The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2007 23: 29–110
Review of the Dinematichthyini (Teleostei: Bythitidae) of the Indo-west Paciic.
Part III. Beaglichthys, Brosmolus, Monothrix and eight new genera with
description of 20 new species
Werner SchWarzhanS1 and Peter raSk Møller2
1
Ahrensburger Weg 103 D, 22359 Hamburg, GeRMANy
wwschwarz@aol.com
² Natural History Museum of Denmark, University of Copenhagen,
Universitetsparken 15, DK-2100 Copenhagen Ø, DeNMARK
pdrmoller@snm.ku.dk
abStract
An ongoing revision of the dinematichthyine ishes (Ophidiiformes, Bythitidae, Brosmophycinae) of the Indo-west
Paciic based on ca. 5000 specimens is published in several parts. Part III includes 957 identiied specimens in the
genera Beaglichthys Machida, 1993 (with one described and two new species), Brosmolus Machida, 1993 (with one
described species), Dactylosurculus (new genus with one new species), Didymothallus (new genus with one described
and two new species), eusurculus (new genus with three new species), Lapitaichthys (new genus with one new species),
Majungaichthys (new genus with one new species), Mascarenichthys (new genus with two new species), Monothrix
Ogilby, 1897 (with one described species), Ungusurculus (new genus with six new species) and Zephyrichthys (new
genus with one new species). In addition, a third new species is described of the genus Paradiancistrus Schwarzhans,
Møller and Nielsen, 2005, which had been established in part I of the review of the Dinematichthyini of the Indowest Paciic. The genera reviewed here are not necessarily closely related to each other, but are compiled in this
volume because they are obviously not related to the three other groups reviewed or under review – Diancistrus
and two related genera (see part I, Schwarzhans et al., 2005), the Dermatopsis – Dipulus group (see part II, Møller
and Schwarzhans, 2006) and Dinematichthys sensu latu (in preparation). These 11 genera are mainly distinguished
by pseudoclasper morphology, meristics, otolith morphology and presence or absence of the upper preopercular
pore. The main distinguishing characters of the species contained in these genera are vertebrae and in ray counts,
morphometric characters, head squamation and the morphology of otoliths and pseudoclaspers.
k eyWordS: viviparous brotulas, coral reef ishes, Indo-west Paciic, Australia, Philippines, Indonesia, Andaman Islands,
Mascarenes, Madagascar, South Africa, new species, new genera.
CONTeNTS
INTrODuCTION.....................................................................................................................30
MATerIAl AND MeThODS................................................................................................ 31
COMPArATIve MATerIAl ................................................................................................ 32
SySTeMATICS........................................................................................................................ 32
Tribe Dinematichthyini Cohen and Nielsen, 1978 .............................................................. 32
Key to the genera of the Dinematichthyini and the species reviewed ............................... 32
Beaglichthys Machida, 1993 ...............................................................................................34
Beaglichthys bleekeri sp. nov. .......................................................................................36
Beaglichthys larsonae sp. nov. ...................................................................................... 41
Beaglichthys macrophthalmus Machida, 1993 .............................................................44
Brosmolus Machida, 1993 ..................................................................................................46
Brosmolus longicaudus Machida, 1993 ........................................................................ 47
Dactylosurculus gen. nov. ..................................................................................................49
Dactylosurculus gomoni sp. nov. ..................................................................................49
Didymothallus gen. nov. ..................................................................................................... 52
Didymothallus criniceps sp. nov. .................................................................................. 53
Didymothallus mizolepis (Günther, 1867) .....................................................................56
Didymothallus pruvosti sp. nov. .................................................................................... 61
29
W. Schwarzhans and P. r. Møller
eusurculus gen. nov. ...........................................................................................................63
eusurculus andamanensis sp. nov. ...............................................................................63
eusurculus pistillum sp. nov. ........................................................................................66
eusurculus pristinus sp. nov. ........................................................................................70
Lapitaichthys gen. nov. .......................................................................................................72
Lapitaichthys frickei sp. nov. ........................................................................................73
Majungaichthys gen. nov. ................................................................................................... 75
Majungaichthys simplex sp. nov. .................................................................................. 75
Mascarenichthys gen. nov. .................................................................................................78
Mascarenichthys heemstrai sp. nov. .............................................................................78
Mascarenichthys microphthalmus sp. nov. ................................................................... 81
Mascarenichthys sp. ......................................................................................................83
Monothrix Ogilby, 1897 ......................................................................................................84
Monothrix polylepis Ogilby, 1897 .................................................................................84
Ungusurculus gen. nov. ......................................................................................................86
Ungusurculus collettei sp. nov. .....................................................................................86
Ungusurculus komodoensis sp. nov. .............................................................................89
Ungusurculus philippinensis sp. nov. ...........................................................................90
Ungusurculus riauensis sp. nov. ...................................................................................93
Ungusurculus sundaensis sp. nov. ................................................................................95
Ungusurculus williamsi sp. nov. ...................................................................................97
Zephyrichthys gen. nov. ..................................................................................................... 100
Zephyrichthys barryi sp. nov. ...................................................................................... 100
GeOGrAPhIC DISTrIBuTION ................................................................................ 103
ADDeNDuM to Part I: Paradiancistrus lombokensis sp. nov. ................................... 105
ADDeNDuM to Part II: Dermatopsis greenieldi Møller and Schwarzhans, 2006 ... 108
ACKNOWleDGMeNTS ............................................................................................ .109
reFerenceS.............................................................................................................. 109
were originally interpreted as members of the tribe
Brosmophycini: Brosmolus owing to the lack of ossiied
parts in the male intromittant organ, Beaglichthys
tentatively since it was known from a single female.
Machida’s judgement was based on Cohen and Nielsen
(1978) who used ossiication of the male copulatory organ
(pseudoclaspers) as the main diagnostic character for the
bythitid tribe Dinematichthyini. Møller et al. (2004a)
discussed the deinition of the tribe at length and followed
Sedor’s (1985) proposal to deine the Dinematichthyini by
the apomorphic position of the copulatory organ below a
covering leshy hood in a cavity of the ventral body wall.
In Schwarzhans et al. (2005) both genera were already
mentioned as part of a forthcoming part of the ongoing
review of the Dinematichthyini. This reassignment was
owed to the fact that males of Beaglichthys macrophthalmus
have now become available (see later) and that in both
instances (Beaglichthys and Brosmolus) the position of the
male copulatory organ fulils the diagnostic requirement
of the tribe Dinematichthyini. In fact it appears that
both genera are related to each other judging from their
unusual elongate shape and the high number of vertebrae.
