about 1.7% SL,
in 32 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, with small knob. Anterior
nostril positioned high, 1/2.5–1/3 distance from upper lip
to anterior margin of eye. Posterior nostril small, about 1/5
to 1/6 the size of eye. Opercular spine with free tip, thin and
sharply pointed. Anterior gill arch with 14–18 (15) rakers,
3 elongate in row. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 15 A–C). Supraorbital
pores 3. Infraorbital pores 6 (3 anterior and 3 posterior),
three posterior pores about one-quarter the size of three
anterior pores. Mandibular pores 6 (3 anterior and 3
posterior): irst anterior pore large, tubular, without cirri.
Preopercular pores: 3 lower, irst and second with joined
opening, covered by dermal lap in lateral view; third
tubular; upper preopercular pore absent. [See description of
Alionematichthys ceylonensis for position of pores.]
Dentition (of holotype). Premaxilla with 7 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.
Vomer horseshoe-shaped, with 2 outer row of small teeth
and 1 inner row of large teeth up to 1/4 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of long teeth up to 1/3 diameter of pupil. Dentary with 5
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/2 diameter of pupil.
Otolith (Fig. 15I, J). Moderately elongate in shape,
length to height 2.1–2.3 and thin (otolith height to otolith
thickness about 3). Anterior tip broadly rounded to
slightly angular, posterior tip much expanded and slightly
ornamented. Dorsal rim gently curved, without distinct
angles. Inner face moderately convex, outer face concave
to almost lat, both smooth. Otolith length to sulcus length
1.6–1.7. Sulcus slightly supramedially positioned, slightly
inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.5–4.5. Ventral furrow feeble and
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined
and 5–8 depressed. Parapophyses present from vertebrae
6 to 11 (12). Pleural ribs on vertebrae 2 to 10 (11). First
anal in pterygiophore slightly longer than subsequent
pterygiophore.
Male copulatory organ (Fig. 15D–H). Two pairs of
moderately large pseudoclaspers. Outer pseudoclasper large,
sometimes extending beyond hood in resting position, with
thickened tip (particularly in large specimens) expressed
as massive knob on inner face; inner pseudoclasper large,
thick, with posterior lobe very large and folded over anterior
pointed lobe. Isthmus moderately narrow; penis curved,
about as long as outer pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly brownish.
Comparison. Alionematichthys plicatosurculus
belongs to the species group without an upper preopercular
pore together with A. ceylonensis, A. phuketensis and
A. shinoharai sp. nov., from which it differs in the peculiar
shape of the thick inner pseudoclasper with the posterior
lobe folded over the anterior lobe and the massive tip of the
inner face of the outer pseudoclasper. For further distinction
from A. ceylonensis and A. phuketensis see respective
descriptions. From A. shinoharai sp. nov. it differs further in
the lack of scales above the opercular spine (vs 2), the long
sulcus (otolith length to sulcus length 1.5–1.6 vs 1.9–2.0)
and the notch at the ventral sulcus margin at the joint of
ostium and cauda (vs no notch).
Distribution. Alionematichthys plicatosurculus is
known from the Philippines south of 12°N, Ambon and
along the northern coast of New Guinea to the Bismarck
Archipelago and Solomon Islands (Fig. 1).
Etymology. Named after the characteristic folded inner
pseudoclasper by a combination of plicatus (Latin = folded)
and surculus (Latin = grapevine tendril) used for the term
pseudoclasper. It is a noun in apposition.
111
W. Schwarzhans and P. R. Møller
Fig. 16. Sample sites of Alionematichthys riukiuensis (Aoyagi, 1954), Alionematichthys aff. riukiuensis 1, Alionematichthys aff.
riukiuensis 2, A. samoaensis sp. nov., A. winterbottomi sp. nov., Alionematichthys sp. 1, Alionematichthys sp. 2. One symbol may
represent several samples.
Fig. 17. Alionematichthys riukiuensis (Aoyagi, 1954), A, fresh dead, WAM P.30909-002 [exact specimen and size not known],
B, NMST-P 55637, male, 74 mm SL.
112
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Reef, Australia, 1–10 m, D. F. Hoese and party, 17 Nov.
1975; AMS I.20770-015, 42 specimens, 11°55’S, 143°27’E,
Sir Charles Hardy Island, Queensland, Australia; AMS
I.20770-073, 11 specimens, 11°55’S, 143°27’E, Sir Charles
Hardy Island, Queensland, Australia; AMS I. 21422-029, 5
specimens, 14°S, 145°E, Coral Sea; AMS I.33740-026, 1
male, 10°S 144°E, Coral Sea; AMS I.34311-010, 1 male,
5 females, 22°14’S, 150°19’E, Australia, Queensland,
Cannibal Group; AMS I.37315-032, 3 males and 4 females,
18°44’S, 169°12’E, Vanuatu, Erromango; AMS I.37322009, 4 females, 19°31’S, 169°29’E, Vanuatu, Tanna; AMS
I.37339-076, 1 male and 1 female, 16°47’S, 168°21’E,
Vanuatu, Epi; AMS I.37922-021, 1 male, and 1 female,
13°52’S, 167°32’E, Vanuatu, Banks Islands; BPBM 5921,
1 female, Vanuatu, Efate; BPBM 11465, 2 females, New
Caledonia; BPBM 40934, 1 male, 2 females, Ryukyu
Islands; BPBM 40936, 2 females, Similan Island Thailand;
CAS 65664, 1 male, Papua New Guinea, Madang; KAUM-I.
11459, 1 male, 61 mm SL, 30°27’23’’N, 130°29’59’’E, Isso,
Yakushima Island, Kagoshima, Japan; KAUM-I. 10661,
1 male, 48 mm SL, off Okinoerabu Island, Kagoshima,
Japan; KAUM-I. 11484, 1 male, 55 mm SL, 30°27’23’’N,
130°29’59’’E, Isso, Yakushima Island, Kagoshima, Japan;
KSHS 3650 (otolith only); MNHN 1980-0851, 3 females,
New Caledonia; NSMT-P 55636, female 74 mm SL,
China, Hainan Island, China; NSMT-P 55637, 1 male,
74 mm SL, Hainan Island, China; NTM S.13425-007, 1
female, 11°58’S, 123°21’E, Ashmore Islands, Australia;
ROM 71841, 9 specimens, Vietnam; ROM 71843, 11
specimens, Vietnam; ROM 71850, 5 specimens, 20°N
107°E; SMNS 26429, 5 specimens, eastsoutheast of Thio,
New Caledonia, 21°38’57’’S, 166°18’31’’E, 0–8.5 m;
SMNS 26430, 6 specimens, Maré, Cap Wabao, Loyalty
Islands, 21°35’45’’S 167°50’06’’E, 2–3.8 m; SMNS 22705,
6 specimens, northnorthwest of Ouégoa, New Caledonia,
20°15’20’’S, 164°24’10’’E, 0.5–3.5 m; SMNS 22804, 5
specimens, east-south-east of Ponérihuen, New Caledonia,
21°04’55’’S 165°28’10’’E, 0.3–2.8 m; SMNS 26431, 2
specimens, north-west of Hienghène, New Caledonia,
20°31’49’’S 164°47’01’’E, 0.2–3.5 m; USNM 199541, 9
males and 14 females, 42–66 mm SL, 04°01’S, 101°01’E,
Mentawai, Sumatra; USNM 394978, 5 specimens, 21°N
120°E; EX USNM 345618, 7 males, 29–65 mm SL, 1
female, 40 mm SL, juvenile 20 mm SL, 11°25’N, 123°15’E,
Panay, Philippines; USNM 394982, 31 specimens, 16°47’S,
168°21’E, Epiu, Vanuatu; USNM 355385, 38 specimens,
Tanna, Vanuau; USNM 394984, 1 female, 75 mm SL,
17°31’S, 168°19’, Efate, Vanuatu; USNM 362434, 1
male, 64 mm SL, 3 females, 4 juveniles, 14°S 167°E,
Banks Islands, Vanuatu; USNM 366490, 3 males, 72–78
mm SL, 3 females, 57–72 mm SL, 08°29’N, 97°39’E,
Similan Islands, Thailand; USNM 366600, 1 male, 62
mm SL, 09°19’N, 123°18’E, Negros, Philippines; USNM
376188, 1 male, 150 mm SL, 2 females, 116–119 mm SL,
Taiwan; WAM P.30305-037, 1 male, 2 females, 13°52’S,
126°56’E, Sir Graham Moore Islands, Western Australia;
Alionematichthys riukiuensis (Aoyagi, 1954)
(Figs 16–19, Tables 1, 8)
Dinematichthys riukiuensis Aoyagi, 1954: 235;
Machida 1992: 270; Machida 1994: 461; Hayashi 1995:
11; Nielsen et al. 1999: 130.
Brotulina fusca. — Matsubara 1955: 800 (not Diancistrus
fuscus Fowler, 1946).
Dinematichthys megasoma Machida, 1994: 459; Nielsen
et al. 1999: 130; Schwarzhans and Møller 2005: 78.
Material examined. 536 specimens, 19–150 mm SL.
PARALECTOTYPE – YCM-P 30001, female, 72 mm,
Okinawa Island, Ryukyu Islands; NON-TYPES – AMS
I.37399-076, male, 73 mm SL, Vanuatu; AMS I.18739-037,
1 male and 2 females, 14°42’S, 145°27’E, Lizard Island,
Queensland, Australia; AMS I.19108-032, 2 males and 3
females, 14°40’S, 145°28’E, Lizard Island, Great Barrier
Table 8. Meristic and morphometric characters of Alionematichthys
riukiuensis (after Aoyagi 1954).
Holotype +
n
non-types
Holotype
YCM-P
30023*
Mean (range)
Standard length in mm
90
72.3 (20-148) 121
Meristic characters
Dorsal in rays
79
83.9 (77-92) 107
Caudal in rays
16
15.7 (14-16)
40
Anal in rays
61
65.1 (59-72) 107
Pectoral in rays
22
20.5 (19-23)
11
Precaudal vertebrae
11
11.3 (11-12) 121
Caudal vertebrae
31
31.5 (29-33) 120
Total vertebrae
42
42.8 (41-44) 120
Rakers on anterior gill arch
19.1 (15-21)
10
Pseudobranchial ilaments
1.9 (0-3)
9
D/V
5.7 (5-7)
111
D/A
23.7 (19-27) 112
V/A
13.1 (12-14) 112
Morphometric characters in % of SL
Head length
29.3
26.8 (25.7-29.3) 12
Head width
14.8 (11.6-18.6) 11
Head height
17.1 (14.3-20.5) 11
Snout length
7.3
5.8 (4.7-7.3)
12
Upper jaw length
14.3
13.9 (12.4-15.1) 11
Diameter of pigmented eye
3.0
2.6 (2.5-3.0)
12
Diameter of pupil
1.4 (1.2-1.6)
10
Interorbital width
7.7
7.3 (6.2-8.0)
11
Posterior maxilla height
5.3
4.6 (3.8-5.3)
11
Postorbital length
20.6
19.5 (19.0-20.6) 12
Preanal length
49.8
47.1 (45.7-49.8) 11
Predorsal length
33.0
31.2 (29.9-33.0) 11
Body depth at origin of anal in 21.4
19.9 (16.8-23.4) 11
Pectoral in length
15.7
14.4 (12.8-16.1) 10
Pectoral in base height
7.5
6.1 (4.4-7.5)
11
Ventral in length
20.4
21.6 (20.0-24.8) 12
Base ventral in - anal in origin
26.5 (21.8-30.9) 10
* Data from Machida (1994) included.
113
W. Schwarzhans and P. R. Møller
Fig. 18. Alionematichthys riukiuensis (Aoyagi, 1954), A, lateral view of head, ROM 71841, male, 73 mm SL; B, lateral view of head, WAM
P.30308-001, male, 112 mm SL; C, ventral view of head, WAM P.30308-001, male, 112 mm SL; D, lateral view of head, AMS I.34311010, female, 69 mm SL; E, inclined lateral view of male copulatory organ, USNM 394982, 92 mm SL; F, view of left pseudoclaspers from
inside, USNM 394982, 92 mm SL; G, ventral view of male copulatory organ, SMNS 26430, 130 mm SL; H, view of left pseudoclaspers
from inside, USNM 376188, 150 mm SL; I, ventral view of left pseudoclaspers, WAM P.30308-001, 112 mm SL; J, median view of right
otolith, WAM P.30308-001, 112 mm SL.
114
Dinematichthyine ishes of the Indo-west Paciic, Part IV
WAM P.30308-001, 7 specimens, 13°S, 125°E, Timor Sea;
WAM P.30909-002, 8 specimens, 16°S, 123°E, North West
Australia; YCM-P44101, 1 male and 2 females, Ishigaki,
Ryukyu Islands; YCM-P44102, 3 specimens, Kakeroma
Island, Ryukyu Islands.
Additional material. USNM 151418, 1 male, 59 mm SL,
Philippines; USNM 151420, 1 male, 74 mm SL, Philippines;
USNM 151422, 1 female 73 mm SL, Philippines; USNM
151423, 1 female, 82 mm SL, Philippines; USNM 151424,
1 male, 84 mm SL, Philippines; USNM 151425, 1 female,
102 mm SL, Philippines; USNM 151426, 1 female, 50
mm SL, Philippines; USNM 164589, 1 female, 78 mm
SL, Espiritu Santo, Vanuatu; USNM 376160, 1 male and
3 females, 62–70 mm SL, Taiwan; USNM 377258, 15
specimens, 38–83 mm SL, 02°35’S, 150°47’E, Kavieng,
New Ireland, Papua New Guinea; USNM 263666, 5 males
and 9 females, 34–83 mm SL, 16°25’N, 119°54’E, Luzon,
Philippines; USNM 263680, 2 males, 59–109 mm SL,
23°29’S, 152°05’E, One Tree Island, Queensland, Australia;
USNM 377214, 25 specimens, 47–109 mm SL, 09°06’N,
122°55’E, Negros, Philippines; USNM 376347, 1 female,
61 mm SL, 10°52’N, 120°56’E, Palawan, Philippines;
USNM 376225, 1 female, 49 mm SL, 10°55’N, 121°02’E,
Palawan, Philippines; USNM 263700, 1 male, 68 mm
SL, 09°04’N, 123°16’E, Apo Island, Philippines; USNM
263743, 5 specimens, 34–76 mm SL, Milne Bay, Papua
New Guinea; USNM 263755, 15 specimens, 31–119 mm
SL, Taiwan; USNM 377208, 1 male and 3 females, 79–104
mm SL, 10°35’N, 122°08’E, Panay, Philippines; USNM
376163, 6 females, 68–92 mm SL, 18°44’S, 169°12’E,
Erromango, Vanuatu; USNM 355152, 1 female, 100 mm
SL, 17°03’S, 168°21’E, Emae, Vanuatu; USNM 356612,
10 specimens, 19–95 mm SL, Tomotu, Santa Cruz Islands,
Solomon Islands; USNM 376167, 3 females, 35–51 mm
SL, 13°32’S, 167°20’, Banks Islands, Vanuatu; USNM
363192, 1 male and 1 female, 33–75 mm SL, Banks
Islands, Vanuatu; USNM 376166, 3 females, 30–88 mm
SL, 13°04’S, 167°39’E, Banks Islands, Vanuatu; USNM
365819, 7 specimens, 44–72 mm SL, Bagabag Island, Papua
New Guinea; USNM 376223, 2 males and 4 females, 55–80
mm SL, 05°14’S, 145°47’E, Madang, Papua New Guinea;
USNM 366482, 16 males and 39 females, 32–75 mm SL,
10°47’S, 152°24’E, Louisade Archipelago, Deboyne Atoll,
Papua New Guinea; USNM 376202, 1 female, 55 mm SL,
05°14’S, 145°47’E, Madang, Papua New Guinea; USNM
376190, 1 female, 65 mm SL, 05°14’S, 145°47’E, Madang,
Papua New Guinea; USNM 366521, 10 specimens, 32–80
mm SL, 05°10’S, 145°51’E, Kranket Island, Papua New
Guinea; USNM 366522, 2 males and 1 female, 40 mm SL,
01°31’S, 145°01’E, Hermit Islands, Papua New Guinea;
USNM 366524, 1 male, 67 mm SL, 05°14’S, 145°47’E,
Madang, Papua New Guinea; USNM 366557, 1 male and
1 female, 51–73 mm SL, 21°55’N, 120°44’E, Taiwan;
USNM 366559, 1 female, 56 mm SL, Taiwan; USNM
374226, 6 specimens, 35–103 mm SL, Taiwan; USNM
366576, 1 female, 52 mm SL, 01°12’S, 144°16’E, Ninigo
Atoll, Papua New Guinea; USNM 376173, 3 males and 3
females, 64–104 mm SL, 23°29’S, 152°05’E, One Tree
Island, Queensland, Australia; USNM 376171, 1 male and
1 female, 63 mm SL, 23°25’S, 151°55’E, Heron Island,
Queensland, Australia; USNM 366682, 1 female, 82
mm SL, 23°25’S, 151°55’E, Heron Island, Queensland,
Australia; USNM 366688, 1 male, 38 mm SL, Taiwan;
USNM 366693, 7 specimens, 21–82 mm SL, Taiwan.
Tentatively assigned specimens (A. aff. riukiuensis).
BPBM 40938, 5 males and 2 females,Taiwan; USNM
309643, 3 males and 3 females, 40–60 mm SL, 04°52’N,
119°26’E, Sibutu Island, Philippines.
Diagnosis. Vertebrae 11–12+30–32=41–44, dorsal in
rays 77–92 (average 84), anal in rays 59–72 (average 65);
eyes moderately large (2.5–3.0% SL); body moderately
slender, massive, snout with many cirri; scales on cheeks
and large scale patch above opercular spine (6–17), large
specimens with few scales also below opercular spine;
upper preopercular pore present; outer pseudoclasper broadbased, not very large, not extending beyond hood in resting
position; inner pseudoclasper not much smaller than outer
pseudoclasper, anterior and posterior lobes of equal size,
anterior lobe broadly connected to anterior rim of outer
pseudoclasper; otolith slender, length to height 2.2–2.4,
with gently curved dorsal rim, otolith length to sulcus length
1.6–1.7, ostium length to cauda length 3.5–4.0.