A similarly high number of precaudal vertebrae are found
in Monothrix and Dactylosurculus gen. nov.
INTrODuCTION
This review of the dinematichthyine ishes, a tribe within
the subfamily Brosmophycinae of the family Bythitidae
(viviparous brotulas), is the ifth part since its inception,
dealing irst with the American Dinematichthyini (Møller
et al. 2004a, 2005) and then with Dinematichthyini from
the Indo-west Paciic (Schwarzhans et al. 2005; Møller
and Schwarzhans 2006). The ongoing review of the Indowest Paciic Dinematichthyini will result in one more
publication before the world-wide review of the group is
completed.
The fishes described in this volume represent a
combination of dinematichthyine genera, many of them
new and containing mostly few species, that are not closely
related to any of the larger groups of Indo-west Paciic
genera that have been revised by us nor the outstanding
one within the genus Dinematichthys sensu lato.
Three genera dealt with in this review have been
described previously, Beaglichthys Machida, 1993,
Brosmolus Machida, 1993 and Monothrix Ogilby, 1897,
all of them by monospecific designation. Monothrix
was described 110 years ago from the Sydney suburb of
Maroubra, and has received almost no attention since
then. The two genera that Machida described in 1993
30
Dinematichthyine ishes of the Indo-west Paciic III
Another pair of probably related genera is eusurculus
gen. nov. and Ungusurculus gen. nov., which are both
characterised by a specialised claw or sucker-disk-like
development of the inner pseudoclasper. Didymothallus
gen. nov. is characterised by the presence of a single (outer)
pseudoclasper with two almost equally long supporters, a
character that it shares only with Gunterichthys Dawson,
1966, from American waters. Majungaichthys n. gen.
and Mascarenichthys gen. nov., both from the western
Indian Ocean, come closest in features of all Indo-west
Paciic Dinematichthyini to the most abundant New World
genus Ogilbia Jordan and evermann, 1898. The two
remaining monospeciic genera, Lapitaichthys gen. nov.
and Zephyrichthys gen. nov., are set apart by an unusual
combination of characters, but do not possess a single
apomorphic character.
Dinematichthyine ishes live in shallow tropical to
subtropical waters, hidden in holes and crevices of coral
reefs, algae beds and rocky shores. The species of the
genera reviewed here generally occur in reef environments
of the western Indian Ocean or the West Paciic as well as
along the tropical shores of the Philippines to Australia,
but there are also three genera included that are typical
of the subtropical rocky shores of Australia, namely
Dactylosurculus gen. nov., Monothrix and Zephyrichthys
gen. nov.. Different reef and non-reef environments can be
dominated by different groups of the Dinematichthyini.
For instance, the species of Mascarenichthys gen. nov. are
typically found in back reef lagoonal environments. Also
Beaglichthys, Lapitaichthys gen. nov. and Ungusurculus
gen. nov. seem to be more typical for inshore environments.
More typical reef dwellers are Didymothallus gen. nov. and
eusurculus gen. nov. along with the two most common
dinematichthyine genera of the Indo-west Pacific,
Diancistrus Ogilby, 1899 and Dinematichthys Bleeker,
1855.
The geographical distribution of most dinematichthyine
species is very restricted. The most extreme forms of the
Dinematichthyini reviewed here in terms of geographic
restriction are Lapitaichthys frickei gen. nov. sp. nov.
from the reef and back reef environment of southern New
Caledonia around Nouméa and a few species of the genus
Ungusurculus gen. nov. These narrow distribution ranges
might be related to the exceptionally low fecundity. In the
case of Didymothallus gen. nov., which is also known from
rather restricted geographic distribution patterns along
Australian and New Caledonian shores, exceptionally low
numbers (one to three, usually two) of (large) embryos have
been found in gravid females. This genus is also noted for
containing a species (Didymothallus criniceps sp. nov.)
with gravid females bearing eggs in the ovary rather
than embryos. Subject to further veriication through live
observations, this species could indeed represent the irst
case of oviparity in the Bythitoidei, which has been deined
on the basis of viviparity and associated anatomical
features by Cohen and Nielsen (1978).
The species reviewed here show a moderate degree
of variation in their general appearance, morphometric
measurements and meristic counts, but do show the usual
high degree of variance in pseudoclasper morphology.
Also, head pores were found to be useful at the generic
level and otolith morphology at generic and specific
levels. Many species of the various genera are notable
for their moderate body size, which rarely exceeds 50
mm Sl and mature from about 30 to 40 mm Sl. Only
species of Beaglichthys and Dactylosurculus gen. nov.
regularly exceed 60 mm Sl in size and almost reach 100
mm in the case of Beaglichthys macrophthalmus. Their
live colouration, where known, tends to be uniform, the
prevailing colours being red, yellow, orange and violet,
but in one case the vertical ins are known to be bright,
though translucent, dark yellow in colour while the body
is violet-red (Beaglichthys macrophthalmus).
MATerIAl AND MeThODS
examination of ca. 5000 specimens of Indo-west
Paciic Dinematichthyini yielded 957 specimens which
could be identiied to the genera treated herein. Also
included are additional specimens identified 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 (Bernice P.
Bishop Museum, honolulu), CAS (California Academy
of Sciences, San Francisco), MNhN (Museum Nationale
d’histoire Naturelle, Paris), NhMG (Natural history
Museum of Göteborg, Sweden), NMNZ (Museum of
New Zealand Te Papa Tongarewa, Wellington), NMv
(National Museum of victoria, Melbourne), NSMT
(National Science Museum, Tokyo), NTM (Museum
and Art Gallery of the Northern Territory, Darwin), QM
(Queensland Museum, Brisbane), rOM (royal Ontario
Museum, Toronto), SAIAB (South African Institute for
Aquatic Biodiversity, formerly ruSI (JlB Smith Institute
of Ichthyology), Grahamstown), SMNS (Staatliches
Museum für Naturkunde, Stuttgart), uSNM (National
Museum of Natural history, Smithsonian Institution,
Washington), WAM (Western Australian Museum, Perth),
yCM (yokosuka City Museum), and ZMuC (Zoological
Museum, university of Copenhagen).
The methodology used in analysing dinematichthyine
ishes follows Møller et al. (2004a) and Schwarzhans et
al. (2005). Morphometric characters are given as percent
of standard length (Sl) throughout. In the descriptions
holotype values are given irst, followed by the range in
paratypes in brackets. Size of eye is measured as horizontal
diameter of pigmented eyeball. Meristic counts were made
31
W. Schwarzhans and P. r. Møller
from radiographs, except pectoral in rays, gill rakers, teeth
and scale rows. Abbreviations used in meristic counts
are: D/v = anterior dorsal in ray above vertebra number;
D/A = anterior anal in ray below dorsal in ray number;
v/A = anterior anal in ray below vertebrae number; DA = number of dorsal in rays minus number of anal in
rays; v in D = number of dorsal in rays per ray-bearing
vertebra.
Otoliths were removed through the gill cavity by
making a small cut above the gills on the right side. Size
of body scales was measured on holotypes at mid-body
above anal in origin.
Pseudoclaspers were observed by bending forward the
leshy hood covering the copulatory organ and thereafter
by bending outwards the pseudoclaspers or spreading
them and ixing them with a thin needle. In drawings, the
pseudoclaspers and penis are shaded; other parts, such
as the leshy hood, isthmus or outline of the copulatory
cavity are simple line drawings.
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 on behaviour, live colouration and feeding.
A number of females were examined for reproductive
data, for example, number and size of embryos.
The distribution maps were created using the
Microsoft encarta 2001 digital world atlas.
Key to the genera of the Dinematichthyini and the
species reviewed herein
1a. Anterior nostril positioned high (less than 1/3 the
distance from upper lip to aggregate distance to
anterior margin of eye) ...........................................
..............Dinematichthys s.l. (revision outstanding)
1b. Anterior nostril positioned low (1/3.5 to 1/6 the
distance from upper lip to aggregate distance to
anterior margin of eye) .......................................... 2
2a. Termination of maxilla low, not expanded ........... 3
2b. Termination of maxilla expanded, angular or with
knob ....................................................................... 6
3a. Opercular spine hidden ........................................ 4
3b. Opercular spine exposed ....................................... 5
4a. Single pair of pseudoclaspers with two equally long
supporters; upper preopercular pore present ..........
.......................................................... Gunterichthys
4b. Single pair of pseudoclaspers with single supporter;
upper preopercular pore absent .....Dermatopsoides
5a. Precaudal vertebrae 11–14; dorsal in rays 64–85;
penis without hook near tip ............... Dermatopsis
5b. Precaudal vertebrae 13–25; dorsal in rays 86–191;
penis with hook near tip ............................. Dipulus
6a. A single pair of (outer) pseudoclaspers ................. 7
6b. Two or three pairs of pseudoclaspers .................. 13
7a.
COMPArATIve MATerIAl
Two long supporters in pseudoclaspers ..................
.....................................(Didymothallus gen. nov.) 8
7b. Single supporter in pseudoclasper ...................... 10
Indo-west Paciic Dinematichthyini: see Schwarzhans
et al. (2005) and Møller and Schwarzhans (2006).
American Dinematichthyini: see Møller et al. (2004a)
and Møller et al. (2005).
Brosmophycinae and Bythitinae: see Møller et al.
(2004b).
8a. Anterior supporter slightly shorter than posterior
supporter; dorsal in rays 78–97; v in D 2.2–2.6;
otolith length to sulcus length 1.9–2.1 ....................
......................................... Didymothallus mizolepis
8b. Both supporters about equally long; dorsal in rays
69–77; v in D 1.9–2.1; otolith length to sulcus length
2.2–2.6 ................................................................... 9
SySTeMATICS
9a. Posterior supporter slender; D/A 23–26; otolith length
to sulcus length 2.2–2.4; cirri on occiput ................
............................ Didymothallus criniceps sp. nov.
9b. Posterior supporter with club-like expanded tip; D/A
20–22; otolith length to sulcus length 2.6; no cirri on
occiput ..................Didymothallus pruvosti sp. nov.
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 unreduced, 6 mandibular, 2–4 preopercular,
5–7 infraorbital and 3–4 supraorbital pores, including
supraorbital pore above opercular spine. Posteriormost
supraorbital head-pore tubular.
Tables 1 to 2 summarise meristic and selected other
morphological key characters used in distinguishing the
species of the genera described here.
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 frickei gen. nov. sp. nov.
11b. Pseudoclasper long and stick-like ...........................
................................................... Ogilbia mccoskeri
[note: this is the only species of genus Ogilbia with
single pseudoclasper, see also 36a]
32
Dinematichthyine ishes of the Indo-west Paciic III
12a. upper preopercular pore absent; dorsal in rays
124–129; anal in rays 90–94; total vertebrae 56–59;
D/A 37–42 ......................... Brosmolus longicaudus
12b. upper preopercular pore present; dorsal in rays
93–104; anal in rays 64–76; total vertebrae 45–47;
D/A 28–35................................Monothrix polylepis
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 in rays 16–21; irst 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; irst and second preopercular pores with
separate openings ....................................................
............. Dactylosurculus gomoni gen. nov. sp. nov.
15a. upper preopercular pore absent ...........................16
15b. upper preopercular pore present ........................ 27
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 lower preopercular pores with
separate openings .................................................18
17b. First and second lower preopercular pore joined in
a single opening .................................................. 21
18a. Inner pseudoclasper anteriorly connected to outer
pseudoclasper; sulcus of otolith with undivided
colliculi........................................ (Beaglichthys) 19
18b. Inner pseudoclasper branched, medially connected to
outer pseudoclasper; sulcus of otolith with separate
colliculi, cauda short ...............................................
.................. Zephyrichthys barryi gen. nov. sp. nov.
19a. Dorsal in rays 111–113; precaudal vertebrae 14; total
vertebrae 51–56; D/A 36–37 ....................................
................................ Beaglichthys macrophthalmus
19b. Dorsal in rays 84–98; precaudal vertebrae 12; total
vertebrae 44–46; D/A 23–29 ............................... 20
20a. Dorsal in rays 84–86; anal in rays 63–68; v in D
2.2–2.3; inner pseudoclasper anteriorly inclined with
lateral appendices, joined to the outer pseudoclasper
at base; outer pseudoclasper with distal knob on its
inner face ................. Beaglichthys bleekeri sp. nov.
20b. Dorsal in rays 93–98; anal in rays 69–78; v in
D 2.4–2.6; inner pseudoclasper broadly attached
anteriorly, simple hook shape; outer pseudoclasper
without knob on inner face .....................................
................................ Beaglichthys larsonae sp. nov.