Description (Figs 17–19).The principal meristic and
morphometric characters are shown in Table 8. One of the
largest species attaining up to 150 mm SL; mature at about
50–55 mm SL. Body moderately slender, massive, with
moderately pointed head proile. Many cirri on snout. Eye
size 2.5–3.0 (3.0)% SL. Head with scale patch on cheek
containing 8–13 vertical rows of scales on upper part and
4–5 vertical rows on lower part, scale patch above opercular
spine with 6–17 scales, in adults, scaled patch below
opercular spine with up to 8 scales. Horizontal diameter
of scales on body of a 74 mm SL male, about 1.5% SL, in
about 40 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded with small knob. Anterior
nostril positioned high, 1/2–2.5 distance from upper lip to
anterior margin of eye. Posterior nostril small, about 1/6 to
1/8 the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 15–21 rakers, 3 elongate. Small
plate-like raker between lower two elongate rakers in most
specimens. Pseudobranchial ilaments 0–3 (usually 2).
Head sensory pores (Fig. 18A–D). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): irst anterior
pore large, tubular, with cirri. Preopercular pores: 3 lower,
irst and second with joined opening, covered by dermal
lap in lateral view; third tubular; upper preopercular pore
present. [See description of Alionematichthys ceylonensis
for position of pores.]
115
W. Schwarzhans and P. R. Møller
Fig. 19. Alionematichthys aff. riukiuensis (Aoyagi, 1954), A, lateral view of head, BPBM 40938, male, 62 mm SL; B, ventral view of head,
BPBM 40938, male, 62 mm SL; C, lateral view of head, USNM 309643, male, 58 mm SL; D, inclined lateral view of male copulatory organ,
BPBM 40938, male, 80 mm SL; E, ventral view of male copulatory organ, BPBM 40938, male, 80 mm SL; F, view of left pseudoclaspers from
inside, USNM 309643, 58 mm SL; G, view of left pseudoclaspers from inside, BPBM 40938, male, 62 mm SL; H, view of left pseudoclaspers
from inside, BPBM 40938, male, 80 mm SL; I, median view of right otolith, USNM 309643, male, 58 mm SL.
116
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Dentition (of a 74 mm SL male, NSMT-P 55637).
Premaxilla with 6 outer rows of granular teeth and 2 inner
rows of larger teeth anteriorly. Anteriormost teeth in inner
row up to 1/2 diameter of pupil. Vomer horseshoe-shaped,
with 6 outer rows of small teeth and 1 inner row of large
teeth up to 1/4 diameter of pupil. Palatine with 4 outer
rows of small teeth and 1 inner row of long teeth up to 1/4
diameter of pupil. Dentary with 6 outer rows of granular
teeth and 1 inner row of larger teeth anteriorly, up to about
1/2 diameter of pupil.
Otolith (Fig. 18J). Elongate in shape, length to height
2.2–2.4 and thin (otolith height to otolith thickness about 3).
Anterior tip slightly pointed, posterior tip pointed, expanded
and irregularly ornamented. Dorsal rim gently curved,
without distinct angles. Inner face moderately convex, outer
face slightly concave to almost lat, both smooth. Otolith
length to sulcus length 1.6–1.7. Sulcus medianly positioned,
not inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.5–4.0. Ventral furrow distinct and
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to
11 (12). Pleural ribs on vertebrae 2 to 11 (12). First anal in
pterygiophore not or only slightly longer than subsequent
pterygiophore.
Male copulatory organ (Fig. 18E–I). Two pairs of
rather small pseudoclaspers, not extending beyond hood
in resting position. Outer pseudoclasper a broad based
lap; inner pseudoclasper slightly shorter, very thick (best
seen in ventral view: Fig. 18G), with anterior pointed lobe
broadly connected by ligament to anterior rim of outer
pseudoclasper, posterior lobe broad, about equal in size.
Isthmus moderately narrow; penis curved, slightly longer
than outer pseudoclaspers.
Colour. Live colour brownish-red or orange-brown
(SMNS 22804), sometimes with yellowish vertical ins
(Fig. 17A). Preserved colour medium to dark brown.
Variability. Alionematichthys riukiuensis shows some
variation in the number of scales on the opercle above and
below the opercular spine and a certain variation of the
dorsal and anal in ray counts, both of which does not seem
to follow a geographic pattern.
Remarks. Machida (1994) described Dinematichthys
megasoma from Australia. These specimens agree well
with the variability observed in A. riukiuensis, including
the characters regarded by Machida as distinctive from
Dinematichthys riukiuensis, i.e. the short tubular posterior
nostril, the widely scaled opercle and the lateral scale row
counts. We have, however, observed two speciic lots which
somewhat depart from the general range of variability
(see Fig. 19). They are both notable for few or no cirri on
the snout and few scales above the opercular spine (2–6)
(Fig. 19A–C). They vary amongst themselves in meristics
(see Table 1, 8). Their pseudoclaspers resemble those of
A. riukiuensis (Fig. 19D–H) perfectly as do their otoliths
(Fig. 19I). Due to the limited amount of specimens available
and the high degree of similarity with A. riukiuensis they are
here referred to as Alionematichthys aff. riukiuensis.
Comparison. Alionematichthys riukiuensis belongs to
the species group with an upper preopercular pore, large
eyes and (many) scales above the opercular spine and cirri
on the snout (comprising also A. winterbottomi sp. nov.). It
differs mainly in the characters of the pseudoclaspers and
the presence of scales below the opercular spine in adults
(vs always absent in A. winterbottomi sp. nov.). It differs
from A. piger and A. samoaensis sp. nov., which likewise
show many cirri on the snout in the presence of many scales
above the opercular spine (vs none or rarely 1–2) and the
pseudoclasper morphology. Also, A. riukiuensis is notable
for the large size it can attain and at which it matures, greater
than in all other species of Alionematichthys.
Distribution. Alionematichthys riukiuensis is widely
distributed in the Indo-west Pacific from the Ryukyu
Islands, Hainan Island and the west coast of Thailand in the
north to the northern shores of Australia, New Caledonia
and Vanuatu in the south (Fig. 16).
117
Alionematichthys samoaensis sp. nov.
(Figs 16, 20, 21, Tables 1, 8)
Material examined. (36 specimens, 22–59 mm SL).
HOLOTYPE – CAS 81486, male, 46 mm SL, ca. 11°03'S,
171°04'W, seaward face of leeward reef on Swains Island,
American Samoa, coll. L. R. Taylor Jr., 23 May 1967.
PARATYPES – AMS IB. 6513, male, 54 mm SL, 14°27'S,
178°05'W, Aloi, Niue, W. N. McDowall, May 1963; BPBM
24122, 2 females, 56 and 59 mm SL, Samoa, Tutuila, R.
C. Wass, 1976-1977; CAS 227305, 20 females, 22–53 mm
SL, 11 males, 25–47 mm SL, juvenile, 22 mm SL, same
data as holotype.
Diagnosis. Vertebrae 11–12+29–31=40–43, dorsal in
rays 73–80, anal in rays 57–64; eyes moderately large
(2.2–2.8% SL); body slender, snout with many cirri;
scales on cheeks, no scales above opercular spine; upper
preopercular pore present; posterior nostril funnel-shaped;
outer pseudoclasper broad-based, not extending beyond
hood in resting position, with thick distal knob on inner face;
inner pseudoclasper about half size of outer pseudoclasper,
anterior thorn slightly bent, posterior lobe broader than
anterior thorn.
Description (Figs 20, 21).The principal meristic and
morphometric characters are shown in Table 9. Body
moderately slender, with moderately pointed head proile.
Many cirri on snout. Eye size 2.2–2.8 (2.2)% SL. Head
with scale patch on cheeks containing 6–7 vertical rows of
scales on upper part and 2–3 vertical rows on lower part,
no scale patch on opercle. Horizontal diameter of scales
on body in holotype 1.7% SL, in about 24 horizontal rows.
Maxillary ending far behind eye, dorsal margin of maxillary
covered by upper lip dermal lobe, posterior end expanded.
Anterior nostril positioned high, one-half distance from
upper lip to anterior margin of eye. Posterior nostril with
W. Schwarzhans and P. R. Møller
Fig. 20. Alionematichthys samoaensis sp. nov., CAS 81486, holotype, male, 46 mm SL.
Fig. 21. Alionematichthys samoaensis sp. nov., A, lateral view of head, holotype; B, ventral view of head, holotype; C, inclined lateral view
of male copulatory organ, CAS 81486, male, 39 mm SL; D, ventral view of male copulatory organ, holotype; E, view of left pseudoclaspers
from inside, holotype; F, view of left pseudoclaspers from inside, CAS 81486, 39 mm SL.
broad funnel-shaped rim, about one-eighth the size of eye.
Opercular spine with free tip, pointed. Anterior gill arch
with 14–18 (15) rakers, 3 elongate. In holotype and a few
paratypes lower two elongate rakers interrupted by one
plate-like raker. In other paratypes, three elongate rakers in
row. Pseudobranchial ilaments 1–2 (usually 2).
118
Head sensory pores (Fig. 21A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): irst
anterior pore large, tubular, with cirri. Preopercular pores:
3 lower, irst and second with joined opening, covered by
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Dentition (of holotype). Premaxilla with 5 outer
rows of granular teeth and 1 inner row of larger teeth
anteriorly. Anteriormost teeth in inner row up to 1/3
diameter of pupil. Vomer horseshoe-shaped, with 2 outer
rows of small teeth and 1 inner row of large teeth up to
1/3 diameter of pupil. Palatine with 1 outer row of small
teeth and 1 inner row of long teeth up to 1/3 diameter of
pupil. Dentary with 5 outer rows of granular teeth and
1 inner row of larger teeth anteriorly, up to about 2/3
diameter of pupil.
Otolith (not known).
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to 11
(12). Pleural ribs on vertebrae 2 to 10–11 (12). First anal in
pterygiophore not or only slightly longer than subsequent
pterygiophore in females, much longer in males, but not
reaching last precaudal parapophysis.
Male copulatory organ (Fig. 21C–F). Two pairs of rather
small pseudoclaspers, not extending beyond hood in resting
position. Outer pseudoclasper a broad based lap, with
thick knob distally on inner face, its supporter somewhat
bent backward at tip; inner pseudoclasper thick, about half
the size of outer pseudoclasper, anterior lobe slightly bent,
posterior lobe broader than anterior thorn. Isthmus narrow;
penis curved, slightly longer than outer pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly light brown.
Comparison. Alionematichthys samoaensis belongs to
the species group with an upper preopercular pore, large
eyes and no scales above the opercular spine, comprising
also A. piger and A. suluensis sp. nov. Alionematichthys
samoaensis obviously is closely related to A. piger, which
it replaces in region of Samoa, differing mainly in the
distinct knob on the inner face of the outer pseudoclasper
(vs thin outer pseudoclasper without knob) and the funnelshaped posterior nostril (vs anterior rim elevated only).
From A. suluensis, it differs readily in the presence of many
delicate cirri on the snout (vs absent) and the anterior lobe
of the inner pseudocasper being as long as its posterior lobe
(vs twice as long).
Table 9. Meristic and morphometric characters of Alionematichthys
samoaensis sp. nov.
Holotype
CAS
81486
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
46
76
16
59
22
12
30
42
15
2
5
23
13
SL
27.2
14.2
16.4
5.5
14.7
2.2
1.4
7.8
4.7
19.8
48.7
30.7
19.4
13.4
6.5
26.0
29.6
Holotype +
35 non-types
n
Mean (range)
37.3 (22-59)
36
76.6 (73-80)
16.1 (16-17)
59.4 (57-64)
21.6 (20-22)
11.2 (11-12)
30.7 (29-31)
41.9 (40-43)
15.6 (14-18)
1.9 (1-2)
5.6 (5-6)
22.6 (20-26)
13.2 (13-14)
33
24
33
12
33
33
33
12
10
33
33
33
27.2 (26.2-28.9)
13.8 (11.8-16.2)
16.6 (15.3-17.9)
6.1 (5.5-6.5)
14.3 (13.3-15.6)
2.5 (2.2-2.8)
1.4 (1.2-1.5)
7.6 (7.2-8.0)
4.7 (3.9-5.3)
19.4 (18.5-21.2)
47.7 (44.8-50.4)
31.1 (28.9-32.5)
18.8 (16.3-20.4)
13.5 (12.7-15.7)
6.6 (6.2-6.8)
25.5 (22.5-27.1)
28.3 (25.8-31.6)
10
10
10
9
10
9
9
10
10
10
10
10
10
10
10
10
10
dermal lap in lateral view; third tubular; upper preopercular
pore present. [See description of Alionematichthys
ceylonensis for position of pores.]
Fig. 22. Alionematichthys shinoharai sp. nov., NSMT-P 34895, holotype, male, 34 mm SL.
119
W. Schwarzhans and P. R. Møller
Fig. 23. Alionematichthys shinoharai sp. nov., A, lateral view of head, holotype; B, ventral view of head, holotype; C, view of left pseudoclaspers
from inside, holotype; D, inclined lateral view of male copulatory organ, holotype; E, median view of right otolith, holotype; F, ventral view
of right otolith, holotype; G, median view of right otolith, YCM-P36416, female, 42 mm SL.
Distribution. Alionematichthys samoaensis is endemic
to the Islands of the Samoa group, and is in fact the
only endemic Dinematichthyini restricted to that region
(Fig. 16).
Biology. A 46 mm SL female (CAS 81486) contains ca.
100 embryos, length 4.6 mm TL.
Each embryo with three short pigmented rows on the
posterior part of the body.
Etymology. Named after the type locality, American
Samoa. It is an adjective.
Alionematichthys shinoharai sp. nov.
(Figs 1, 22, 23, Tables 1, 10)
Material examined. (2 specimens, 34–42 mm SL).
HOLOTYPE – NSMT-P 34895, male, 34 mm SL,
28°11’2’’N, 129°16’E, Sakinome beach, Amami-Oshima
Island, Ryukyu Islands, Japan, 5 m, K. Matsuura and M.
Aizawa, 14 June 1991. PARATYPES – YCM-P 36416,
female, 42 mm SL, Amami Islands, Kakeroma Island,
Ryukyu Islands, Japan, 24 Aug. 1995.
Diagnosis. Vertebrae 11+32–33=43–44, dorsal in rays
83–85, anal in rays 67; eyes large (3.0–3.3% SL); body
slender, snout pointed, without cirri; scales on cheeks,
two scales above opercular spine; upper preopercular
pore absent; outer pseudoclasper broad-based simple lap;
inner pseudoclasper very small, somewhat depressed;
otolith moderately elongate, length to height 2.1–2.2, with
regularly curved dorsal rim, sulcus small, otolith length
to sulcus length 1.9–2.0, cauda very small, ostium length
to cauda length 5.5–7, ostium height to cauda height
2.2–2.7.
Description (Figs 22–23).The principal meristic and
morphometric characters are shown in Table 10. Mature at
120
about 35–40 mm SL (male of 34 mm SL probably subadult).
Body slender, with pointed head proile. No cirri on snout.
Eye size 3.3 (3.0)% SL. Head with scale patch on cheeks
containing 7 vertical rows of scales on upper part and 4
vertical rows on lower part, 2 scales on opercle above
opercular spine. Horizontal diameter of scales on body of
holotype 1.5% SL, in about 24 horizontal rows. Maxillary
ending far behind eye, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded with small
knob. Anterior nostril positioned high, one-third distance
from upper lip to anterior margin of eye. Posterior nostril
with narrow elevated rim, about one-eighth the size of eye.
Opercular spine with free tip, sharply pointed. Anterior
gill arch with 15 (16) rakers, 3 elongate. Pseudobranchial
ilaments 2.
Head sensory pores (Fig. 23A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): irst
anterior pore large, tubular, with single cirrus anteriorly.
Preopercular pores: 3 lower, irst and second with joined
opening, covered by dermal lap in lateral view; third
tubular; upper preopercular pore absent. [See description
of Alionematichthys ceylonensis for position of pores.]
Dentition (of holotype). Premaxilla with 5 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.
Vomer horseshoe-shaped, with 1 outer row of small teeth
and 1 inner row of larger teeth up to 1/5 diameter of pupil.
Palatine with 1 outer row of small teeth and 1 inner row
of longer teeth up to 1/4 diameter of pupil. Dentary with 5
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Dorsal rim gently curved, with postdorsal angle in male
specimen (Fig. 23E) and absent in female (Fig. 23G),
exhibiting slight sexual dimorphism. Inner and outer face
moderately convex, smooth. Otolith length to sulcus length
1.9–2.0. Sulcus medianly positioned, slightly inclined, with
separated colliculi, without marked notch at ventral rim
of sulcus at joint of ostium with cauda. Cauda very small,
ostium length to cauda length 5.5–7, ostium height to cauda
height 2.2–2.7. Ventral furrow feeble, close to ventral rim
of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to 11.
Pleural ribs on vertebrae 2 to 10. First anal in pterygiophore
only slightly longer than subsequent pterygiophore.
Male copulatory organ (Fig. 23C, D). Two pairs of
small pseudoclaspers, which may not be fully mature in
single male specimen investigated. Outer pseudoclasper a
simple lap with broad base; inner pseudoclasper a small
depressed lap without signiicant indentation. Isthmus
moderately narrow; penis slightly curved, longer than outer
pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly light brown.
Comparison. Alionematichthys shinoharai belongs
to the species group without an upper preopercular
pore together with A. ceylonensis, A. phuketensis and
A. plicatosurculus. It differs from all these in the presence
of two scales above the opercular spine (vs none), the
somewhat less highly positioned anterior nostril with onethird distance from upper lip to anterior margin of eye (vs
1/2–2.5) and the peculiar otolith with its short sulcus and the
very small cauda. The combination of these characters is so
distinctive that they could indicate a separate genus, which,
however, will be subject to review of a larger male specimen
in order to fully evaluate the pseudoclasper pattern.