21a. Inner pseudoclasper concave, anteriorly connected
to outer pseudoclasper; precaudal vertebrae 11 ......
..........................................Diancistrus manciporus
[note: only species of genus Diancistrus without
upper preopercular pore, see also 30b]
21b. Inner pseudoclasper claw-like; precaudal vertebrae
12–13 ..........................(Ungusurculus gen. nov.) 22
22a. Dorsal in rays 70–77; D/A 20–23; otolith with spiny
postdorsal angle... Ungusurculus riauensis sp. nov.
22b. Dorsal in rays 76–87; D/A 22–27; otolith without
spiny postdorsal angle ......................................... 23
23a. Supporter of outer pseudoclasper without anterior
hook; inner pseudoclasper with inwardly directed
spines .................................................................. 24
23b. Supporter of outer pseudoclasper with anterior hook;
inner pseudoclasper claw-like, bifurcate ............ 25
24a. Inner pseudoclasper large, with strong spines; otolith
length to otolith height 2.0–2.1; otolith length to
sulcus length 2.0–2.1 ...............................................
......................Ungusurculus philippinensis sp. nov.
24b. In ner pseudoclasper half the size of outer
pseudoclasper, with week spines; otolith length to
otolith height 1.8–2.0; otolith length to sulcus length
2.2–2.4 ..................Ungusurculus williamsi sp. nov.
25a. Total vertebrae 41; otolith length to otolith height
1.8 .......................... Ungusurculus collettei sp. nov.
25b. Total vertebrae 42–44; otolith length to otolith height
2.3–2.4 (not known for Ungusurculus komodoensis
sp. nov.) ................................................................ 26
26a. Total vertebrae 44; head with cirri; no knob on inner
face of outer pseudoclasper itting into opening of
claw-like tip of inner pseudoclasper .......................
....................... Ungusurculus komodoensis sp. nov.
26b. Total vertebrae 42–43; head without cirri; knob on
inner face of outer pseudoclasper itting into opening
of claw-like tip of inner pseudoclasper ...................
.......................... Ungusurculus sundaensis sp. nov.
27a. Single lower preopercular pore .......................... 28
27b. Three lower preopercular pores, but irst and second
pore open into single opening ............................. 29
28a. Inner pseudoclasper about as large as outer
pseudoclasper and separate ................ Pseudogilbia
28b. In ner pseudoclasper half the size of outer
pseudoclasper and anteriorly connected to u-shaped
structure ........................................ Paradiancistrus
29a. Inner pseudoclasper anteriorly joined to outer
pseudoclasper ...................................................... 30
29b. Inner and outer pseudoclaspers free ....................31
33
W. Schwarzhans and P. r. Møller
30a. 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
30b. 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
Beaglichthys Machida, 1993
(Tables 1–5)
Beaglichthys Machida, 1993: 284 (type species
Beaglichthys macrophthalmus Machida, 1993, by
monotypy); Neilsen and Cohen in Nielsen et al. 1999:
117, 125.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed
low on snout; male copulatory organ with two pairs
of pseudoclaspers, the outer large, twice as large as
inner, both joined anteriorly; relatively large (> 50 mm
Sl in mature males); precaudal vertebrae 12–14, total
vertebrae 44–56, v in D 2.2–2.7; caudal in rays 12–16;
branchiostegal rays 6–8; pseudobranchial filaments
0–3; head with scale patch only on cheek, no scales on
operculum; no upper preopercular pore; irst and second
lower preopercular pore with separate openings; otolith
elongate (otolith length to height 2.2–2.3), its sulcus
straight, large (otolith length to sulcus length 1.7–2.2),
colliculi fused; maxilla expanded postero-ventrally;
anterior anal in pterygiophore long.
Remarks. Beaglichthys was originally described by
Machida (1993) as a monospeciic genus and deined by
the following combination of characters: anal in origin
at midpoint of body; cheek scaly; eye diameter longer
than snout length; opercular spine strong; developed
rakers on irst gill arch 3; branchiostegal rays 8; caudal
in rays 12; precaudal vertebrae 14. Of these, the anal
in origin, scaly cheek, opercular spine and three gill
rakers on irst arch are all widespread characters found
in the Dinematichthyini and not of particular diagnostic
value. The 12 caudal in rays are the lowest number now
observed within the genus, which ranges from 12–16
(12–14 within the type species B. macrophthalmus). The
large eye diameter is now considered to be diagnostic for
the species B. macrophthalmus but not for the genus. The
number of branchiostegal rays varies from 6 to 8. The
number of precaudal vertebrae is 12 in the two new species
and 14 in B. macrophthalmus. This character, along with
the dependant number of total vertebrae (44–46 vs 51–56
in B. macrophthalmus) and the predorsal length in % of
Sl (28.9–35.2 vs 25.7–26.8 in B. macrophthalmus) could
be used to separate the two new species – B. bleekeri
sp. nov. and B. larsonae sp. nov. – from the type species
B. macrophthalmus in another genus. For the purpose of
this review we have refrained from such a solution until
the nature and distinction of the two established genera
Beaglichthys Machida, 1993 and Brosmolus Machida,
1993 (see below), can be investigated on a broader base
of specimens than currently available.
Comparison. Beaglichthys appears to be related
most closely to Brosmolus Machida, 1993, from which
it differs in being less slender with lower numbers of
dorsal and anal in rays (84–113 and 63–83 vs 124–129
and 90–94 respectively). The other main difference is the
presence of two pairs of pseudoclaspers in Beaglichthys
31a. Inner pseudoclasper hidden in pocket of isthmus
when in resting position; penis hooked...................
............................... (Mascarenichthys gen. nov.) 32
31b. Inner pseudoclasper open; penis bent, but not
hooked ................................................................. 33
32a. eye size 1.5–2.7 % Sl; pectoral in rays 16–18; outer
pseudoclasper longer than inner pseudoclasper......
....................... Mascarenichthys heemstrai sp. nov.
32b. eye size 0.8–1.2 % Sl; pectoral in rays 19; outer
pseudoclasper shorter than inner pseudoclasper ....
............. Mascarenichthys microphthalmus sp. nov.
33a. Inner pseudoclasper sucker-disk-like; supporter of
outer pseudoclasper distally expanded and usually
with anterior hook .......... (eusurculus gen. nov.) 34
33b. Inner pseudoclasper simple lap or diversiied, but not
sucker-disk-like; supporter of outer pseudoclasper
not expanded and without anterior hook............. 36
34a. Position of anterior nostril at distance of 1/3.5–1/4
to aggregate distance to anterior margin of eye;
precaudal vertebrae 13–14 (rarely 12); inner
pseudoclasper stalked with folded sucker-disk tip;
otolith length to height 2.2–2.4 ...............................
...................................eusurculus pistillum sp. nov.
34b. Position of anterior nostril at distance of 1/4–1/5
to aggregate distance to anterior margin of eye;
precaudal vertebrae 11–12; inner pseudoclasper
stalked with simple or miniature sucker-disk tip;
otolith length to height 2.0–2.1 (not known for e.
andamanensis) .................................................... 35
35a. Inner pseudoclasper with simple sucker-disk tip ....
......................... eusurculus andamanensis sp. nov.
35b. Inner pseudoclasper with miniature sucker-disk tip
with two small hooks ..............................................
...................................eusurculus pristinus sp. nov.
36a. Sulcus of otolith with separated colliculi................
..................................................................... Ogilbia
36b. Sulcus of otolith with undivided colliculi ...............
............. Majungaichthys simplex gen. nov. sp. nov.
34
Dinematichthyine ishes of the Indo-west Paciic III
Table 1. Comparison matrices of selected characters used for distinguishing dinematichthyine genera: A, summarises selected morphological
and meristic characters; B, summarises pseudoclasper and otolith characters. roman numbers in the second column indicate parts of the
review of the Dinematichthyini for America (two parts) and the Indo-west Paciic (four parts, part Iv outstanding).
II
I
Indo-west Paciic
II
III
Iv
east Paciic –
West Atlantic
Gunterichthys
Indo-west Paciic
cheeks
cheeks + above
opercul. spine
entire
few on cheeks
Scales on head
none
3
2*
1
absent
upper lower preoperc.
preop. pore opening
pore
(*1st + 2nd
joined)
present
15+
14
13
12
11
low
intermediate
high
shallow
Precaudal vertebrae
anterior
nostril
Gunterichthys
Ogilbichthys
Pseudogilbia
Typhliasina
Ogilbia
Brotulinella
Diancistrus
Paradiancistrus
Dermatopsis
Dermatopsoides
Dipulus
Beaglichthys
Brosmolus
Dactylosurculus
Didymothallus
eusurculus
Lapitaichthys
Majungaichthys
Mascarenichthys
Monothrix
Ungusurculus
Zephyrichthys
Dinematichthys s.l.
B
I
expanded
free
east Paciic –
West Atlantic
I
hidden
Opercular Maxilla
spine
termination
A
Ogilbichthys
Pseudogilbia
Typhliasina
II Ogilbia
Brotulinella
I Diancistrus
Paradiancistrus
Dermatopsis
II Dermatopsoides
Dipulus
Beaglichthys
Brosmolus
Dactylosurculus
Didymothallus
eusurculus
Lapitaichthys
III
Majungaichthys
Mascarenichthys
Monothrix
Pseudoclaspers
1 2 3
special characters of the
special characters of the
pair pairs pairs inner pseudoclasper(s) (i.p.) outer pseudoclasper (o.p.)
two long supporters
separated in anterior and
posterior i.p.
as large as o.p.
very diversiied (atrophied
in one species)
leshy appendix of o.p.
anteriorly connected to o.p. hook, ear-lobe or wing shape
anteriorly connected to o.p.
one or two supporters
2nd inger-like i.p. inserted all pseudoclaspers joined
between 1st i.p. + o.p.
at base
two long supporters
sucker-disk-like tip
supporter with anterior hook
hidden in pocket of isthmus
massive to claw-like tip
Zephyrichthys
branched + connected to
o.p.
Iv Dinematichthys s.l.
Other characters
penis straight
blind
slender body
penis with thorn
anteriorly connected to o.p.
Ungusurculus
Otoliths
separate undivided
colliculi colliculi
elongate body
8 branchiostegal rays
penis straight
penis hooked
supporter often with anterior
hook
sulcus with short cauda
35
W. Schwarzhans and P. r. Møller
Beaglichthys bleekeri sp. nov.
(Figs 1–3; Tables 2–3)
Material examined. (4 specimens, 49–72 mm Sl).
hOlOTyPe – SMNS 10671, male, 72 mm Sl, Java (no
further detail given), Indonesia; specimen donated by
P. Bleeker in Jan. 1860. ParatyPeS – uSNM 374173,
female, 62 mm Sl, 12°05’S, 131°06’e, east vernon
Island, Northern Territory, Australia, tide pools, coll. B.
B. Collette, 31 May 1979; WAM P. 31250-046, female, 66
mm Sl, 15°17’S, 124°10’e, Kimberley, Western Australia,
depth 0.5 m, coll. S. M. Morrison, 2 Dec. 1996; WAM
P.31832–003, female, 49 mm Sl, 20°37’S, 116°33’e,
enderby Island, Dampier Archipelago, Australia, coll. J.
B. hutchins, 9 Aug. 2000.
Diagnosis. vertebrae 12+32-33=44-45, dorsal in rays
84–86, anal in rays 63–68, pectoral in rays 21–22, caudal
in rays 14–15, v in D 2.2–2.3; pseudobranchial ilaments
0–1; eyes small (1.8–2.4 % Sl); outer pseudoclasper broadbased with pointed tip and distal knob on inner side; inner
pseudoclasper nearly as large as outer pseudoclasper,
anteriorly connected to it, forwardly inclined with broad
leshy appendices; cheek with 5–6 rows of scales on upper
part and 2–3 rows on lower part; otolith with centrally
positioned undivided sulcus, otolith length to otolith height
= 2.3, otolith length to sulcus length = 2.2.
Description (Figs 2, 3).The principal meristic and
morphometric characters are shown in Table 3. Body
slender, but with massive head. Mature at about 50 mm
Sl. head with scale patch on cheek containing 5 to 6
vertical rows of scales on upper part and 2–3 vertical
rows on lower part. horizontal diameter of scales on
body about 1.3 % Sl, in 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
low, 1/5 distance from upper lip to aggregate distance
to anterior margin of eye. Posterior nostril moderate in
size, about half the size of eye. Opercular spine with
free tip, thin and sharply pointed. Anterior gill arch with
15–18 (18) rakers, 3–4 (3) elongate. In holotype, row
of elongate rakers interrupted by one small plate-like
raker (not seen in paratypes). Pseudobranchial ilaments
single or absent (1).