Distribution. Alionematichthys shinoharai has so far
only been found at the northern islands of the Ryukyu Island
chain of Japan, i.e. Amami-Oshima (Fig. 1).
Etymology. Named in honour of Gento Shinohara,
NSMT, Tokyo, in recognition of his many contributions to
ichthyology and for his kind support of the present revision.
It is a noun in apposition.
Table 10. Meristic and morphometric characters of Alionematichthys
shinoharai sp. nov.
Holotype
Paratype
NSMTYCMP34895
P36416
Standard length in mm
33
42
Meristic characters
Dorsal in rays
83
85
Caudal in rays
14
14
Anal in rays
67
67
Pectoral in rays
22
22
Precaudal vertebrae
11
11
Caudal vertebrae
33
32
Total vertebrae
44
43
Rakers on anterior gill arch
16
15
Pseudobranchial ilaments
2
2
D/V
6
5
D/A
20
24
V/A
13
14
Morphometric characters in % of SL
Head length
27.2
26.7
Head width
11.9
12.8
Head height
15.9
16.2
Snout length
5.6
6.5
Upper jaw length
13.8
13.8
Diameter of pigmented eye
3.0
3.3
Diameter of pupil
2.0
2.1
Interorbital width
6.5
7.0
Posterior maxilla height
4.0
4.0
Postorbital length
19.8
18.0
Preanal length
44.8
44.9
Predorsal length
32.0
32.6
Body depth at origin of anal in
16.1
18.1
Pectoral in length
14.1
15.1
Pectoral in base height
6.6
5.5
Ventral in length
24.7
22.6
Base ventral in - anal in origin
25.2
24.1
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/3 diameter of pupil.
Otolith (Fig. 23E–G). Moderately elongate in shape,
length to height 2.1–2.2 (34–40 mm SL), rather thick
(otolith height to otolith thickness about 2). Anterior tip
sharply pointed, posterior tip pointed, slightly expanded.
Fig. 24. Alionematichthys suluensis sp. nov., AMS I.40149-028, holotype, male, 40 mm SL.
121
W. Schwarzhans and P. R. Møller
Alionematichthys suluensis sp. nov.
(Figs 9, 24, 25, Tables 1, 11)
Material examined. (57 specimens, 26–49 mm SL).
HOLOTYPE – AMS I.40149-028, male, 40 mm SL,
12°21.70N, 121°27.57E, 0–20 m, rocky surge area at base
of cliffs, S tip of Buyamao Island off SE Mindoro Island,
Philippines, coll. MIN team, rotenone, 30 May 2000.
PARATYPES – AMS I.40149-028), 1 male, 30 mm SL, 5
females, 30–42 mm SL, 3 juveniles, 13–17 mm SL, same
data as holotype; USNM 263691, 1 male, 43 mm SL and
5 females, 41–45 mm SL, 08°51’42’’N, 123°24’36’’E,
Solino Island, Zamboanga, Mindanao, Philippines, 0–5 m,
A. Alcala et al., 4 May 1979; USNM 263694, 6 males and
12 females, 27–49 mm SL, 10°52’N, 121°12’E, Palawan,
Philippines; USNM 376216, 4 males and 1 female, 29–45
mm SL, 09°04’N, 123°08’E, Negros, Philippines; USNM
366601, 1 male, 46 mm SL, 09°04’N, 123°16’E, Apo Island,
Philippines; USNM 366602, 1 female, 34 mm SL, 09°23’N,
123°15’E, Negros, Philippines; USNM 374183, 1 male,
31 mm SL, 10°35’05’’N, 122°08’30’’E, rocky tidepool,
Talisayan Point, San Joaquin, Lawigan, Iloilo Province,
Philippines, 0–7 m, J. T. Williams et al., 25 Sep. 1995;
USNM 376179, 3 males and 6 females, 26–48 mm SL,
09°03’N, 122°59’E, Negros, Philippines; USNM 376181,
3 males, 37–44 mm SL, 10°55’05’’N, 121°02’03’’E, Putic
Island, Palawan, Philippines, 0–4.6 m, V. Springer et al.,
22 May 1978; USNM 376182, 2 males, 34 and 44 mm SL,
09°06’30’’N, 122°55’24’’E, near Giligaon, north of Maloh,
Negros, Philippines, 0–2 m, L. W. Knapp et al., 26 April
1979; USNM 376186, 2 males, 32 and 34 mm SL and 1
female, 41 mm SL, 08°51’24’’N, 123°24’36’’E, Solino
Island, Zamboanga, Mindanao, Philippines, 0–5 m, L. W.
Knapp et al., 3 May 1979.
Diagnosis. Vertebrae 11–12+30–32=41–43, dorsal in
rays 72–82, anal in rays 56–64; eyes moderately large
(2.1–2.5% SL); body slender, snout without cirri; scales
only on cheeks, no scales above opercular spine; upper
preopercular pore present; outer pseudoclasper moderately
broadbased, long, sometimes extending beyond hood in
resting position; inner pseudoclasper half as long as outer
pseudoclasper, anterior lobe very long, twice as long as
posterior lobe; otolith length to height 2.1–2.2, with shallow
Fig. 25. Alionematichthys suluensis sp. nov., A, lateral view of head, holotype; B, ventral view of head, holotype; C, view of left pseudoclaspers
from inside, holotype; D, inclined lateral view of male copulatory organ, holotype; E, median view of right otolith, AMS I.40149-028, female,
40 mm SL; F, ventral view of right otolith, AMS I.40149-028, female, 40 mm SL.
122
Dinematichthyine ishes of the Indo-west Paciic, Part IV
sixth the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 12–16 (14) rakers, 3 elongate in a
row. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 25A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about half the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): first
anterior pore large, tubular, with single cirrus anteriorly.
Preopercular pores: 3 lower, irst and second with joined
opening, covered by dermal lap in lateral view; third
tubular; upper preopercular pore present. [See description
of Alionematichthys ceylonensis for position of pores.]
Dentition (of holotype). Premaxilla with 4 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 3/4 diameter of pupil.
Vomer horseshoe-shaped, with 1 outer row of small teeth
and 1 inner row of larger teeth up to 1/3 diameter of pupil.
Palatine with an outer row of small teeth and an inner row
of larger teeth up to 1/3 diameter of pupil. Dentary with 3
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, merging into 1 row of larger teeth posteriorly.
Large dentary teeth up to about 3/4 diameter of pupil.
Otolith (Fig. 25E, F). Moderately elongate in shape,
length to height 2.1–2.2 (30–49 mm SL) and moderately
thick (otolith height to otolith thickness about 1.5). Anterior
tip slightly pointed, posterior tip pointed, expanded and
irregularly ornamented. Dorsal rim shallow, with notch
above anterior tip and behind marked postdorsal angle.
Inner face convex, outer face slightly concave to almost lat,
both smooth. Otolith length to sulcus length 1.7–1.8. Sulcus
slightly supramedially positioned and slightly inclined,
with separated colliculi and marked notch at ventral rim
of sulcus at joint of ostium with cauda. Ostium length to
cauda length 3.3–3.7. Ventral furrow very close to ventral
rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined
and 5–8 depressed. Parapophyses present from vertebrae
6 to 11 (12). Pleural ribs on vertebrae 2 to 11 (12). First
anal in pterygiophore markedly longer than subsequent
pterygiophore, reaching tip of last precaudal parapophysis
in males.
Male copulatory organ (Fig. 25C, D). Two pairs of
pseudoclaspers, outer sometimes extending beyond hood
in resting position. Outer pseudoclasper moderately broad
based with thick supporter bent backwards at tip; inner
pseudoclasper about half the length, thin, with very long
anterior inclined lobe about twice as long as small posterior
lobe. Isthmus narrow; penis curved, slightly shorter than
outer pseudoclaspers.
Colour. Live colour unknown. Preserved colour
uniformly brown.
Comparison. Alionematichthys suluensis belongs to
the species group with an upper preopercular pore, large
eyes and no scales above the opercular spine as A. piger
and A. samoaensis. It differs from both in the lack of cirri
on the snout (except for a single cirrus associated with the
Table 11. Meristic and morphometric characters of Alionematichthys
suluensis sp. nov.
Holotype
AMS I.
40149028
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
40
80
16
63
20
11
32
43
14
2
6
23
13
SL
24.4
12.2
14.5
5.7
12.3
2.2
1.2
6.3
4.0
16.9
46.0
?
15.7
12.1
4.4
25.6
28.6
Holotype +
56 paratypes
n
Mean (range)
36.7 (13-48)
57
77.5 (72-82)
15.8 (15-16)
60.2 (56-64)
20.2 (19-22)
11.5 (11-12)
30.7 (30-32)
42.1 (41-43)
14.5 (12-16)
2
6.1 (6-7)
22.3 (20-25)
13.6 (13-15)
28
21
28
12
28
28
28
11
11
28
28
28
25.2 (23.9-27.0)
11.9 (10.2-13.6)
15.1 (14.3-16.6)
5.5 (4.9-6.1)
13.0 (12.3-13.9)
2.2 (2.1-2.5)
1.4 (1.2-1.7)
6.6 (6.0-7.4)
4.2 (3.7-4.7)
18.0 (16.9-18.9)
46.5 (43.6-48.6)
31.0 (29.4-32.5)
15.8 (13.5-18.4)
12.9 (10.8-14.5)
5.3 (4.4-5.8)
24.0 (21.2-25.9)
28.4 (26.0-31.6)
11
11
11
11
11
11
11
11
11
11
11
10
11
11
11
11
11
dorsal rim and postdorsal angle, otolith length to sulcus
length 1.7–1.8, ostium length to cauda length 3.3–3.7.
Description (Figs 24, 25).The principal meristic and
morphometric characters are shown in Table 11. One of
the smallest species in the genus; mature at about 30 mm
SL, maximum size about 50 mm SL. Body slender, with
moderately pointed head proile. No cirri on snout. Eye
size 2.1–2.5 (2.2)% SL. Head with scale patch on cheek
containing 5–7 (5) vertical rows of scales on upper part
and 3 vertical rows on lower part, no scales on opercle.
Horizontal diameter of scales on body about 2.0% SL,
in 24 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded with little knob. Anterior
nostril positioned high, one-third distance from upper lip to
anterior margin of eye. Posterior nostril small, about one123
W. Schwarzhans and P. R. Møller
Fig. 26. Alionematichthys winterbottomi sp. nov., ROM 47597, holotype, male, 54 mm SL.
Fig. 27. Alionematichthys winterbottomi sp. nov., A, lateral view of head, ROM 47597, holotype, male, 54 mm SL; B, ventral view of head,
holotype; C, ventral view of male copulatory organ, ROM 82975, paratype, 58 mm SL; D, inclined lateral view of male copulatory organ, ROM
82975, paratype, 58 mm SL; E, view of left pseudoclaspers from inside, ROM 82975, paratype, 65 mm SL; F, view of left pseudoclaspers
from inside, holotype; G, median view of right otolith, CAS 227284, male, 76 mm SL.
124
Dinematichthyine ishes of the Indo-west Paciic, Part IV
irst anterior mandibular pore) and the very peculiar shape
of the inner pseudoclasper with its long anterior thorn.
Distribution. Alionematichthys suluensis appears to
be restricted in distribution to the Philippines between 13°
and 8° N, mainly in the Sulu Sea, where it co-occurs with
A. piger, A. plicatosurculus and A. riukiuensis (Fig. 16).
Biology. A 39 mm SL female (USNM 376179), contains
325 embryos, length 3.7 mm TL.
Etymology. Named after the type locality, the Sulu Sea
of the Philippines. It is an adjective.
Table 12. Meristic and morphometric characters of Alionematichthys
winterbottomi sp. nov.
Holotype +
n
Holotype 198 paratypes
ROM
47597
Mean (range)
Standard length in mm
80
48.9 (10-88) 199
Meristic characters
Dorsal in rays
81
82.5 (80-86)
24
Caudal in rays
16
16
24
Anal in rays
62
63.9 (61-65)
24
Pectoral in rays
22
21.3 (20-22)
11
Precaudal vertebrae
12
11.0 (11-12) 26
Caudal vertebrae
31
31.0 (29-32)
26
Total vertebrae
43
42.0 (40-43)
26
Rakers on anterior gill arch
17
18.3 (17-20)
11
Pseudobranchial ilaments
2
1.9 (1-2)
11
D/V
5
5.7 (5-6)
26
D/A
22
23.1 (21-24)
26
V/A
13
13
26
Morphometric characters in % of SL
Head length
27.2
27.3 (26.3-27.8) 11
Head width
15.9
15.5 (14.0-17.2) 11
Head height
16.4
16.9(15.9-18.3) 11
Snout length
5.2
6.0 (5.2-6.5)
11
Upper jaw length
14.4
14.4 (13.6-15.1) 11
Diameter of pigmented eye
2.2
2.5 (2.2-3.0)
11
Diameter of pupil
1.4
1.5 (1.2-1.7)
11
Interorbital width
6.8
7.1 (6.2-7.9)
11
Posterior maxilla height
5.0
4.9 (4.5-5.4)
11
Postorbital length
20.4
20.0 (19.5-20.6) 11
Preanal length
47.1
46.9 (44.7-48.6) 11
Predorsal length
30.7
31.5 (30.4-32.5) 11
Body depth at origin of anal in 19.7
18.2 (17.2-19.7) 11
Pectoral in length
15.3
14.6 (13.2-15.5) 11
Pectoral in base height
6.7
6.6 (6.3-7.2)
11
Ventral in length
22.8
23.0 (21.4-24.4) 11
Base ventral in - anal in origin 26.3
26.7 (23.4-29.5) 11
Alionematichthys winterbottomi sp. nov.
(Figs 16, 26, 27, Tables 1, 12)
Material examined. (217 specimens, 10–88 mm SL).
HOLOTYPE – ROM 47597, male, 80 mm SL, 18°45’52’’S,
178°31’13’’E, shallow reef off irst black rock 300 m S of
University of the South Paciic Research Station, Dravuni
Island, Fiji, coll. R. Winterbottom, A. R. Emery, F. Emery
and R. McKinnon, 20 March 1983. PARATYPES – CAS
222530, 2 males, 56 and 57 mm SL and 1 female, 42 mm
SL, Vatumatau Bay, Fiji, D. W. Greenield et al., 12 Nov.
2002; CAS 227291, 1 male, 65 mm SL and 1 female, 57
mm SL, Viti Levu north, Fiji, D. W. Greenield et al., 31
March 2002; CAS 222532, 1 male, 48 mm SL, 1 female,
49 mm SL and 1 juvenile, 29 mm SL, Fiji, D. W. Greenield
et al., 14 Nov. 2002; CAS 222540, 1 female, 52 mm SL,
Fiji, 3 April 2002; CAS 222544, 2 males, 42 and 48 mm
SL and 2 females, 45 and 55 mm SL, Fiji, D. W. Greenield
et al., 14 March 1982; CAS 222552, 4 males, 51–70 mm
SL, 2 females, 48–54 mm SL, and 2 juveniles, 22–23 mm
SL, Great Sea reef, Kia Island, Fiji, D. W. Greenield et al.
31 March 2002; CAS 222558, 1 male, 63 mm SL, and 1
juvenile, 35 mm SL, Fiji, D. W. Greenield et al. 15 nov.
2002; CAS 227284, 7 males, 54–75 mm SL and 5 females,
41–64, Fiji, reef NE of Yaqaga Island and NW of Ovatova
reef, D. W. Greenield et al. 25 March 2002; CAS 227307,
3 males, 42–57 mm SL, 3 females, 40–59 mm SL, Bua Bay,
Fiji, D. W. Greenield et al. 24 March 2002; CAS 222586,
2 males, 42–44 mm SL and 3 females, 43, 50 and 50 mm
SL, Nasau Bay, Vanua Levu, Fiji, D. W. Greenield et al.,
25 May 2003; CAS 222589, 1 female, 57 mm SL, Fiji, D.
W. Greenield et al. 25 May 2003; CAS 222590, 2 males,
51 and 65 mm SL, 1 juvenile, 25 mm SL; Fiji east shore, D.
W. Greenield et al. 15 March 2002; CAS 222592, 1 male,
46 mm SL, 2 females, 31 and 44 mm SL, Budd Reef, Fiji,
D. W. Greenield et al. 23 May 2003; CAS 227294, 1 male,
57 mm SL, Fiji, D. W. Greenield et al., 20 May 2003; CAS
227295, 4 males, 45–57 mm SL, 12 females, 29–58 mm
SL, Budd Reef, Fiji, D. W. Greenield et al., 22 May 2003;
CAS 227297, 1 male, 53 mm SL and 3 females, 35–52
mm SL, Fiji, D. W. Greenield et al., 23 May 2003; CAS
222605, 2 females, 44 and 73 mm SL, Fiji, D. W. Greenield
et al., 5 January 2003; CAS 227298, 1 female, 88 mm SL,
Fiji, D. W. Greenield et al., 28 May 1999; CAS 227299, 3
males, 55–75 mm SL and 1 female, 53 mm SL, Fiji, D. W.
Greenield et al., 29 January 2002; CAS 222608, 3 males,
38–44 mm SL, Fiji, D. W. Greenield et al., 7 February
2002; CAS 222613, 3 males, 43–60 mm SL, Barrier reef
of Suva, Fiji, D. W. Greenield et al. 27 Jan. 2002; CAS
222619, 1 male, 64 mm SL, Fiji, D. W. Greenield et al.,
31 May 1999; CAS 222621, male, 46 mm SL, female, 50
mm SL, Nukalau Island, off Suva, D. W. Greenield et al.,
28 Jan. 200?; ROM 82975, 26 males, 34–78 mm SL, 57
females, 22–75 mm SL, 16 juveniles, 10–34 mm SL, same
data as holotype; ROM 82977, 5 females 45–70 mm SL, 2
males, 60 and 77 mm SL, 18°46’30’’S, 178°30’28’’E, off
Yanu-Yanu-Sau Island, south of Dravuni, Fiji, A. R. Emery
and party, 30 Mar. 1983.