Head sensory pores (Fig. 3A). Supraorbital pores 2
to 3: irst pore in front of second anterior infraorbital
pore, second pore indistinct above and behind eye, 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,
less than 1/3 the size of three anterior pores. Mandibular
pores 6 (3 anterior and 3 posterior): irst anterior pore large
and tubular, with single cirrus anteriorly, second anterior
pore positioned in lateral skin fold, small, third anterior
pore at anterior termination of jugular isthmus, three
vs one pair in Brosmolus. In the case of Brosmolus,
however, only minute pseudoclaspers are known from
the single male holotype, which appears to be pre-adult
(see below). Without this difference in pseudoclaspers, we
would suggest synonymising both genera, in which case
Brosmolus would gain priority.
Beaglichthys shares the combination of two pairs of
pseudoclaspers and the lack of an upper preopercular
pore with Ungusurculus gen. nov., Zephyrichthys gen.
nov., some species of Dinematichthys Bleeker, 1855,
and Diancistrus manciporus Schwarzhans, Møller and
Nielsen, 2005, in the Indo-west Paciic and some species
of Ogilbichthys Møller, Schwarzhans and Nielsen, 2004
(although with two inner pseudoclaspers) and Typhliasina
Whitley, 1951, of the American Dinematichthyini.
Ungusurculus gen. nov. differs from Beaglichthys in the
claw-like development of the free inner pseudoclaspers,
where the inner pseudoclasper is anteriorly positioned
and joined to the outer pseudoclasper. A similar anteriorly
joined inner pseudoclasper is seen in Zephyrichthys gen.
nov. and Diancistrus Ogilby, 1899. Zephyrichthys gen. nov.
has a bifurcate branched inner pseudoclasper (vs single
unbranched in Beaglichthys) and otoliths with separated
colliculi (vs fused in Beaglichthys). Beaglichthys is
distinguished from the single Diancistrus species without
upper preopercular pore (D. manciporus) by having
12–14 precaudal vertebrae (vs 11). It is distinguished from
species of Dinematichthys without an upper preopercular
pore by the low position of the anterior nostril (vs high),
the anteriorly joined inner pseudoclasper (vs free), the
higher number of precaudal vertebrae (12–14 vs 11) and
the fused colliculi on the otolith (vs separated). The two
New World genera Ogilbichthys and Typhliasina have
11 precaudal vertebrae (vs 12–14 in Beaglichthys) and
free inner pseudoclaspers (two inner pseudoclaspers in
the case of Ogilbichthys) (vs anteriorly joined to outer
pseudoclasper in Beaglichthys).
Beaglichthys, Brosmolus, Dactylosurculus gen. nov.
and Zephyrichthys gen. nov. are unique amongst the
dinematichthyine genera of this part of the review in
exhibiting separate openings of the irst and the second
lower preopercular pores, a character otherwise only found
in the genera Dermatopsoides Smith, 1947, and Dipulus
Waite, 1905, which however differ in many other signiicant
characters (see Møller and Schwarzhans 2006).
Distribution. Beaglichthys was originally described as
a monospeciic genus based on a single female specimen
from northern Australia (Beagle Gulf) described by
Machida (1993) as B. macrophthalmus. With the two
new species described here – B. bleekeri sp. nov. and B.
larsonae sp. nov. – the range of the distribution of the
genus now extends from Java in Indonesia and along the
northern Australian shores to Daru, southern New Guinea,
Gulf of Papua (Fig. 1).
36
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 1. Sample sites of Beaglichthys bleekeri sp. nov., B. larsonae sp. nov., B. macrophthalmus and Brosmolus longicaudus.
One symbol may represent several samples.
posterior pores on rear part of lower jaw. Preopercular
pores: 3 lower, irst and second with separate opening;
third non-tubular; no upper preopercular pores. [This
description of the position of head sensory pores is used
as reference for all subsequent descriptions.]
Dentition (of holotype). Premaxilla with six outer
rows of granular teeth and two inner rows of larger teeth
anteriorly. Anteriormost teeth in inner row up to 2/3
diameter of pupil. vomer horseshoe-shaped, with four
outer rows of small teeth and one inner row of large teeth
up to 1/2 diameter of pupil. Palatine with three outher rows
of small teeth and one inner row of large teeth up to 1/2
diameter of pupil. Dentary with four outer rows of granular
teeth and one inner row of larger teeth anteriorly, merging
into one row of larger teeth posteriorly, up to about size
of pupil diameter.
Fig. 2. Beaglichthys bleekeri sp. nov. A, fresh dead, WAM P.31832-003, female, 49 mm Sl; B, SMNS 10671, holotype, male, 72 mm Sl.
37
W. Schwarzhans and P. r. Møller
Colour. Beaglichthys bleekeri is known from a
photograph of a freshly caught specimen (WAM P.31832003, Fig. 2A), which shows a milky yellowish, partly
translucent ish, with slightly darker dorsum and yellow
vertical in bases. The rear of the body is red-orange in a
narrow strip above the vertebral column. Preserved colour
medium brown.
Remarks. The holotype was originally part of the
Bleeker collection, who also described the first ever
dinematichthyine in 1855 from a single specimen of
Dinematichthys iluocoeteoides from Batu Island (now
Kepulauan Batu, Sumatera Barat Province), off western
Sumatra, Indonesia. Bleeker’s holotype has been lost
(see extensive discussion in Cohen and Nielsen (1978)).
The re-deinition of the genus Dinematichthys and the
species D. iluocoeteoides are therefore still pending and
will be included in the next part of this world-wide review
of the Dinematichthyini. Cohen and Nielsen (1978) also
discuss the nature of a specimen from Bleeker’s original
collection (and assigned by Günther (1862)) kept in london
(BMNh 1868.2.28.65) but they 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
Otolith (Fig. 3 B, C). elongate in shape, length to height
2.3 (50–66 mm Sl) and moderately thin (otolith height
to otolith thickness about 2.5). Anterior and posterior
tips moderately pointed. Dorsal rim gently curved, with
indistinct pre- and postdorsal angles; ventral rim likewise
gently curved. Inner face moderately convex, outer face
lat to slightly convex, both smooth. Otolith length to
sulcus length 2.2. Sulcus centrally positioned, with fused
colliculi, not inclined to otolith. ventral furrow indistinct,
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined
and 5–8 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11. First anal
in pterygophore elongate, but not reaching tip of last
precaudal parapophysis.
Male copulatory organ (Fig. 3 D, e). Two pairs of
large pseudoclaspers, outer pair only about 50% larger
than inner; outer pseudoclasper with broad base and
pointed tip and distinct knob at inner face near tip; inner
pseudoclasper strongly forwardly inclined, joined to outer
pseudoclasper at base, with single strong supporter in
middle and large leshy appendices on either side. Penis
curved, shorter than outer pseudoclaspers.
Fig. 3. Beaglichthys bleekeri sp. nov. A, lateral view of head, WAM P.31832-003, female, 49 mm Sl; B, median view of right otolith,
WAM P.31250-046, female, 66 mm Sl; C, ventral view of right otolith, WAM P.31250-046, female, 66 mm Sl; D, inclined lateral view
of male copulatory organ, holotype (note right outer pseudoclasper detached anteriorly through rupture of ligament); E, view of left
pseudoclasper from inside, holotype.
38
Dinematichthyine ishes of the Indo-west Paciic III
Table 2. Comparison of selected meristic and morphometric characters of the species of the genera treated in Part III of the study:
A, frequency distribution of dorsal in rays counts; B, frequency distribution of anal in rays counts; C, frequency distributions of
caudal in rays counts, precaudal vertebrae and total vertebrae counts; D, Frequency distribution of D/A, v/A and D in v; E, frequency
distribution of selected morphometric characters.
Frequency distribution of dorsal in rays counts
62-63
64-65
66-67
68-69
70-71
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
104-105
106-107
108-109
110-111
112-113
114-115
116-117
118-119
120-121
122-123
124-125
126-127
128-129
A
Beaglichthys bleekeri
B. larsonae
B. macrophthalmus
Brosmolus longicaudus
Dactylosurculus gomoni
2 5 6
Didymothallus criniceps
D. mizolepis
2 2
D. pruvosti
eusurculus andamanensis
e. pistillum
e. pristinus
Lapitaichthys frickei
1 2
Majungaichthys simplex
Mascarenichthys heemstrai 5 15 19 21 7 5
1 1
M. microphthalmus
1 1
M. sp.
Monothrix polylepis
Ungusurculus collettei
U. komodoensis
U. philippinensis
5 5
U. riauensis
U. sundaensis
U. williamsi
Zephyrichthys barryi
1
Paradiancistrus lombokensis
3 1
2 2 2 1
9 4
2 - 1
1 1 2 10 6 7 5 2 3
4 20 21 25 27 14 10 8 1 2
2 - 2 1
5 6 11 20 19 6 2
3 - 4 5 8 4 1
1 - 6 11 7 4
1 - 2
1 7 9 5 3 5 1
1
7 4
1
4 15 16 11 4
1 - - - 1
1 4 2 4
2 6 10 15 8 5 1
2
78-79
-
1
6
5
14
10 1 1
3 10 29 31 20 19 10 4 3
1 2 - 1
1 1 3
5 13 18 17 13 1
3 4 11 4 1 1 1
10 6 6 5
2 2 1 1
11 23 23 11 2
1 1
2
2 4 4 11 7
1
1
1 1 4 13 19 9 2
1 5 11 2 2
1 - 1
2 5 4
8 8 11 10
1
39
9
5
1
1
6
2
1
2
2
3
94-95
76-77
-
92-93
74-75
-
90-91
72-73
4
88-89
70-71
1
2
86-87
68-69
1
84-85
66-67
1
82-83
64-65
1
80-81
62-63
58-59
56-57
54-55
52-53
50-51
48-49
46-47
7
60-61
Frequency distribution of anal in rays counts
B
Beaglichthys bleekeri
B. larsonae
B. macrophthalmus
Brosmolus longicaudus
Dactylosurculus gomoni
Didymothallus criniceps
D. mizolepis
D. pruvosti
eusurculus andamanensis
e. pistillum
e. pristinus
Lapitaichthys frickei
Majungaichthys simplex
Mascarenichthys heemstrai
M. microphthalmus
M. sp.
Monothrix polylepis
Ungusurculus collettei
U. komodoensis
U. philippinensis
U. riauensis
U. sundaensis
U. williamsi
Zephyrichthys barryi
Paradiancistrus lombokensis
1 3
1
1
1
W. Schwarzhans and P. r. Møller
C
Caudal in rays
Precaudal
vertebrae
Frequency distribution of total vertebrae
12 13 14 15 16 11 12 13 14 15 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59