Additional material. USNM 377255, 15 specimens,
Vuro, Fiji; USNM 378376, 4 specimens, 27–50 mm SL,
Malolo, Fiji; USNM 257649, 1 male and 2 females,
32–58 mm SL, 17°11’S, 176°54’E, Yasawa Group, Viwa
Island, Fiji; USNM 263659, 2 males and 2 females,
125
W. Schwarzhans and P. R. Møller
30–72 mm SL, 18°52’S, 178°30’E, Kandavu Island, Fiji;
USNM 374211, 1 male, 65 mm SL, 21°20’S, 174°58’W,
Eua, Tonga; USNM 366566, 9 specimens, 36–77 mm
SL, 18°50’S, 178°32’E, Kandavu Island, Fiji; USNM
376196, 2 females, 39–48 mm SL, 18°56’S, 178°21’W,
Lau Group, Yagasa Island, Fiji; USNM 376209, 1 male
and 1 female, 45–49 mm SL, 17°06’S, 177°13’E, Yasawa
Group, Naviti Island, Fiji.
Diagnosis. Vertebrae 11(–12)+29–32=40–43, dorsal
in rays 80–86, anal in rays 61–65; eyes moderately large
(2.2–3.0% SL); body moderately slender, snout with many
cirri; scales on cheeks, patch of scales above opercular spine
with 5–9 scales; upper preopercular pore present; outer
pseudoclasper broad-based, not extending beyond hood in
resting position; inner pseudoclasper short, thin, free from
outer pseudoclasper, bifurcate, anterior pointed lobe short,
posterior lobe much expanded posteriorly; otolith length to
height 2.0–2.2, with shallow dorsal rim and weak postdorsal
angle, otolith length to sulcus length 1.6–1.7, ostium length
to cauda length 3.0–4.5.
Description (Figs 26, 27).The principal meristic and
morphometric characters are shown in Table 12. Mature at
about 45 mm SL. Body moderately slender, with pointed
head proile. Many cirri on snout. Eye size 2.2–3.0 (2.2)%
SL. Head with scale patch on cheek containing 8–10 (9)
vertical rows of scales on upper part and 4 vertical rows
on lower part, 5–9 (9) scales on opercle above opercular
spine. Horizontal diameter of scales on body in holotype
about 1.9% SL, in 30 horizontal rows. Maxillary ending far
behind eye, dorsal margin of maxillary covered by upper
lip dermal lobe, posterior end expanded. Anterior nostril
positioned high, 1/2.5 distance from upper lip to anterior
margin of eye. Posterior nostril small, about one-quarter the
size of eye. Opercular spine with free tip, pointed. Anterior
gill arch with 17–20 (17) rakers, Upper branch with one
knob-like raker and 3–5 (3) plate-shaped rakers and lower
branch with 13–15 (13) rakers, all plate-shaped except for
knob-shaped irst and third. Pseudobranchial ilaments 1–2
(usually 2).
Head sensory pores (Fig. 27A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): irst
anterior pore large, tubular. Preopercular pores: 3 lower,
irst and second with joined opening, covered by dermal
lap in lateral view; third tubular; upper preopercular pore
present. [See description of Alionematichthys ceylonensis
for position of pores.]
Dentition (of holotype). Premaxilla with 7 outer rows
of granular teeth and 2 inner rows of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/3 diameter of pupil.
Vomer horseshoe-shaped, with 3 outer rows of small teeth
and 1 inner row of large teeth up to 1/3 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of 8 large teeth up to 1/3 diameter of pupil. Dentary with 5
outer rows of granular teeth and 1 inner row of larger teeth
126
anteriorly, merging into 1 row of larger teeth posteriorly.
Thirteen large dentary teeth, up to about 2/3 diameter of
pupil.
Otolith (Fig. 27G). Moderately elongate in shape,
length to height 2.0–2.2 and moderately thin (otolith
height to otolith thickness about 2.2–2.8). Anterior tip
slightly pointed, posterior tip pointed, expanded and
irregularly ornamented. Dorsal rim shallow, with notch
above anterior tip and with indistinct postdorsal angle.
Inner face convex, outer face slightly concave to lat, both
smooth. Otolith length to sulcus length 1.6–1.7. Sulcus
slightly supramedially positioned and slightly inclined,
with separated colliculi and marked notch at ventral rim
of sulcus at joint of ostium with cauda. Ostium length
to cauda length 3.0–4.0 (4.5). Ventral furrow weak, very
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to
11 (12). Pleural ribs on vertebrae 2 to 11 (12). First anal in
pterygiophore slightly to markedly longer than subsequent
pterygiophore, but not reaching tip of last precaudal
parapophysis.
Male copulatory organ (Fig. 27C–F). Two pairs of
pseudoclaspers, outer not extending beyond hood in
resting position. Outer pseudoclasper broad based, nearly
triangular in shape, somewhat thickened towards tip; inner
pseudoclasper less half the length, thin, bifurcate, anterior
pointed lobe short, anteriorly free from outer pseudoclasper
halfway up from base, posterior lobe much expanded
posteriorly. Isthmus narrow; penis curved, slightly longer
than outer pseudoclaspers.
Colour. After preservation the specimens are uniformly
brownish.
Comparison. Alionematichthys winterbottomi belongs
to the species group with an upper preopercular pore, large
eyes, many cirri on the snout and within that group to the
subgroup with 5 or more scales above the opercular spine
together with A. riukiuensis, from which it differs in the
speciic shape of the thin, free and small inner pseudoclasper
with its bifurcate shape and the lack of scales below the
opercular spine. Both species are obviously is closely
related to each other.
Distribution. Alionematichthys winterbottomi is
restricted in distribution to the Fiji and Tonga Islands, where
it replaces A. riukiuensis (Fig. 16).
Etymology. Named in honour of Richard Winterbottom,
ROM, Toronto, Canada, in recognition of his many
contributions to ichthyology and for his great support of
the present revison. It is a noun in apposition.
Alionematichthys sp. 1
(Figs 16, 28, Table 1)
Material examined. (2 specimens, 75–87 mm SL).
AMS 19108-032, 1 male, 75 mm SL, 14°40’S, 145°28’E,
Lizard Island, Great Barrier Reef, Australia, 1–10 m, D.
F. Hoese and party, 17 Nov. 1975; WAM P.30305-036, 1
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Australia, Kendrew Island, Dampier Arch., G. R. Allen and
R. Steene, 3. Nov. 1974.
Remarks. A single female specimen from NW
Australia likely represents an undescribed species which
is characterised by the absence of cirri on the snout, the
absence of an upper preopercular pore (transformed to a
wart) and the presence of two scales above the opercular
spine. A similar combination of characters is found in
A. shinoharai, from which it differs in the more stout
and massive body and head and the longer sulcus on the
inner face of the otolith. The scale patch on the cheek is
remarkable for its uniform width with six vertical rows of
scales on the upper part and ive on the lower part. The
otolith is characterised by a poor distinction of ostium
and cauda and the small size of the cauda as found in
A. crassiceps. Geographically, this record is just outside
the distribution range of other species of Alionematichthys.
We have refrained from establishing a new species due
to the lack of a male specimen for investigation of the
pseudoclasper morphology.
female, 87 mm SL, 13°52’S, 126°56’E, Sir Graham Moore
Islands, Western Australia, depth 0–1 m, G. R. Allen, 15
Aug. 1991.
Remarks. Two specimens from northern Australia are
characterised by the presence of a uniform scale patch on
the opercle reaching from above the opercular spine to well
below it. The single male shows an inner pseudoclasper
resembling that of A. riukiuensis except for the anterior
rim of the inner pseudoclasper being free of a connection
to the outer pseudoclasper. All other characters, such as
cirri on the snout, head pores, otolith measures, meristics
and morphometrics perfectly it within the variation of
A. riukiuensis. More material needs to be studied before it
can be concluded if these specimens represent an extreme
variant of A. riukiuensis or a new species.
Alionematichthys sp. 2
(Figs 16, 29, Table 1)
Material examined. (1 specimen, 83 mm SL). WAM
P.25111-047, 1 female, 83 mm SL, 20°28’30’’S, 116°32’E,
Fig. 28. Alionematichthys sp.1, A, lateral view of head, AMS I.19108-032, male, 77 mm SL; B, ventral view of head, AMS I.19108-032,
male, 77 mm SL; C, view of left pseudoclaspers from inside, AMS I.19108-032, male, 77 mm SL; D, inclined lateral view of male copulatory
organ, AMS I.19108-032, male, 77 mm SL; E, median view of right otolith, WAM P.30305-036, female, 87 mm SL.
127
W. Schwarzhans and P. R. Møller
Fig. 29. Alionematichthys sp.2, WAM P.25111-047, female, 83 mm SL, A, lateral view of head, B, ventral view of head; C, median view of
right otolith.
Dinematichthys Bleeker, 1855
Type species: Dinematichthys iluoecoeteoides Bleeker,
1855 (type locality Kepulauan Batu, Sumatera Barat
Province, Indonesia). Gender masculine.
Dinematichthys Bleeker, 1855 in part. — Cohen and
Nielsen 1978; Cohen and Hutchins 1982; Machida 1994;
Nielsen et al. 1999.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed high on
snout at about 1/2–1/2.5 distance from lip to anterior rim
of eye; head completely and continuously covered with
scales on cheeks, opercle and occiput; cirri on snout; male
copulatory organ with two pairs of small pseudoclaspers;
outer pseudoclasper a simple lap, inner pseudoclasper a
lap of about half size of outer, with two or three lobes;
size up to about 110 mm SL; precaudal vertebrae usually
11 (10–12), total vertebrae 41–45, dorsal in rays 75–92,
anal in rays 59–71, V in D 2.1–2.4; upper preopercular pore
absent; sulcus of otolith with ostium at least two times as
long as cauda; maxilla expanded postero-ventrally.
128
Comparison. Dinematichthys is readily recognised
by its uniform and continuous head squamation including
cheeks, opercle and occiput, usually visible without removal
of mucus. The only other genus sharing this character is
Porocephalichthys gen. nov., which does not overlap in
distribution with Dinematichthys and which differs in
many other characters (see Porocephalichthys gen. nov.).
Even when in doubt, the combination of the absence of the
upper preopercular pore, the small and simple shaped inner
and outer pseudoclaspers, and the otolith with the sulcus
divided into ostium and cauda, usually allow a reliable
identiication.
Species. Dinematichthys contains two species, the
widely distributed D. iluocoeteoides known throughout
the Indo-west Paciic and D. trilobatus sp. nov., endemic
to the isolated Christmas and Cocos Islands of the Indian
Ocean.
Remarks. The holotype was originally part of Bleeker’s
collection, who described the irst ever dinematichthyine
in 1855, Dinematichthys iluocoeteoides, from a single
Dinematichthyine ishes of the Indo-west Paciic, Part IV
specimen from Batu Island (now Kepulauan Batu, Sumatra
Barat Province), off western Sumatra, Indonesia.
Bleeker’s holotype has been lost (see extensive
discussion in Cohen and Nielsen (1978)). A redeinition of
the genus Dinematichthys and the species D. iluocoeteoides
is therefore necessary.
Cohen and Nielsen (1978) discussed the nature of a
specimen from Bleeker’s original collection (assigned by
Günther (1862)) kept in London (BMNH 1868.2.28.65), but
concluded that it could not be the original type (though it
was later referred to as the holotype by Eschmeyer (1998)
(also Online Catalog of Fishes, version 19 Sept. 2008,
http://www.calacademy.org/research/ichthyology/catalog/
fishcatmain.asp). It is a male specimen and, although
dried and shrivelled, is here designated as neotype since it
resembles Bleeker’s original description in all important
aspects, and comes from the type locality and Bleeker’s
original collection.
Dinematichthys iluocoeteoides Bleeker, 1855
(Figs 30–32, Tables 1, 13)
Dinematichthys iluocoeteoides Bleeker, 1855: 319;
Günther 1862; Wourmes and Bayne 1973: 32; Cohen and
Nielsen 1978: 58; Dor 1984: 59; Winterbottom et al. 1989:
14; Nielsen et al. 1999: 130.
Dinematichthys indicus Machida, 1994: 451; Nielsen et
al. 1999: 130; Schwarzhans and Møller 2005: 78.
Dinematichthys randalli Machida, 1994: 456; Nielsen
et al. 1999: 130.
Material examined. (1207 specimens, 11–112 mm
SL). NEOTYPE – BMNH 1862.2.28.65, male, 61 mm
SL, Batu Island, Indonesia, purchased from P. Bleeker.
PARATyPES of D. indicus: ROM 37813-2, 3 males 45,
56 and 92 mm SL, 11 females, 49–77 mm SL, Chagos
Archipelago, Indian Ocean, 7°17’30”N, 7°23’56”E, depth
1–2 m, ROM 58269, 5 males, 48–81 mm SL, 3 females,
36–64 mm SL, 12°23’52’’S, 43°30’00’’E, Chissioua Dzaha,
Comoros, Indian Ocean.
Additional specimens. AMS I. 17491-006, 1 male, 56
mm SL, 1 female, 59 mm SL, 9°09’S, 159°48’E, Savo
Island, Solomon Islands, 0–10 m; AMS I. 17492-015, 2
females, 41–48 mm SL, 09°0’S, 160°06’E, Florida Island,
Solomon Islands; AMS I. 20775-092, 1 male, 69 mm SL,
Raine Island, Great Barrier Reef, Queensland, Australia;
AMS I. 22612-010, 1 male 56 mm SL and 2 females, 48
and 55 mm SL, 15°49’S, 145°50’E, Escape Reef, Great
Barrier Reef, Queensland, Australia; AMS I. 33693-008, 1
male, 66 mm SL, 11°42’45’’S, 144°04’E, Great Detached
Reef, Queensland, Australia, 6–19 m, AMS I. 33708-062,
1 female, 50 mm SL, 10°59’98’’S, 144°01’22’’E, Reef
10-418, Queensland, Australia, 2–9 m, AMS I. 37936-070,
1 female, 71 mm SL, 15°00’43’’S, 168°03’30’’E, Maewo
Island, Vanuatu; BMNH 2000.4.19.370-382, 12 specimens,
57–103 mm SL, Abu Dhabi, Persian Gulf; BPBM 8023, 1
male, 86 mm SL, Marshall Islands; BPBM 8072, 2 females,
49–65 mm SL, Palau; BPBM 40933, 2 females, 36 and 59
mm SL, Chuuk, Caroline Islands, 0–4 m; BPBM 10816, 1
female, 59 mm SL, Guadalcanal, Solomon Isls; BPBM
11358, 1 male, 107 mm SL and 1 female, 93 mm SL, Fiji;
BPBM 40935, 1 male, 47 mm SL, Ryukyu Islands, Japan;
BPBM 17704, 1 female, 53 mm SL, Caroline Islands;
BPBM 21795, 1 female, 72 mm SL, Mauritius; BPBM
40937, 1 female, 55 mm SL, Similan Island, Thailand;
BPBM 40939, 2 females, 32 and 47 mm SL, Seychelles;
BPBM 28591, 1 female, 73 mm SL, Philippines; BPBM
29149, 2 females, 55 and 67 mm SL, Marshall Islands;
BPBM 30491, 1 female, Flores, Indonesia; BPBM 30860,
1 male, 49 mm SL, Persian Gulf; BPBM 30861, 1 female,
54 mm SL, Persian Gulf; BPBM 35199, 1 female, 80 mm
SL, 27° 04’N, 142°12’30’’E, Chichi-jima, Ogasawara
Islands, Japan; BPBM 35276, 1 male, 74 mm SL, 1 juvenile,
Table 13. Meristic and morphometric characters of Dinematichthys
iluocoeteoides Bleeker, 1855.
Neotype +
n
578 non-types
Neotype
BMNH
1862.2.28.65
Mean (range)
61
55.7 (11-112) 579
129
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal
in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in
origin
84
16
69
22
11
31
42
15
6
22
13
SL
26.2
15.2
11.4
4.9
13.5
2.8
1.2
4.9
6.2
18.8
45.7
31.6
83.1 (75-92)
16.0 (15-16)
65.3 (59-71)
22.7 (21-24)
11.0 (10-12)
31.5 (30-34)
42.6 (41-45)
15.4 (13-19)
2
6.0 (5-7)
22.7 (19-26)
13.1 (12-14)
137
86
141
15
151
153
153
20
12
145
144
145
26.4 (25.8-27.1)
15.1 (12.2-20.9)
16.7 (11.4-18.9)
5.9 (4.9-6.5)
13.4 (12.7-13.8)
2.7 (2.2-3.3)
1.6 (1.2-2.0)
6.5 (4.9-7.5)
4.6 (4.1-6.2)
18.6 (17.8-19.2)
47.2 (44.3-50.6)
30.8 (29.2-31.8)
11
11
11
11
11
11
11
11
11
10
11
11
19.4
19.3 (16.3-21.4) 11
10.5
6.1
17.7
14.6 (10.5-16.2) 11
6.0 (5.2-6.9)
11
23.5 (17.7-25.6) 11
27.4
27.1 (24.1-37.0) 10
W. Schwarzhans and P. R. Møller
Fig. 30. Sample sites of Dinematichthys iluocoeteoides Bleeker, 1855, holotype of D. iluocoeteoides, holotype of D. indicus,
holotype of D. randalli, D. trilobatus sp. nov. and Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982). One symbol may
represent several samples.
Fig. 31. Dinematichthys iluocoeteoides Bleeker, 1855, A, fresh dead, BPBM 35594, female, 80 mm SL; B, neotype, BMNH 1862.2.28.65,
male, 61 mm SL.
21 mm SL, Ogasawara Islands, Japan; BPBM 35594, 1
female, 80 mm SL, Seychelles; BPBM 38149, 3 males, 66,
71 and 102 mm SL and 2 females, 56 and 65 mm SL, Tonga;
CAS 14255, 6 males, 41–83 mm SL, 6 females 50–102 mm
SL, 7°14’32”N, 144°27’12”E, Ifalik Atoll, Micronesia, 0–4
m; CAS 35340, 13 female, 34–72 mm SL, 10 males, 28–60
mm SL, Maldives; CAS 35341,19 males, 32–66 mm SL,
31 females, 29–66 mm SL, 1 juvenile, 23 mm SL,
130
Seychelles; CAS 227304, 1 male, 70 mm SL, Madang,
Papua New Guinea; CAS 65665, 4 males, 50–75 mm SL,
1 female, 55 mm SL, Madang, Papua New Guinea; CAS
65669, 1 male, 67 mm SL and 2 females, 53 and 63 mm
SL, Madang, Papua New Guinea; CAS 227289, 1 female,
76 mm SL, Madang, Papua New Guinea; CAS 227281, 1
male, 59 mm SL and 1 female, 64 mm SL, Kapingamarangi
Atoll, Micronesia; CAS 227290, 1 female, 46 mm SL, Viti
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 32. Dinematichthys iluocoeteoides Bleeker, 1855, A, lateral view of head, BPBM 40937, female, 55 mm SL; B, lateral view of head,
USNM 384592, male, 72 mm SL; C, dorsal view of head, USNM 384592, male, 72 mm SL; D, ventral view of head, USNM 384592, male
72 mm SL; E, lateral view of head, USNM 358342, male, 36 mm SL; F, view of left pseudoclaspers from inside, holotype; G, ventral view of
male copulatory organ, BPBM 35276, 74 mm SL; H, inclined lateral view of male copulatory organ, BPBM 35276, 74 mm SL; I, view of left
pseudoclaspers from inside, BPBM 35276, 74 mm SL; J, view of left pseudoclaspers from inside, USNM 319902, male, 71 mm SL; K, view
of left pseudoclaspers from inside, SMNS 22544, 52 mm SL; L, median view of right otolith, USNM 199680, male, 78 mm SL; M, ventral
view of right otolith, USNM 199680, male, 78 mm SL; N, median view of right otolith, WAM P.31213-007, female, 52 mm SL.
131
W. Schwarzhans and P. R. Møller
Levu north, Fiji; CAS 222534, 2 females, 58 and 63 mm
SL, Fiji; CAS 222535, 2 females, 24 and 48 mm SL, Fiji;
CAS 222539, 4 females, 55–68 mm SL, Fiji; CAS 222541,
female 70 mm SL, Fiji; CAS 227292, 1 female 65 mm SL,
Fiji; CAS 222546, 2 males, 41 and 64 mm SL, 1 female,
43 mm SL, Fiji; CAS 222548, 2 males, 52 and 58 mm SL,
4 females, 41–70 mm SL, 4 juveniles, 15–26 mm SL, Fiji;
CAS 222549, 1 male, 60 mm SL, 4 females, 43–65 mm
SL, Fiji; CAS 222557, 1 male, 74 mm SL, Fiji; CAS
222560, 1 male, 58 mm SL, 1 female, 55 mm SL, Fiji; CAS
222562, 2 males, 47 and 76 mm SL, 1 female with embryos,
50 mm SL, Fiji; CAS 222564, 1 male, 49 mm SL, Fiji; CAS
222568, 1 female, 45 mm SL, Fiji; CAS 227306, 1 male,
60 mm SL, Fiji; CAS 227287, 3 males, 38, 45 and 75 mm
SL, 11 female, 40–73 mm SL, 2 juveniles, 19 and 23 mm
SL, Fiji; CAS 222573, 2 females, 58 and 74 mm SL, Fiji;
CAS 222574, 1 female, 38 mm SL, Fiji ; CAS 222575, 1
male, 52 mm SL, 2 females, 41 and 73 mm SL, Fiji; CAS
222577, 3 females, 30–72 mm SL, 2 juveniles, 18 and 22
mm SL, Fiji; CAS 222578, male, 47 mm SL, Fiji; CAS
222579, 1 male, 53 mm SL, Fiji; CAS 222580, 1 female,
50 mm SL, Fiji; CAS 222581, 1 female, 52 mm SL, Fiji;
CAS 222587, 1 male, 78 mm SL, 1 female, 32 mm SL, Fiji;
CAS 222591, female, 86 mm SL, Fiji; CAS 222591, female,
86 mm SL, Fiji; CAS 227308, 2 females, 57 and 59 mm
SL, Fiji; CAS 222593, 2 males, 36–52 mm SL, 3 females,
32–51 mm SL, Fiji; CAS 222597, 8 females, 35–82 mm
SL, Fiji; CAS 222599, 4 males, 48–75 mm SL, 3 females,
65–75 mm SL, Fiji; CAS 222600, 2 males, 78 and 87 mm
SL, 1 female, 78 mm SL, Fiji; CAS 227296, 1 male, 48 mm
SL, Fiji; CAS 222602, 4 females, 61–80 mm SL, Fiji; CAS
222603, 1 male, 81 mm SL, 6 females, 65–90 mm SL, Fiji;
CAS 227309, 4 males, 51–83 mm SL, 2 females, 65–73
mm SL, Fiji; CAS 227300, 1 female, 82 mm SL, Fiji; CAS
222609, 2 females, 61 and 80 mm SL, 8 juveniles, 14–33
mm SL, Fiji; CAS 222611, 4 females, 35–72 mm SL, 2
males, 44–86 mm SL, 4 juveniles, 20–25 mm SL, Fiji; CAS
227310, 1 male, 53 mm SL, 1 female, 73 mm SL; CAS
222615, 2 females, 60 and 80 mm SL, Fiji; CAS 222617,
2 females, 49–75 mm SL, 2 males, 44 and 50 mm SL, Fiji;
CAS 222620, 2 females, 39 and 75 mm SL, Fiji; CAS
227302, 2 females, 65 and 75 mm SL, Fiji; CAS 222624,
1 male, 75 mm SL, Fiji; CAS 222627, 1 male, 81 mm SL,
Fiji; CAS 222634, 2 males, 59 and 85 mm SL, 1 juvenile,
19 mm SL; CAS 222639, 1 female, 60 mm SL, 1 male, 58
mm SL, Fiji; KAUM-I. 10660, 1 male, 42 mm SL,
Okinoerabu Island, Kagoshima, Japan; MNHN 1965-0431,
2 females, 54 and 70 mm SL, 23°20’0”S, 43°31’0”E,
Madagascar; MNHN 1977-0893, 1 female, 42 mm SL,
29°15’0”N, 34°45’0”E, Gulf of Aqaba, Israel; MNHN
1980-0243, 1 female, 90 mm SL, 22°26’0”S, 166°26’0”E,
New Caledonia; MNHN 1980-0563, 1 male, 112 mm SL,
17°30’0”S, 167°30’0”E, Pleiades Nord, Loyalty Islands,
15–17 m; EX MNHN 1980-0851, 1 male, 100 mm SL, 1
female, 66 mm SL, 17°30’0“S, 167°30’0”E, Ile Solitaire,
off Nouméa, New Caledonia; NTM S.13676-031, 2 females,
132
58 and 70 mm SL, 05°10’S, 145°50’3”E, Tab Island,
Madang, Papua New Guinea, 15–24 m; ROM 37811, 3
males, 50, 50 and 60 mm SL, 05°25’21”S, 071°46’52”E,
Isle du Coin, Chagos Archipelago; ROM 37812, 22 males,
30–81 mm SL, 18 females, 33–68 mm SL, 3 juveniles,
20–24 mm SL, 05°25’00”S, 071°46’00”E, Isle du Anglaise,
Chagos Archipelago; ROM 42314,1 female, 42 mm SL, 1
male, 48 mm SL, 09°28’00”S, 159°49’00”E, Honiara,
Solomon Islands; ROM 50299, female, 42 mm SL, male
40 mm SL, 09°28’00”S, 159°42’00”E, 10 km W of Honiara,
Solomon Islands; ROM 55144, 63 male, 09°12’16”S,
123°27’15”E, Tonga Point, Philippines; ROM 58267, 6
females, 32–53 mm SL, 5 males, 32–58 mm SL, 2 juveniles,
25 and 25 mm SL, 12°23’52”S, 43°30’00”E, Point
Chongochahari, Comoros; ROM 68122, 4 females, 28–65
mm SL, 3 juveniles, 12–22 mm SL, 12°23’52”S,
43°30’00”E, Chissioua Dzaha, Comoros; SAIAB 35108,
male, 63 mm SL, female, 54 mm SL, 22°00’N, 120°45’E,
Wanlitong, Taiwan; SAIAB 53381, 2 females, 27 and 41
mm SL, 16°18’N, 119°54’E, Luzon, Bolinao, Philippine
Islands; SMNS 13691, 2 females, 61 and 72 mm SL,
27°41’30’’N, 34°08’10’’E, Ras Mohammed, Red Sea,
0.5–4.5 m; SMNS 17126, 1 female, 28 mm, SL, 19°40’23’’S,
63°25’58’’E, Rodrigues Island, Mascarene Islands, 0.3–0.9
m; SMNS 17206, 4 females, 40–48 mm SL, 3 males, 39–87
mm SL, 19°40’25’’S, 63°25’59’’E, Rodrigues Island,
Mascarene Islands, 0–2 m; SMNS 18674, 2 males, 59–62
mm SL, 1 female, 80 mm SL, 08°22’06’’S, 116°04’46’’E,
Lombok, Indonesia, 1.5–3.8 m; SMNS 22544, 1 male, 54
mm SL, 29°09’44’’S, 34°41’34’’E, Gulf of Aqaba, Red
Sea, 0–3 m; TAU 11692, 2 females, 46 and 51 mm SL, 4
males, 42–56 mm SL, Seychelles; TAU 11808, 1 male, 70
mm SL, 1 female, 52 mm SL, Seychelles; TAU 11830, 3
specimens, Seychelles; UF 173089, 1 females, 81 mm SL,
1 male, 55 mm SL, Enewetak Atoll, Marshall Islands;
USNM 376199, 1 female, 53 mm SL, Buru Island,
Indonesia; USNM 99066, 1 female, 55 mm SL, Cagayanes,
Philippines; USNM 99223, 1 female, 34 mm SL, Ragay
Gulf, Philippines; USNM 99226, 1 male, 63 mm SL,
Dalaganem, Philippines; USNM 133865, 1 male, 80 mm
SL, Tahiti; USNM 142011, 2 females, 62–68 mm SL,
Rongelap Atoll, Marshall Islands; USNM 142016, 1 male,
82 mm SL, Bikini Atoll, Marshall Islands; USNM 374223,
1 male and 1 female, 46–85 mm SL, Bikini Atoll, Marshall
Islands; USNM 142019, 4 specimens, 89–108 mm SL,
Bikini Atoll, Marshall Islands; USNM 374224, 1 female,
91 mm SL, Bikini Atoll, Marshall Islands; USNM 166807,
2 males and 2 females, 60–82 mm SL, Arno Atoll, Marshall
Islands; USNM 376193, 3 females, 52–80 mm SL, Onotoa,
Gilbert Islands, Kiribati; USNM 199680, 5 females, 52–65
mm SL, 5 males, 40–83 mm SL, 12°10’S, 44°23’E,
Comoros; USNM 206304, 25 specimens, 34–83mm SL,
Saint Anne Island, Seychelles; USNM 201252, 1 male, 42
mm SL, Ambon, Maluku Islands, Indonesia; USNM
394973, 3 females, 31–71 mm SL, Tutuila, Samoa; USNM
223324, 2 specimens, 22–38 mm SL, Pohnpei, Micronesia;
Dinematichthyine ishes of the Indo-west Paciic, Part IV
USNM 223412, 6 specimens, 37–62 mm SL, Pohnpei,
Micronesia; USNM 223426, 1 female, 63 mm SL, Pohnpei,
Micronesia; USNM 223508, 14 specimens, 25–81 mm SL,
Pohnpei, Micronesia; USNM 223558, 3 specimens, 60–80
mm SL, Pohnpei, Micronesia; USNM 224330, 5 specimens,
28–75 mm SL, Pohnpei, Micronesia; USNM 224331, 5
specimens, 23–77 mm SL, Pohnpei, Micronesia; USNM
244012, 1 male, 77 mm SL, 1 female, 32 mm SL, 17°45’S,
177°04’W, Malolo Island, Fiji; USNM 244013, 1 male and
4 females, 24–63 mm SL, Malolo Island, Fiji; USNM
263660, 21 specimens, 20–75 mm SL, Lemus, Kavieng,
Papua New Guinea; USNM 263667, 5 specimens, 30–65
mm SL, Red Sea; USNM 263668, 1 female, 50 mm SL,
Marshall Islands; USNM 263673, 7 specimens, 40–86 mm
SL, 16°21’S, 43°59’E, Madagascar; USNM 377198, 15
specimens, 45–83 mm SL, 01°33’S, 144°59’E, Papua New
Guinea, 0–15 m; USNM 377209, 1 male and 1 female,
53–64 mm SL, 10°52’N, 120°56’E, Palawan, Philippines,
0–14 m; USNM 377191, 1 female, 58 mm SL, 08°51’N,
123°24’E, Zamboanga, Philippines; USNM 263692, 15
specimens, 51–80 mm SL, 09°10’N, 123°26’E, Negros,
Philippines, 0–3 m; USNM 377200, 3 males and 4 females,
38–82 mm SL, Weligama, Sri Lanka; USNM 263708, 1
specimen, 56 mm SL, Vuro Island, Fiji; USNM 263712,
27 specimens, 27–75 mm SL, 10°30’S, 44°21’E, Comores;
USNM 263750, 12 specimens, 22–71 mm SL, Kiriwina,
Papua New Guinea; USNM 263756, 18 specimens, 39–83
mm SL, Bougainville, Solomon Islands; USNM 267192,
4 specimens, 17–58 mm SL, Seychelles; USNM 394974,
4 males, 27–36 mm SL, 2 females, 42 and 43 mm SL, Mahe,
Seychelles; USNM 300092, 2 males and 1 female, 44–69
mm SL, 20°24’N, 121°55’E, Batanes, Philippines; USNM
300094, 1 female, 21°07’N, 121°56’E, Batanes, Philippines;
USNM 300100, 1 female, 20°17’N, 121°50’E, Batanes,
Philippines; USNM 300103, 1 male, 50 mm SL, 22°54’N,
121°54’E, Batanes, Philippines; USNM 374228, 1 male,
57 mm SL, 04°52’N, 119°26’E, Sibutu, Philippines; USNM
319898, 1 female, 78 mm SL, 20°34’S 166°14’E, Loyalty
Islands, New Caledonia, 0–5 m; USNM 319902, 2 males
50 and 71 mm SL, 5 females, 20°34’S, 166°14’E, Loyalty
Islands, New Caledonia; USNM 334123, 20 specimens,
26–89 mm SL, Tongatapu, Tonga; USNM 334125, 9
specimens, 28–75 mm SL, Tongatapu, Tonga; USNM
334126, 2 specimens, 23–93, mm SL, Tongatapu, Tonga;
USNM 334127, 13 specimens, Tongatapu, Tonga; USNM
374209, 11 specimens, 35–79 mm SL, 21°20’S, 174°58’W,
Tonga; USNM 336510, 9 specimens, 19°16’S, 174°22’W,
Tonga; USNM 338465, 4 females, 31–98 mm SL, Vava’u
Group, Tonga; USNM 394977, 6 specimens, 54–67 mm
SL, 18°44’31’’S, 174°06’36’’W, Vava’u Group, Tonga;
USNM 377199, 5 specimens, 42–75 mm SL, 10°35’N,
122°08’E, Panay, Philippines; USNM 376156, 1 female,
47 mm SL, 10°28’N, 122°28’E, Guimaraes, Philippines;
USNM 394983, 5 females, 43–72 mm SL, 1 male, 58 mm
SL, 16°47’13’’S, 168°21’36’’E, Epi, Vanuatu, 1–10 m;
USNM 352643, 6 specimens, 52–55 mm SL, Ryukyu
Islands; USNM 355820, 1 male and 1 female, 40–66 mm
SL, Papua New Guinea; USNM 357210, 1 juvenile, 23 mm
SL, 10°40’S 165°47’E, Solomon Islands; USNM 358342,
1 male, 36 mm SL, 10°16’S, 166°18’E, Solomon Islands;
USNM 384592, 1 female, 63 mm SL, 1 male, 72 mm SL,
Vanuatu, Efate; USNM 363675, 1 female, 79 mm SL,
Vanuatu; USNM 363908, 2 females, 60–88 mm SL,
15°00’S, 168°03’E, Maewo, Vanuatu; USNM 365322, 1
male and 1 female, 80–102 mm SL, 08°23’S, 162°51’E,
Stewart Island, Solomon Islands; USNM 365613, 4
specimens, 42–57 mm SL, 13°21’30’’S, 176°10’10’’W,
Wallis Island, Wallis and Futuna Islands, 1–13 m; USNM
377236, 2 males, 41–80 mm SL, Bugabag, Papua New
Guinea; USNM 365823, 4 specimens, 42–75 mm SL,
Hermit Island, Papua New Guinea; USNM 365826, 7
specimens, 58–66 mm SL, Hermit Island, Papua New
Guinea; USNM 365829, 1 female, 37 mm SL, Massas,
Papua New Guinea; USNM 366216, 6 specimens, 51–83
mm SL, Chagos Archipelago; USNM 366227, 3 females,
25–50 mm SL, 27°16’N, 33°47’E, Egypt; USNM 366228,
1 male, 45 mm SL, 26°08’N, 34°16’E, Egypt; USNM
366229, 1 female, 68 mm SL, 27°18’N, 33°47’E, Egypt;
USNM 366230, 1 male and 1 female, 40–60 mm SL,
Mauritius; USNM 366231, 2 juveniles, 17–26 mm SL,
16°25’S, 59°36’E, Cargados Carajos, Mascarene Islands;
USNM 366232, 3 females, 38–68 mm SL, 16°43’S,
59°35’E, Cargados Carajos, Mascarene Islands; USNM
366233, 7 specimens, 35–65 mm SL, 16°27’S, 59°36’E,
Cargados Carajos, Mascarene Islands; USNM 366234, 9
males and 24 females, 31–70 mm SL, 10°19’S, 56°35’E,
Agalega, Mauritius; USNM 366457, 2 specimens, Agalega,
Mauritius; USNM 366459, 8 males and 4 females, 32–86
mm SL, 07°15’S, 72°22’E, Chagos Archipelago; USNM
366460, 1 male, 54 mm SL, 07°15’S, 72°22’E, Chagos
Archipelago; USNM 366461, 5 males and 8 females, 42–84
mm SL, 07°15’S, 72°22’E, Chagos Archipelago; USNM
366462, 1 female, 45 mm SL, 13°49’S, 72°24’E, Chagos
Archipelago; USNM 366463, 2 females, 57–60 mm SL,
07°14’S, 72°23’E, Chagos Archipelago; USNM 366464, 3
males and 2 females, 39–69 mm SL, 07°20’S, 72°27’E,
Chagos Archipelago; USNM 366465, 5 males and 1 female,
40–59 mm SL, Marsa Mokrakh, Egypt; USNM 366466, 4
females, 27–41 mm SL, Egypt; USNM 366467, 1 female,
68 mm SL, Hikkaduwa, Sri Lanka; USNM 366468, 6
specimens, 40–82 mm SL, Hikkaduwa, Sri Lanka; USNM
366469, 1 male, 72 mm SL, Hikkaduwa, Sri Lanka; USNM
366472, 33 specimens, 22–92 mm SL, Egypt; USNM
376219, 6 males and 1 female, 21–65 mm SL, 05°52’S,
112°37’E, Java, Bawaean Island; USNM 366476, 47
specimens, 16°28’S, 59°40’E, Cargados Carajos, Mascarene
Islands; USNM 366477, 1 male, 71 mm SL, Hikkaduwa,
Sri Lanka; USNM 366478, 1 male and 1 female, 71–84
mm SL, Galle, Sri Lanka; USNM 366479, 1 female, 60
mm SL, Hikkaduwa, Sri Lanka; USNM 366483, 1 female,
58 mm SL, 07°14’S, 72°23’E, Chagos Archipelago; USNM
366484, 2 females, 42–57 mm SL, 07°20’S, 72°27’E,
133
W. Schwarzhans and P. R. Møller
Chagos Archipelago; USNM 366485, 2 males and 6
females, 44–67 mm SL, 07°13’S, 72°25’E, Chagos
Archipelago; USNM 366486, 1 female, 84 mm SL,
07°14’S, 72°23’E, Chagos Archipelago; USNM 366487, 1
male and 4 females, 44–69 mm SL, 07°16’S, 72°28’E,
Chagos Archipelago; USNM 366491, 1 female, 41 mm SL,
02°50’N, 95°56’E, Sumatra; USNM 366492, 3 males,
35–67 mm SL, 04°14’S, 152°10’E, Bismarck Archipelago,
Papua New Guinea; USNM 376220, 1 male, 65 mm SL,
03°47’S, 128°06’E, Maluku, Indonesia; USNM 376200, 1
female, 39 mm SL, 05°52’S, 110°25’E, Karimundjawa,
Java, Indonesia; USNM 366500, 1 male and 7 females,
39–75 mm SL, 05°51’S, 106°34’E, Pulau Seribu, Indonesia;
USNM 366501, 19 specimens, 16–48 mm SL, 01°33’S,
144°59’E, Papua New Guinea; USNM 376205, 5 specimens,
48–74 mm SL, Madang, Papua New Guinea; USNM
366504, 3 specimens, Madang, Papua New Guinea; USNM
366506, 1 male and 3 females, 68–87 mm SL, Briwadi
Island, Kiriwina Islands (Trobriand), Papua New Guinea;
USNM 376162, 31 specimens, 07°15’S, 72°22’E, Chagos
Archipelago; USNM 366513, 1 juvenile, 29 mm SL, Buroa,
Egypt; USNM 366514, 1 female, 61 mm SL, 15°32’N,
40°00’E, Ethiopia; USNM 366515, 14 specimens, 15–67
mm SL, Gulf of Aqaba, Egypt; USNM 366516, 1 female,
74 mm SL, Buroa, Egypt; USNM 366517, 2 males and 1
female, 43–71 mm SL, Sinai, Egypt; USNM 366519, 4
females, 41–58 mm SL, 16°28’S, 59°37’E, Cargados
Carajos, Mascarene Islands; USNM 366520, 15 specimens,
34–67 mm SL, North Island, Agalega, Mauritius; USNM
376201, 10 specimens, 34–69 mm SL, 05°10’S, 145°51’E,
Papua New Guinea; USNM 366523, 1 female, 61 mm SL,
Madang, Papua New Guinea; USNM 376191, 1 female, 45
mm SL, Madang, Papua New Guinea; USNM 366526, 13
specimens, 39–61 mm SL, Kiriwina Islands (Trobriand),
Papua New Guinea; USNM 366527, 11 specimens, 59–91
mm SL, 12°46’N, 44°59’E, Aden; USNM 366529, 3
specimens, 12°53’S, 45°16’E, Comores; USNM 366534,
4 specimens, 07°56’N, 81°34’E, Sri Lanka; USNM 366535,
21 specimens, 26–75 mm SL, 05°24’S, 53°13’E, Comores;
USNM 366551, 5 specimens, 27–47 mm SL, Israel; USNM
366552, 22 specimens, 11–98 mm SL, Gulf of Aqaba,
Egypt; USNM 366553, 13 specimens, 51–72 mm SL,
15°30’N, 39°54’E, Ethiopia; USNM 366571, 1 male, 54
mm SL, Guadalcanal, Solomon Islands; USNM 366573, 1
male, 73 mm SL, New Georgia, Solomon Islands, USNM
366590, 20 specimens, 30–80 mm SL, 16°36’S, 59°31’E,
Cargados Carajos, Mascarene Islands; USNM 366591, 31
specimesn, 28–75 mm SL, 16°26’S, 59°36’E, Cargados
Carajos, Mascarene Islands; USNM 366593, 30 specimens,
28–74 mm SL, 16°25’S, 59°36’E, Cargados Carajos,
Mascarene Islands; USNM 366594, 5 specimens, 35.81
mm SL, 16°15’S, 59°33’E, Cargados Carajos, Mascarene
Islands; USNM 374229, 1 female, 51 mm SL, 09°13’N,
123°28’E, Negros, Philippines; USNM 374225, 1 male, 66
mm SL, 09°31’N, 123°40’E, Philippines; USNM 366679,
1 female, 86 mm SL, Kwajalein Atoll, Marshall Islands;
134
USNM 366692, 2 females, 68–72 mm SL, 21°55’N,
120°50’E, Taiwan; USNM 366693, 2 specimens, 48–64
mm SL, 21°55’N, 120°48’E, Taiwan; USNM 366697, 20
specimens, 37–74 mm SL, Red Sea; USNM 366701, 1
specimen, 47 mm SL, 09°03’N, 123°07’E, Philippines;
USNM 366702, 4 specimens, 27–43 mm SL, Bararin,
Philippines; USNM 366704, 1 male, 49 mm SL, Tagauyan,
Philippines; USNM 366720, 1 male, 55 mm SL, Balabac,
Philippines; USNM 366721, 1 female, 44 mm SL, 11°42’S,
167°51’E, Vanikolo Islands, Santa Cruz Islands, Solomon
Islands,; USNM 366839, 7 specimens, 59–80 mm SL,
18°08’S, 178°24’E, Fiji; USNM 366840, 1 male, 78 mm
SL, 19°09’S, 179°45’E, Fiji; USNM 366842, 6 specimens,
45–77 mm SL, 18°58’S, 179°52’W, Fiji; USNM 366844,
4 specimens, 55–76 mm SL, 21°38’S, 178°45’W, Fiji;
USNM 366845, 6 males and 1 female, 44–81 mm SL,
18°55’S, 178°33’W, Fiji; USNM 366846, 6 specimens,
45–75 mm SL, 18°57’S, 178°17’E, Fiji; USNM 376206, 6
specimens, 40–111 mm SL, 17°06’S, 177°13’E, Fiji;
USNM 374208, 3 females, 32–60 mm SL, 13°23’S,
176°11’W, Wallis Island, Wallis and Futuna Islands; USNM
263698, 1 female, 57 mm SL, 01°33’S, 144°59’E, Papua
New Guinea; WAM P.27469-005, 5 males, 65–70 mm SL,
15°50’S, 145°50’E, Escape Reef, Queensland, Australia;
WAM P.27663-003, 2 males 65 and 65 mm SL, 1 female,
35 mm SL, NW Australia; WAM P.27825-031, 2 males, 35
and 42 mm SL, 2 females, 39 and 44 mm SL, Bismarck
Sea, Papua New Guinea; WAM P.28191-005, 1 male, 52
mm SL, Coral Sea; WAM P.29627-048, 2 females, Coral
Sea; WAM P.29642-010, 2 females, 42–53 mm SL, Coral
Sea; WAM P.30618-002, 1 male and 3 females, 67–79 mm
SL, Papua New Guinea; WAM P.30633-023, 1 male, 58
mm SL, Papua New Guinea; WAM P.30719-036, 1 female,
42 mm SL, Flores Sea, Indonesia; WAM P.30791-002, 1
female, 50 mm SL, New Caledonia; WAM P.30844-050, 3
specimens, Timor Sea; WAM P.31213-058, 1 female, 52
mm SL, Papua New Guinea; WAM P.31437-043, 1 male,
60 mm SL, Timor Sea; YCM-P 34028, 1 male, 54 mm SL,
Kakeroma Island, northern Ryukyu Islands, Japan; ZMUC
P 771654-56, 2 males, 66 and 85 mm SL and 1 female, 77
mm SL, Seychelles; ZMUC P 771657-58, 1 male, 61 mm
SL, and 1 female, 56 mm SL, Mauritius.
Diagnosis. Vertebrae 10–12+30–34=41–45, dorsal
in rays 75–92, anal in rays 59–71; eyes large (2.2–3.3%
SL); body moderately slender, snout with many cirri; head
continuously covered with scales, ventrally terminating
behind maxilla; upper preopercular pore absent; outer
pseudoclasper broad-based, small, not extending beyond
hood in resting position; inner pseudoclasper small, with
two lobes; otolith length to height 2.1–2.3 with regularly
curved dorsal rim, otolith length to sulcus length 1.6–1.7,
ostium length to cauda length 3.8–4.8.
Description (Figs 31, 32).The principal meristic and
morphometric characters are shown in Table 13. Mature at
about 45 mm SL. Body slender, with moderately pointed
head proile. Many cirri on snout. Eye size 2.2–3.3 (2.8)%
Dinematichthyine ishes of the Indo-west Paciic, Part IV
SL. Head continuously covered with scales on cheeks,
opercle and occiput, ventrally terminating behind maxilla.
Horizontal diameter of scales on body of 85 mm SL male
(ZMUC P771656) about 2.0% SL, in about 28 horizontal
rows. Maxillary ending far behind eye, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior
end expanded with little knob. Anterior nostril positioned
high, 1/2–1/2.5 distance from upper lip to anterior margin
of eye. Posterior nostril small, about one-ifth to one-sixth
the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 13–19 (15) rakers, 3 elongate in a
row. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 32A–E). Supraorbital pores
2–3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior):
irst anterior pore large, tubular, with cirrus anteriorly.
Preopercular pores: 3 lower, irst and second with joined
opening, covered by dermal lap in lateral view, third
tubular; no upper preopercular pore. [See description of
Alionematichthys ceylonensis for position of pores.]
Dentition (of holotype). Premaxilla with 7 outer rows of
granular teeth and 2 inner rows of larger teeth. Anteriormost
teeth in inner row up to 3/4 diameter of pupil. Vomer
horseshoe-shaped, with 3 outer rows of small teeth and 1
inner row of larger teeth up to 2/3 diameter of pupil. Palatine
with 2 outer rows of small teeth and 1 inner row of larger
teeth up to 1/2 diameter of pupil. Dentary with 5 outer rows
of granular teeth and 1 inner row of larger teeth, up to about
size of diameter of pupil.
Otolith (Fig. 32L–N). Elongate in shape, length to
height 2.1–2.3 and moderately thin (otolith height to otolith
thickness about 2.5). Anterior tip rounded, posterior tip
slightly pointed, expanded. Dorsal rim shallow, gently
curved. Inner face convex, outer face slightly concave to
lat, both smooth. Otolith length to sulcus length 1.6–1.7.
Sulcus medianly positioned, slightly inclined, with
separated colliculi and small notch at ventral rim of sulcus
at joint of ostium with cauda. Ostium length to cauda length
3.8–4.8. Ventral furrow weak, moderately close to ventral
rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6
to 11 (10–12). Pleural ribs on vertebrae 2 to 11 (10–12).
First anal in pterygiophore not longer than subsequent
pterygiophore.
Male copulatory organ (Fig. 32G–K). Two pairs of
small pseudoclaspers, outer not extending beyond hood in
resting position. Outer pseudoclasper broad based, simple
lap-shaped; inner pseudoclasper about half length of outer
pseudoclasper, with two lobes. Isthmus moderately narrow;
penis curved, slightly longer than outer pseudoclaspers.
Colour. Live colour reported as to yellow or bright to
red (Fig. 31A). Preserved colour mostly dark brown.
Variability and ontogeny. Despite the wide distribution
of D. iluocoeteoides there are few, if any, regional
variations to be observed. For instance, in life, colouration
of specimens from the Western Indian Ocean is usually
described as bright red while a record from the most northeastern location at the Japanese Bonin Islands was reported
as yellow (personal communication by J. E. Randall).
Another character showing some variation is the amount
of cirri on the snout, which appears to be denser in western
Indian Ocean specimens than in western Paciic specimens.
Specimens from furthest to the southwest (Vanuatu, Fiji,
Tonga) are special for their generally somewhat higher
meristic counts, both in vertebrae (43–45) as well as
dorsal in rays (84–92). Ontogenetic changes are most
strongly expressed in the squamation of the head, which
is continuous on cheeks, opercle and occiput in adults and
subadults from about 40 mm SL. Smaller juvenile and larval
specimens show some interruption of the scale coverage on
the head, particularly between cheeks and opercle, while
already complete between cheeks and occiput (Fig. 32E).
The density of cirri on the snout also increases with growth,
i.e. is absent in small specimens below 40 mm SL.
Remarks. Machida (1994) described D. indicus from
the Chagos Archipelago and Comores Islands, i.e. from
the central and western Indian Ocean and D. randalli from
Kosrae, Micronesia. He distinguished D. indicus from
D. randalli by the presence of cirri on the top and sides of
the head behind the eye (vs no cirri on head) and by lower
scale row counts (85–100 vs 107–113). In our review of
more than 1000 specimens from an almost continuous
geographical range, the main diagnostic character, i.e.
presence or absence of cirri on the head could not be
veriied. It is true that western Paciic specimens appear to
have generally fewer cirri, but we regard this character as
too weak and too irregularly distributed to be of diagnostic
value. Similarly, the scale row count is considered by us as
a not suficiently reliable character.
Distinction of these two species from D. iluocoeteoides
was based on comparative data from Bleeker (1855) and
Sedor and Cohen (1987), with D. iluocoeteoides and was
described as having larger eyes (a little over 5 in head
length / eye diameter vs 7.8–9.4) and having a sheathed
maxillary (vs unsheathed). Again, in our evaluation the eye
size differences are well within the variations observed,
while we failed to recognise a difference of sheathed versus
unsheathed maxillary in any of the specimens other than
possibly caused by preservation. In consequence, all three
nominal species are here regarded as representing a single
valid species for which D. iluocoeteoides is the senior
synonym.
Comparison. Dinematichthys iluocoeteoides is
closely related to D. trilobatus sp. nov. (see below for
differentiation of the two species). The head squamation
continuing on the cheeks, opercle and occiput, which
usually is easily visible without the removal of mucus,
distinguishes D. iluocoeteoides readily from any other cooccurring dinematichthyine species. Confusion may only
be possible in the case of juveniles, which show some gaps
135
W. Schwarzhans and P. R. Møller
in head squamation between the cheeks and the opercle
(Fig. 32E). Also, the absence of the upper preopercular
pore and connection of the cheek squamation with the
occiput squamation still allow a secure identiication in
most instances.
Distribution. Dinematichthys iluocoeteoides is the
most widely distributed species of all Dinematichthyini,
ranging from the western Indian Ocean (Red Sea, Arabian
Gulf, East Africa) to the Ryukyu and Ogasawara Islands
in the north-western Paciic and Marshall and Kiribati
Islands, Fiji, Tonga and Samoa in the central and southwestern Paciic (Fig. 30). The reason for its unusually
wide distribution is unknown, but due to the low degree of
regional variation (except for the sympatric D. trilobatus
sp. nov. of the Christmas and Cocos Islands; see below) is
likely to be of recent origin. Despite this wide distribution,
D. iluocoeteoides usually does not occur in large numbers
at most sampled locations, except for some areas in the
western and central Indian Ocean, e.g. the Comores or the
Chagos Archipelago. Dinematichthys iluocoeteoides is also
notable for its strict association with true reef environments
and particularly so with outer reefs and reef cores.
Dinematichthys trilobatus sp. nov.
(Figs 30, 33, 34, Tables 1, 14)
Material examined. (16 specimens, 37–58 mm SL).
HOLOTYPE – WAM P.29927-005, male, 46 mm SL,
12°05’S, 096°53’E, Direction Island, Cocos-Keeling
Islands, 0.1–2.0 m, G. R. Allen, 24 Feb. 1989. PARATYPES
– ANSP 162877, 49 mm SL, 12°06’30’’S, 096°49’35’’E;
Turk Reef, Cocos-Keeling Islands, 45–49 m, W.F. SmithVaniz and P. L. Colin, 8 March 1974; WAM P.26106-012,
male, 40 mm SL, 7 females, 45–57 mm SL, 10°29’S,
105°40’E, Rhonda Beach, Christmas Island, 10–12 m, G.
R. Allen and R. Steene, 2 June 1978; WAM P.29002-021,
males, 42, and 50 mm SL, female 58 mm SL, 10°25’S,
105°40’E, Kiritimati, 13–26 m, G. R. Allen and R. Steene,
28 June 1986; WAM P.29927-013), 3 females, 37, 37 and
38 mm SL, same data as the holotype.
Diagnosis. Vertebrae 11+31–32=42–43, dorsal fin
rays 80–87, anal in rays 60–67; eyes large (2.4–3.0%
SL); body slender, snout with few cirri; head continuously
covered with scales, ventrally extending beyond tip of
maxilla forward of third posterior mandibular pore; upper
preopercular pore absent; outer pseudoclasper broad-based,
small, not extending beyond hood in resting position; inner
pseudoclasper small, with three lobes; otolith length to
height 2.1–2.2, with regularly curved dorsal rim, otolith
length to sulcus length 1.7–1.8, ostium length to cauda
length 3.7–4.2.
Description (Figs 33, 34).The principal meristic and
morphometric characters are shown in Table 14. Mature at
about 40 mm SL. Body slender, with pointed head proile.
Few cirri on snout. Eye size 2.4–3.0 (2.8)% SL. Head
continuously covered with scales on cheeks, opercle and
occiput, ventrally extending beyond tip of maxilla forward
136
Table 14. Meristic and morphometric characters of Dinematichthys
trilobatus sp. nov.
Holotype
WAM
29927005
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
46
87
16
66
22
11
32
43
15
2
6
22
13
SL
26.5
15.0
17.0
5.3
13.4
2.8
1.7
6.2
4.5
18.8
49.1
30.1
16.9
14.6
5.9
23.5
27.2
Holotype +
15 paratypes
n
Mean (range)
47.4 (37-58)
16
82.6 (80-87)
15.9 (15-16)
64.1 (60-67)
22.4 (21-24)
11
31.7 (31-32)
42.7 (42-43)
15.4 (13-17)
2
6
22.0 (19-24)
13
15
8
15
14
15
15
15
14
13
14
14
14
27.1 (25.8-28.0)
13.7 (12.7-16.6)
16.5 (15.6-17.5)
5.9 (4.9-6.9)
14.0 (12.9-15.0)
2.7 (2.4-3.0)
1.8 (1.5-2.6)
6.9 (6.2-7.9)
4.5 (4.1-4.8)
19.0 (17.8-19.8)
45.8 (42.6-49.1)
31.1 (30.1-32.2)
17.6 (15.7-20.0)
14.8 (12.5-16.4)
6.3 (5.8-7.2)
23.6 (20.9-26.1)
25.7 (23.7-27.5)
13
13
13
13
13
13
13
13
13
13
13
13
13
13
13
11
13
of third posterior mandibular pore. Horizontal diameter
of scales on body about 1.3% SL, in 27 horizontal rows.
Maxillary ending far behind eye, dorsal margin of maxillary
covered by upper lip dermal lobe, posterior end expanded
with little knob. Anterior nostril positioned high, 1/2.5
distance from upper lip to anterior margin of eye. Posterior
nostril small, about 1/6 the size of eye. Opercular spine with
free tip, pointed. Anterior gill arch with 13–17 (15) rakers,
3 elongate in a row. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 34A–C). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): irst
anterior pore large, tubular. Preopercular pores: 3 lower,
irst and second with joined opening, covered by dermal
lap in lateral view; third tubular; no upper preopercular
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 33. Dinematichthys trilobatus sp. nov., WAM P.29927-005, holotype, male, 46, mm SL.
Fig. 34. Dinematichthys trilobatus sp. nov., A, lateral view of head, WAM P.29002-021, female, 58 mm SL; B, dorsal view of head, WAM
P.26106-012, female, 57 mm SL; C, ventral view of head, WAM P.26106-012, female, 57 mm SL; D, ventral view of male copulatory organ,
WAM P.29002-021, male, 42 mm SL; E, view of left pseudoclaspers from inside, holotype; F, inclined lateral view of male copulatory organ,
WAM P.29002-021, 42 mm SL; G, median view of right otolith, WAM P.29002-021, male, 42 mm SL; H, ventral view of right otolith, WAM
P.29002-021, male, 42 mm SL.
pore. [See description of Alionematichthys ceylonensis for
position of pores.]
Dentition (of holotype). Premaxilla with 6 outer rows of
granular teeth and 1 inner row of larger teeth. Anteriormost
teeth in inner row up to 1/3 diameter of pupil. Vomer
horseshoe-shaped, with 2 outer rows of small teeth and 1
inner row of larger teeth up to 1/4 diameter of pupil. Palatine
137
with 1 outer row of small teeth and 1 inner row of larger
teeth up to 1/4 diameter of pupil. Dentary with 3 outer rows
of granular teeth and 1 inner row of larger teeth, up to about
1/2 diameter of pupil.
Otolith (Fig. 34G, H). Elongate in shape, length to
height 2.1–2.2 and moderately thin (otolith height to otolith
thickness about 2.3). Anterior tip slightly pointed, posterior
W. Schwarzhans and P. R. Møller
tip pointed, expanded. Dorsal rim shallow, gently curved.
Inner face convex, outer face slightly concave to lat, both
smooth. Otolith length to sulcus length 1.7–1.8. Sulcus
medianly positioned, slightly inclined, with separated
colliculi and marked notch at ventral rim of sulcus at
joint of ostium with cauda. Ostium length to cauda length
3.7–4.2. Ventral furrow weak, moderately close to ventral
rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 depressed. Parapophyses present from vertebrae 6 to 11.
Pleural ribs on vertebrae 2 to 11. First anal in pterygiophore
not longer than subsequent pterygiophore.
Male copulatory organ (Fig. 34D–F). Two pairs of
small pseudoclaspers, the outer not extending beyond
hood in resting position. Outer pseudoclasper broad based,
simple lap-shaped; inner pseudoclasper about half the
length of outer pseudoclasper, with three lobes. Isthmus
moderately wide; penis curved, about as long as outer
pseudoclaspers.
Colour. Live colour unknown. Preserved colour light
brown.
Comparison. Dinematichthys trilobatus obviously is
closely related to D. iluocoeteoides, from which it differs
mainly in the shape of the inner pseudoclaper with three
distinct lobes (vs two), the more expanded head squamation
ventrally extending beyond tip of maxilla forward of third
posterior mandibular pore, and and only few cirri on the
snout (vs many cirri).
Distribution. Dinematichthys trilobatus is endemic to
the Christmas and Cocos-Keeling Islands, where it replaces
the widespread D. iluocoeteoides (Fig. 30). Dinematichthys
trilobatus represents the only dinematichthyine known from
the Christmas and Cocos Islands and its speciic separation
from the widespread D. iluocoeteoides indicates a prolonged
separation history for the fauna of these islands, or a perhaps
a colonisation event with subsequent speciation.
Etymology. Named for the inner pseudoclasper with
three lobes: trilobatus (Latin = three lobed). It is an
adjective.
Porocephalichthys gen. nov.
Type species: Dinematichthys dasyrhynchus Cohen
and Hutchins, 1982 (type locality Rottnest Island, Western
Australia). Gender masculine.
Dinematichthys (non Bleeker, 1855) in part. — Cohen
and Nielsen 1978; Cohen and Hutchins 1982; Machida
1994, Nielsen et al. 1999.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed high on
snout at about 1/2.5 the distance from lip to anterior rim
of eye; head completely and continuously covered with
scales on cheeks, opercle and occiput; abundant cirri on
snout; male copulatory organ with two pairs of equally
long pseudoclaspers; outer pseudoclasper a simple lap,
inner pseudoclasper inwardly curved stick-like; size up
to 100 mm SL; precaudal vertebrae 14, total vertebrae
138
47–49, dorsal in rays 96–104, caudal in rays 17–18,
anal in rays 62–69, V in D 2.3; upper preopercular pore
present; four additional pairs of supraorbital pores, one
pair above and in front of eyes, a second above and behind
eyes, two more pairs of pores on occiput in prolongation
of upper preopercular pore; one or two additional posterior
mandibular pores; jaws without canine teeth; no elongate
gill rakers; sulcus of otolith with cauda nearly of the length
of ostium.
Comparison. When originally described, the species
was tentatively placed in the genus Dinematichthys. Cohen
and Hutchins (1982, p. 342) wrote: “Differences between
the two are numerous and when more species are described
and specimens of the true D. iluocoeteoides are re-collected
and become available for study, it may be necessary to
establish the Rottnest Island species in a separate genus”.
After having reviewed all Dinematicthyine genera, we are
conident that D. dasyrhynchus should be placed in a separate
genus. Porocephalichthys resembles Dinematichthys in the
uniform and continuous head squamation including cheeks,
opercle and occiput and in having the anterior nostril placed
high on snout. The other characters of the diagnosis, such as
the many additional pores, total lack of elongate gill-rakers,
number of vertebrae, dorsal, caudal and pectoral in rays
(Table 1), and the proportions of the sulcus of the otolith
and the pseudoclasper pattern, place Porocephalichthys
well apart from Dinematichthys and any of the other
dinematichthyine genera. The correct phylogenetic position
of the genus needs further investigation.
Species. Porocephalichthys is a monospeciic genus
with P. dasyrhynchus endemic to SW Australia.
Etymology. Combined from porocephalus (Latin =
head with pores) and ichthys (Greek = ish), referring to the
many pores on the head. The gender is masculine.
Porocephalichthys dasyrhynchus (Cohen and
Hutchins, 1982)
(Figs 30, 35, 36, Tables 1, 15)
Dinematichthys iluocoeteoides. — Mees 1960: 18.
Dinematichthys sp. — Hutchins 1979: 93.
Dinematichthys dasyrhynchus Cohen and Hutchins,
1982: 342; Allen 1985: 2268, ig. 49; Paxton et al. 1989:
316; Nielsen et al. 1999: 130.
Material examined (9 specimens, 43–95 mm SL) (all
from Western Australia). Paratypes: AMS I. 20245-016,
1 male, 64 mm SL, 2 females, 77–94 mm SL, Horseshoe
Reef, Rottnest Island, 12–15 m, B. C. Russell and J. B.
Hutchins; Non-types: WAM P.27950-011, 2 males, 61
and 64 mm SL, 30°18’S, 115°00’E, Jurien Bay, 4–6 m, N.
Sinclair et al., 9 April 1983; WAM P.27951-007, 1 female,
95 mm SL, 30°18’S, 115°00’E, Jurien Bay, Osprey Inlet,
Western Australia, 2–5 m, J. B. Hutchins et al., 10 April
1983; ZMUC P77716-18, 3 females, 43, 46 and 89 mm SL,
32°01’S, 115°29’E, Green Island, Rottnest Island, Western
Australia, 10 April 1978.
Diagnosis. See generic diagnosis.
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Fig. 35. Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982), ZMUC P77726, female, 89 mm SL.
Fig. 36. Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982), A, lateral view of head, WAM P.27950-011, male, 64 mm SL; B, ventral
view of head, WAM P.27950-011, male, 64 mm SL; C, dorsal view of head, WAM P.27950-011, male, 64 mm SL; D, inclined lateral view
of male copulatory organ, WAM P.27950-011, male, 61 mm SL; E, view of left pseudoclaspers from inside, WAM P.27950-011, male, 64
mm SL; F, view of left pseudoclaspers from inside, WAM P.27950-011, male, 61 mm SL; G, median view of right otolith, ZMUC P77726,
female, 89 mm SL; H, anterior view of right otolith, ZMUC P77726, female, 89 mm SL; I, dorsal view of right otolith, ZMUC P77726,
female, 89 mm SL.
139
W. Schwarzhans and P. R. Møller
Table 15. Meristic and morphometric characters of Porocephalichthys
dasyrhynchus (Cohen and Hutchins, 1982).
Holotype +
n
Holotype 6 paratypes and
6 non-types
WAM
P.26614010*
Mean (range)
88
74.8 (43-102)
13
Standard length in mm
Meristic characters
Dorsal in rays
Caudal in rays
Anal in rays
Pectoral in rays
Precaudal vertebrae
Caudal vertebrae
Total vertebrae
Rakers on anterior gill arch
Pseudobranchial ilaments
D/V
D/A
V/A
Morphometric characters in % of
Head length
Head width
Head height
Snout length
Upper jaw length
Diameter of pigmented eye
Diameter of pupil
Interorbital width
Posterior maxilla height
Postorbital length
Preanal length
Predorsal length
Body depth at origin of anal in
Pectoral in length
Pectoral in base height
Ventral in length
Base ventral in - anal in origin
96
18
62
25
14
33
47
23
SL
26.1
6.3
13.6
2.8
5.6
4.9
56.8
28.4
20.5
12.5
22.7
-
* Data from Cohen and Hutchins (1982).
99.4 (96-104)
17.3 (17-18)
66.4 (62-69)
26.5 (25-28)
14
34.1 (33-35)
48.1 (47-49)
21.3 (19-26)
2
5
39.2 (37-43)
16.8 (16-17)
9
7
9
13
9
9
9
12
9
5
5
5
26.4 (24.6-28.1)
12.7 (11.1-15.9)
15.4 (14.2-16.6)
6.3 (5.5-7.2)
13.2 (12.5-13.9)
3.0 (2.6-3.4)
1.8 (1.7-2.1)
5.6 (5.1-6.0)
4.3 (3.2-5.4)
18.2 (17.6-18.6)
52.7 (48.8-56.8)
28.1 (25.6-30.4)
16.4 (13.7-20.5)
12.3 (11.7-13.5)
6.4 (6.0-7.7)
23.3 (18.6-28.1)
32.4 (29.5-36.3)
12
9
9
13
13
13
9
13
13
5
13
13
12
13
9
13
9
Description (Figs 35, 36).The principal meristic and
morphometric characters are shown in Table 15. Mature at
about 60 mm SL. Body slender, with pointed head proile.
Many cirri on snout. Eye size 2.6–3.4 (2.8)% SL. Head with
uniform and continuous squamation on cheeks, opercle and
occiput. Horizontal diameter of scales on body of 89 mm
SL female 1.3% SL, in about 33 horizontal rows. Maxillary
ending far behind eye, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded. Anterior
nostril positioned high, 1/2.5 distance from upper lip to
anterior margin of eye. Posterior nostril moderately small,
about one-quarter the size of eye. Opercular spine with free
tip, pointed. Anterior gill arch with 19–26 (23) short rakers,
none elongate. Pseudobranchial ilaments 2.
140
Head sensory pores (Fig. 36A–C). Supraorbital pores
7: 4 additional pairs of supraorbital pores as follows: 1
pair above and in front of eyes, a second above and behind
eyes, 2 more pairs on occiput in prolongation of upper
preopercular pore. Infraorbital pores 6 (3 anterior and 3
posterior), 3 posterior pores about half size of three anterior
pores. Mandibular pores 7–8 (3 anterior and 4–5 posterior):
irst anterior pore large, tubular; one or two additional
posterior mandibular pores oriented in separate, though
incomplete, row parallel to original one and closer to jaw.
Preopercular pores: 3 lower, irst and second with joined
opening, covered by dermal lap in lateral view, third nontubular; upper preopercular pore present. [See description
of Alionematichthys ceylonensis for position of pores.]
Dentition (of holotype). Premaxilla with 11 outer
rows of granular teeth and 1 inner row of larger teeth.
Anteriormost teeth in inner row up to 1/5 diameter of
pupil. Vomer triangular, with 10 outer rows of small teeth
and 1 inner row of larger teeth up to 1/6 diameter of pupil.
Palatine with 7 outer rows of small teeth and 1 inner row
of larger teeth up to 1/6 diameter of pupil. Dentary with 9
outer rows of granular teeth and 1 inner row of larger teeth,
up to about 1/4 diameter of pupil.
Otolith (Fig. 36G–I). Moderately elongate in shape,
length to height 2.2 and moderately thick (otolith height to
otolith thickness about 2). Anterior tip rounded, posterior tip
rounded, slightly expanded. Dorsal rim thick, shallow, with
slight and broad depression at its middle. Inner face convex,
outer face slightly concave to lat, both smooth. Sulcus very
long, almost opening anteriorly, otolith length to sulcus
length 1.2. Sulcus slightly supramedially positioned and
slightly inclined, with separated colliculi and marked notch
at ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 1.2–1.5. Ventral furrow distinct, close
to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–10 depressed. Parapophyses present from vertebrae
6 to 14. Pleural ribs on vertebrae 2 to 13. First anal in
pterygiophore much longer than subsequent pterygiophore,
reaching tip of last precaudal parapophysis.
Male copulatory organ (Fig. 36D–F). Two pairs of
pseudoclaspers, outer not extending beyond hood in resting
position. Outer pseudoclasper simple lap-shaped without
broadened base; inner pseudoclasper about length of outer
pseudoclasper or longer, reduced to stick-like inwardly
bent supporter. Isthmus moderately narrow; penis curved,
slightly longer than outer pseudoclaspers.
Colour. Live colour uniformly orange-brown (Allen
1985: 2268). Preserved colour uniformly light brown.
Comparison. See comparison between
Porocephalichthys and other genera above.
Distribution. Restricted in distribution to SW Australia,
around Perth and Rottnest Island (Fig. 30). The types were
collected in limestore reef habitats, at depths of 3–15 m.
Dinematichthyine ishes of the Indo-west Paciic, Part IV
DISCUSSION
Dinematichthys and Alionematichthys represent the
genera with the widest geographical distribution in the
tribe. Dinematichthys from the western Indian Ocean to
the western Paciic, absent only from the southeastern
Polynesian Islands; Alionematichthys is apparently not
found west of Sri Lanka, but reaches far east to Ducie Atoll,
Pitcairn Group, and constitutes the only dinematichthyine
genus which is represented in both the Indo-west Paciic
and the tropical western Atlantic by A. minyomma. Like
most other Dinematichthyini, the genera dealt with in this
last part of the review exhibit some restricted geographic
distribution patterns, but with fewer species than in the
similarly widely distributed genus Diancistrus. However,
the ishes dealt with in this review also contain some of
the most widespread species of the group, particularly
Dinematichthys iluocoeteoides (East Africa to Ogasawara)
and Alionematichthys piger (Andaman Islands to Ducie
Atoll). Alionematichthys riukiuensis is remarkable for its
large size. It can attain up to 150 mm SL, second in length
only to Dipulus caecus (up to 200 mm SL), but due to its
massive appearance it is certainly the foremost of all the
Dinematichthyini in body mass.
The fishes of the genera Dinematichthys and
Alionematichthys reviewed herein represent by far the
most common Dinematichthyini in the Indo-west Paciic,
comprising more than two-thirds of all investigated
specimens from the area. Both genera are particularly
common in reef core and outer reef environments, where they
often dominate the dinematichthyine samples, but are less
dominant in back reef / inshore and lagoonal environments.
They are entirely missing from locations outside of the
Indo-west Paciic reef belt (except for A. minyomma from
the Caribbean Sea). The tropical western Indian Ocean is
dominated by Dinematichthys iluocoeteoides; less so in the
eastern Indian Ocean and western Paciic, where various
species of the genus Alionematichthys dominate. In the
latter region, the genus Diancistrus (see Schwarzhans
and Møller (2005)) usually occurs associated with, or as
competitor of, Alionematichthys and Dinematichthys. Other
dinematichthyine genera seem to be more adapted to reefassociated niches in lagoon or near-shore environments (see
Schwarzhans and Møller (2007)).
141
W. Schwarzhans and P. R. Møller
REGIONAL CHECKLIST
The purpose of the following regional check list is
to facilitate a quick approach for cross-checking when
analyzing a collection of dinematichthyine ishes. The
regions are chosen to it best with the geographic distribution
patterns observed in ishes of this group. Species considered
endemic to a particular region are underlined. It has to be
mentioned though that certain regions are under-represented
due to limited sampling, such as the Andaman Sea, the
Gulf of Oman, the Banda and Celebes Seas of Indonesia,
the Solomon Islands and Madagascar. It is recommended
Red Sea and Gulf of Aden
Dinematichthys iluocoeteoides
Arabian Sea and Gulf of Oman
Dinematichthys iluocoeteoides
East Africa south of 5°N
Dinematichthys iluocoeteoides
Mascarenichthys sp.
Seychelles and Mascarenes
Dinematichthys iluocoeteoides
Mascarenichthys heemstrai
Madagascar
Dinematichthys iluocoeteoides
Majungaichthys simplex
South Africa south of 30°S
Dermatopsoides andersoni
Dermatopsoides kasougae
Dermatopsoides talboti
Mascarenichthys microphthalmus
Chagos Archipelago and Maldives
Diancistrus alleni
Dinematichthys iluocoeteoides
Sri Lanka
Alionematichthys ceylonensis
Dinematichthys iluocoeteoides
Andamans, Thailand, Malaya and
Sumatra
Alionematichthys phuketensis
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus sp.
Dinematichthys iluocoeteoides
Eusurculus andamanensis
Ungusurculus riauensis
Christmas and Cocos Islands
Dinematichthys trilobatus
that readers consult the check list of neighbouring areas
in these instances as well, since the lack of species is
probably artiicial. Other areas have remained virtually
unsampled for Dinematichthyini, such as the tropical East
African shores, the Spratly Islands, Timor or the Natuna
Islands (South China Sea). Such areas could harbour further
unknown endemic species. Finally, we provide a key to all
currently recognised genera of Dinematichthyini. To key
out a species, readers need to use all six individual papers
in the revision: Møller et al. 2004a, Møller et al. 2005,
Schwarzhans et al. 2005, Møller and Schwarzhans 2006,
Schwarzhans and Møller 2007, plis the present paper.
North Vietnam and Hainan
Island, China
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus vietnamensis
Kagoshima Islands, Japan
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus erythraeus
Diancistrus fuscus
Dinematichthys iluocoeteoides
Ryukyu Islands, Japan
Alionematichthys piger
Alionematichthys riukiuensis
Alionematichthys shinoharai
Diancistrus erythraeus
Diancistrus fuscus
Diancistrus jackrandalli
Diancistrus sp.
Dinematichthys iluocoeteoides
Taiwan and Philippines north of
15°N
Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys riukiuensis
Brotulinella taiwanensis
Diancistrus erythraeus
Diancistrus fuscus
Diancistrus machidai
Dinematichthys iluocoeteoides
Sulu Sea and Philippines south of
15°N
Alionematichthys piger
Alionematichthys plicatosurculus
Alionematichthys suluensis
Alionematichthys riukiuensis
Diancistrus erythraeus
Diancistrus fuscus
Diancistrus karinae
142
Diancistrus machidai
Diancistrus springeri
Dinematichthys iluocoeteoides
Paradiancistrus cuyoensis
Ungusurculus philippinensis
Ungusurculus williamsi
Celebes Sea
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus altidorsalis
Diancistrus beateae
Diancistrus karinae
Diancistrus machidai
Dinematichthys iluocoeteoides
Sunda Archipelago and
Indonesian Timor Sea
Beaglichthys bleekeri
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus beateae
Diancistrus machidai
Diancistrus novaeguineae
Diancistrus springeri
Dinematichthys iluocoeteoides
Paradiancistrus lombokensis
Ungusurculus komodoensis
Ungusurculus sundaensis
Banda Sea and NW New Guinea
Alionematichthys piger
Alionematichthys plicatosurculus
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus machidai
Diancistrus niger
Diancistrus novaeguineae
Dinematichthys iluocoeteoides
Ogasawara Islands, Japan
Dinematichthys iluocoeteoides
Dinematichthyine ishes of the Indo-west Paciic, Part IV
Belau, Guam, Caroline Islands
including Pohnpei
Alionematichthys crassiceps
Alionematichthys piger
Diancistrus atollorum
Diancistrus beateae
Diancistrus karinae
Diancistrus pohnpeiensis
Dinematichthys iluocoeteoides
Marshall and Gilbert Islands
Alionematichthys piger
Diancistrus atollorum
Diancistrus beateae
Diancistrus mennei
Dinematichthys iluocoeteoides
Bismarck Archipelago and NE
New Guinea
Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys plicatosurculus
Alionematichthys riukiuensis
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus atollorum
Diancistrus beateae
Diancistrus eremitus
Diancistrus karinae
Diancistrus mcgroutheri
Diancistrus novaeguineae
Dinematichthys iluocoeteoides
Eusurculus pristinus
Ungusurculus collettei
Solomon Islands
Alionematichthys piger
Alionematichthys plicatosurculus
Alionematichthys riukiuensis
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus beateae
Diancistrus eremitus
Diancistrus novaeguineae
Dinematichthys iluocoeteoides
Eusurculus pristinus
NW Australia W of 142°E & N of
25°S
Alionematichthys piger
Alionematichthys riukiuensis
Alionematichthys sp.
Beaglichthys bleekeri
Beaglichthys larsonae
Beaglichthys macrophthalmus
Brosmolus longicaudus
Diancistrus alleni
Diancistrus beateae
Diancistrus jeffjohnsoni
Diancistrus novaeguineae
Didymothallus criniceps
Didymothallus mizolepis
Dinematichthys iluocoeteoides
Dipulus hutchinsi
Eusurculus pistillum
Great Barrier Reef and SE New
Guinea
Alionematichthys piger
Alionematichthys riukiuensis
Alionematichthys sp.
Beaglichthys macrophthalmus
Diancistrus alleni
Diancistrus beateae
Diancistrus leisi
Diancistrus longiilis
Diancistrus mcgroutheri
Diancistrus novaeguineae
Didymothallus criniceps
Dinematichthys iluocoeteoides
Eusurculus pistillum
SW Australia W of 145°E and S of
25°S
Dactylosurculus gomoni
Dermatopsoides morrisonae
Dipulus caecus
Dipulus hutchinsi
Dipulus multiradiatus
Porocephalichthys dasyrhynchus
Zephyrichthys barryi
SE Australia E of 145°E and S of
25°S
Dermatopsis hoesei
Dermatopsis macrodon
Monothrix polylepis
Lord Howe Island
Alionematichthys piger
Diancistrus longiilis
New Zealand, North Island
Dermatopsis joergennielseni
Norfolk Island
Dipulus norfolkanus
New Caledonia
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus longiilis
Didymothallus pruvosti
Lapitaichthys frickei
143
Loyalty Islands and Vanuatu
Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus alleni
Diancistrus beateae
Diancistrus brevirostris
Diancistrus longiilis
Diancistrus novaeguineae
Diancistrus tongaensis
Dinematichthys iluocoeteoides
Eusurculus pristinus
Paradiancistrus acutirostris
Fiji
Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys winterbottomi
Dermatopsis greenieldi
Diancistrus beateae
Diancistrus eremitus
Diancistrus ijiensis
Diancistrus robustus
Diancistrus tongaensis
Dinematichthys iluocoeteoides
Tonga
Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys winterbottomi
Diancistrus alatus
Diancistrus manciporus
Diancistrus tongaensis
Dinematichthys iluocoeteoides
Samoa
Alionematichthys samoaensis
Diancistrus alleni
Diancistrus beateae
Diancistrus tongaensis
Dinematichthys iluocoeteoides
Cook, Tubuai and Society Islands,
Pitcairn (inc. Ducie Atoll)
Alionematichthys piger
Diancistrus katrineae
Diancistrus tongaensis
Diancistrus sp.
W. Schwarzhans and P. R. Møller
Key to the genera of the Dinematichthyini
1a. Head with continuous squamation on cheeks, opercle
and occiput .............................................................. 2
1b. Head with squamation patches on cheeks, and
occasionally also on opercle or without scales ....... 3
2a. Precaudal vertebrae 14; dorsal in rays >95; no canine
teeth; upper preopercular pore present; 3 pairs of pores
on occiput; sulcus of otolith with separate equally long
ostial and caudal colliculi ........................................
.....................................................Porocephalichthys
2b. Precaudal vertebrae 10–12; dorsal fin rays <95;
canine teeth present; upper preopercular pore absent;
no pores on occiput; sulcus of otolith with separate
colliculi, length ostial colliculum 2.5–4 times caudal
colliculum ........................................Dinematichthys
3a. Anterior nostril positioned high (less than 1/3 the
distance from upper lip to aggregate distance to
anterior margin of eye).................. Alionematichthys
3b. Anterior nostril positioned low (1/3.5 to 1/6 the
distance from upper lip to aggregate distance to
anterior margin of eye) ........................................... 4
4a. Termination of maxilla low, not expanded ............. 5
4b. Termination of maxilla expanded, angular or with
knob......................................................................... 8
5a. Opercular spine hidden ........................................... 6
5b. Opercular spine exposed ......................................... 7
6a. Single pair of pseudoclaspers with two equally long
supporters; upper preopercular pore present ...........
............................................................ Gunterichthys
6b. Single pair of pseudoclaspers with single supporter;
upper preopercular pore absent ......Dermatopsoides
7a. Precaudal vertebrae 11–14; dorsal in rays 64–85;
penis without hook near tip ................. Dermatopsis
7b. Precaudal vertebrae 13–25; dorsal in rays 86–191;
penis with hook near tip ............................... Dipulus
8a. A single pair of (outer) pseudoclaspers .................. 9
8b. Two or three pairs of pseudoclaspers.................... 13
9a. Two long supporters in pseudoclaspers ....................
Didymothallus
9b. Single supporter in pseudoclasper ........................ 10
10a. Precaudal vertebrae 11–12; sulcus of otolith with
separate colliculi .................................................. 11
10b. Precaudal vertebrae 13–15; sulcus of otolith with
undivided colliculi .............................................. 12
11a. Pseudoclasper simple lap with small hook at middle
of anterior rim … ............................... Lapitaichthys
144
11b. Pseudoclasper long and stick-like.............................
..................................................... Ogilbia mccoskeri
[note: only species of genus Ogilbia with single
pseudoclasper]
12a. Upper preopercular pore absent; dorsal fin rays
124–129; anal in rays 90–94; total vertebrae 55–59;
D/A 37–42 ...............................................Brosmolus
12b. Upper preopercular pore present; dorsal fin rays
93–104; anal in rays 64–76; total vertebrae 45–47;
D/A 28–35 ................................................ Monothrix
13a. 3 pairs of pseudoclaspers ...................................... 14
13b. 2 pairs of pseudoclaspers ...................................... 15
14a. Inner pseudoclasper separated into an anterior and a
posterior pseudoclasper, pseudoclaspers not joined at
base; 6–7 branchiostegal rays; 11 precaudal vertebrae;
pectoral fin rays 16–21; first and second lower
preopercular pore joined in a single opening ...........
.............................................................. Ogilbichthys
14b. Second inner pseudoclasper inserted between
irst inner pseudoclasper and outer pseudoclasper;
pseudoclaspers joined at base; 8–9 branchiostegal
rays; 13–14 precaudal vertebrae; pectoral in rays
22–26; first and second preopercular pores with
separate openings ........................... Dactylosurculus
15a. Upper preopercular pore absent ........................... 16
15b. Upper preopercular pore present ......................... 20
16a. No scales on head; specimens longer than 20 mm
SL without visible eyes, only minute black dots in
specimens less than 20 mm SL .............. Typhliasina
16b. Scales on cheeks; all specimens with visible eyes ...
............................................................................... 17
17a. First and second preopercular pores with separate
openings ................................................................ 18
17b. First and second lower preopercular pore joined in a
single opening ....................................................... 19
18a. Inner pseudoclasper anteriorly connected to outer
pseudoclasper; sulcus of otolith with undivided
colliculi ................................................ Beaglichthys
18b. Inner pseudoclasper branched, medially connected to
outer pseudoclasper; sulcus of otolith with separate
colliculi, cauda short ......................... Zephyrichthys
19a. Inner pseudoclasper concave, anteriorly connected to
outer pseudoclasper; precaudal vertebrae 11 ............
............................................ Diancistrus manciporus
[note: only species of genus Diancistrus without upper
preopercular pore]
19b. Inner pseudoclasper claw-like; precaudal vertebrae
12 ........................................................Ungusurculus
Dinematichthyine ishes of the Indo-west Paciic, Part IV
20a. Single lower preopercular pore ............................ 21
20b. Three lower preopercular pores, but irst and second
pore open into single opening ............................... 22
(SAM), J. Barry Hutchins (WAM), Tomio Iwamoto (CAS),
Susan L. Jewett (USNM), Jeffrey W. Johnson (QM), Helen
K. Larson (NTM), Jeff M. Leis (AMS), Yoshihiko Machida
(BSKU), James Maclaine (NHM former BMNH), Mizuki
Matsunuma (KAUM), John E. McCosker (CAS), Mark
A. McGrouther (AMS), Sue Morrison (WAM), Hiroyuki
Motomura (KAUM), Vusi Mthombeni (SAIAB), Jørgen G.
Nielsen (ZMUC), John R. Paxton (AMS), Patrice Pruvost
(MNHN), John E. Randall (BPBM), Sandra Raredon
(USNM), Sally Reader (AMS), Clive Roberts (NMNZ),
Robert H. Robins (UF), Mark Sabaj (ANSP), Jeff Seigel
(LACM), Gento Shinohara (NSMT), Shirleen Smith
(USNM), David G. Smith (USNM), William F. Smith-Vaniz
(USGS), Victor G. Springer (USNM), Andrew L. Stewart
(NMNZ), Arnold Suzumoto (BPBM), Tom Trnski (AMS),
Jeffrey T. Williams (USNM) and Richard Winterbottom
(ROM). Special thanks go to Ronald Fricke (SMNS) and
Helen K. Larson (NTM), for careful reviews of all four
papers on the Dinematichthyini of the Indo-west Paciic.
Also thanks are due to the following colleagues at
ZMUC: Brigitte Rubæk for making the specimen drawings,
Geert Brovad for producing the photos, and Tammes
Menne for help with x-raying and packing. The project was
inanced by the Carlsberg Foundation and by the Visiting
Collection Fellowship grant from the Australian Museum,
Sydney.
21a. Inner pseudoclasper about as large as outer
pseudoclasper and separate ..................Pseudogilbia
21b. Inner pseudoclasper half the size of outer pseudoclasper
and anteriorly connected to U-shaped structure .......
......................................................... Paradiancistrus
22b. Inner pseudoclasper anteriorly joined to outer
pseudoclasper ........................................................ 23
22a. Inner and outer pseudoclaspers free ..................... 24
23a. Body slender (head height ≤ 15% SL, depth at anal
≤ 16% SL); precaudal vertebrae predominantly 12
(rarely 11); maxilla rounded posterior-ventrally with
weak knob in front of rear corner; body scales small,
< 1.2% SL ............................................. Brotulinella
23b. Body robust, moderately slender to deep (head
height > 15% SL except for Diancistrus jeffjohnsoni,
body depth at anal > 16% SL except for Diancistrus
longifilis); precaudal vertebrae 11 (except 12 in
Diancistrus jeffjohnsoni); maxilla with angular
postero-ventral widening close to its termination;
body scales (1.2) 1.3–2.2% SL ............. Diancistrus
24a. Inner pseudoclasper hidden in pocket of isthmus when
in resting position; penis hooked ..............................
........................................................Mascarenichthys
24b. Inner pseudoclasper open; penis bent, but not
hooked ................................................................... 25
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ACKNOWLEDGMENTS
We wish to thank the following people for helping us
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