Beaglichthys bleekeri
B. larsonae
B. macrophthalmus
Brosmolus longicaudus
Dactylosurculus gomoni
Didymothallus criniceps
D. mizolepis
D. pruvosti
eusurculus andamanensis
e. pistillum
e. pristinus
Lapitaichthys frickei
Majungaichthys simplex
Mascarenichthys heemstrai
M. microphthalmus
M. sp.
Monothrix polylepis
Ungusurculus collettei
U. komodoensis
U. philippinensis
U. riauensis
U. sundaensis
U. williamsi
Zephyrichthys barryi
Paradiancistrus lombokensis
4
7
3
1 2
36 3
(1) 25
132 5
4
5
1 64 2
12 13
32
6
69
2
2
16 15
1
1
45 4
22
2
12
51 4
2
2 2
3 3 1
3 15 5
4 42 39 24 27
1 2
3 2
4 32
3 4 14 4
9 17
3 3
14 39 20 1
1 1
2
Beaglichthys bleekeri
B. larsonae
B. macrophthalmus
Brosmolus longicaudus
Dactylosurculus gomoni
Didymothallus criniceps
D. mizolepis
D. pruvosti
eusurculus andamanensis
e. pistillum
e. pristinus
Lapitaichthys frickei
Majungaichthys simplex
Mascarenichthys heemstrai
M. microphthalmus
M. sp.
Monothrix polylepis
Ungusurculus collettei
U. komodoensis
U. philippinensis
U. riauensis
U. sundaensis
U. williamsi
Zephyrichthys barryi
Paradiancistrus lombokensis
13 8
11
3 1
1 - 2
2
6 3 7 6
4 - 2
1 8 44 17 1
1 - 1
1 1
3 5 11 15 5
3 2
33 57 11 3
-
1 7 16 9 3 1
2
1
1
-
- 1
1 1 - 1
28 2 1
6
7 15 9
1
19 25 7
1 11 10
1 1
7 4
7 20 26 1
1 - - 1
D/A (1st dorsal above 1st anal)
1 2 1
2 2 3
1 -
1
v/A (1st anal below
vertebra)
12-13
14-15
16-17
18-19
20-21
22-23
24-25
26-27
28-29
30-31
32-33
34-35
36-37
38-39
40-41
42-43
D
3 1
3 2 1
1 - 1
1
11 1
1 7 10 1
4 14
1 1
2
4 10
1 3
1 9 21
4 1
7 9
2
2
1 7 2
2
1
8 11
12
1 1
5
4 24 6
13 14 15 16 17 18 19 1.8 1.9 2.0 2.1 2.2 2.3 2.4 2.5 2.6 2.7
3
1 - 1 1
2
14 41 11 1
3
4 10 12 4
7
1
25
1
5 12 12 2
1
1
6 30 14
15 7
2
10 1
6 25 13 1
1
1
1
40
D in v (dorsal in rays per rayed
vertebra)
3 - 1
1 6
3
1 - 2
21 18
21 5
49 80 6
2 2
2 3
4 61 4
12 6
4 17 6 1
3 2
43 1
2
1
6 21 4
1
1
12 37 1
20
2
6 5
30 12 1
1
2 2
3
1 2
1 11
8 13 5
4
3 1
1 - 1
1
19 8
9 44 57 20 5
3 2
4 26 25 5
2 4 12 7
9 11 8
3 1 2
2 24 34 11
2
1 1
14 11 6
1
1
24 23 3
1 4 14 3
1 1
2 8 1
7 21 10 3
1
Dinematichthyine ishes of the Indo-west Paciic III
Body depth at
Predorsal length in
Base ventral in - anal in
head height in
eye size in % Sl
origin of anal in
% Sl
origin in % Sl
% Sl
in % Sl
Beaglichthys bleekeri
B. larsonae
B. macrophthalmus
Brosmolus longicaudus
Dactylosurculus gomoni
Didymothallus criniceps
D. mizolepis
D. pruvosti
eusurculus andamanensis
e. pistillum
e. pristinus
Lapitaichthys frickei
Majungaichthys simplex
Mascarenichthys heemstrai
M. microphthalmus
M. sp.
Monothrix polylepis
Ungusurculus collettei
U. komodoensis
U. philippinensis
U. riauensis
U. sundaensis
U. williamsi
Zephyrichthys barryi
Paradiancistrus lombokensis
> 1.5
1.5-1.9
2.0-2.4
2.5-2.9
3.0-3.5
> 26.0
26.0-27.9
28.0-29.9
30.0-31.9
32.0-33.9
34.0-36.0
> 12.0
12.0-13.9
14.0-15.9
16.0-17.9
18.0-19.9
23.0-24.9
25.0-26.9
27.0-28.9
29.0-30.9
31.0-32.9
32.0-33.9
34.0-35.9
head length in
% Sl
> 23.0
23.0-24.4
24.5.-25.9
26.0-27.4
27.5-28.9
> 12.0
12.0-13.4
13.5-14.9
15.0-16.4
16.5-17.5
E
x x x
x x x
x x x
x x
x x x x
x x
x x
x x
x x
x x
x x x
x x x x x
x x x x
x x
x
x x
x x x
x x
x x
x x
x
x
x
x x x
x x x x
x x
x x x
x x
x x
x x x x
x x
x x x
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Fishes, version 19 June 2007, http://www.calacademy.org/
research/ichthyology/catalog/ishcatmain.asp)). It is a male
specimen and although dried and shrivelled may serve
as neotype in our forthcoming review since it resembles
Bleeker’s description in all important aspects.
Another specimen, assigned as D. iluocoeteoides from
Bleeker’s original collection, has been brought to our
attention by r. Fricke from the SMNS collection caught
off Java (SMNS 10671), representing a male of a different
species, selected here as holotype for B. bleekeri.
Bleeker mentioned in his description of D. iluocoeteoides
that the nostrils were close to the eye and that head scales
were present on nape, preopercle and opercle. These
characters are shared by the BMNh specimens, but not
by SMNS 10671.
Comparison. Beaglichthys bleekeri differs from
B. larsonae sp. nov. in the lower number of dorsal and
anal in rays (84–86 and 63–68 vs 93–98 and 69–78), the
lower index v in D (2.2–2.3 vs 2.4–2.6), the anteriorly
inclined inner pseudoclasper with lateral appendices and
joined to the outer pseudoclasper at its base (vs broadly
anteriorly attached inner pseudoclasper with simple
hook shape), the triangular shaped outer pseudoclasper
with the distal knob at its inner face (vs wing shaped)
and the sulcus placed symmetrically on the inner
face of the otolith (vs posteriorly expanded). From B.
macrophthalmus it differs in the lower vertebrae count
(12+32-33 vs 14+37-42), the lower dorsal and anal in
ray counts (84–86 and 63–68 vs 111–113 and 83), the
morphology of the pseudoclaspers (similar to difference
with B. larsonae sp. nov.) and the short sulcus (otolith
length to sulcus length 2.2 vs 1.7).
Distribution. Off Java (further details unknown)
and along the north-western coast of Australia from the
Dampier Archipelago to the vernon Islands north-east of
Darwin (Fig. 1).
Etymology. Named in memory of Pieter Bleeker, the
outstanding ichthyologist of the Indo-west Paciic during
the early years, who also collected the holotype of this
species.
Beaglichthys larsonae sp. nov.
(Figs 1, 4, 5; Tables 2, 4)
Material examined. (7 specimens, 56–76 mm Sl).
hOlOTyPe – uSNM 327954, male, 60 mm Sl, 12°29’S,
130°53’e, Darwin harbour, Northern Territory, Australia,
coll. P. C. heemstra, h. K. larson and r. S. Williams,
18–19 Feb. 1988. PArATyPeS – AMS I.24676-050, 1 male,
56 mm Sl, 2 females, 61–76 mm Sl, 12°29’S, 130°53’e,
Darwin harbour, Northern Territory, Australia, depth 0–1
41
W. Schwarzhans and P. r. Møller
Table 4. Meristic and morphometric characters of Beaglichthys
larsonae sp. nov.
Table 3. Meristic and morphometric characters of Beaglichthys
bleekeri sp. nov.
holotype
SMnS
10671
Standard length in mm
72
Meristic characters
Dorsal in rays
84
Caudal inrays
14
Anal in rays
63
Pectoral in rays
21
Precaudal vertebrae
12
Caudal vertebrae
32
Total vertebrae
44
rakers on anterior gill arch
18
Pseudobranchial ilaments
1
D/v
6
d/a
25
v/A
15
Morphometric characters in % of SL
head length
24.4
head width
10.3
head height
12.9
Snout length
4.4
upper jaw length
12.1
Diameter of pigmented eye
1.8
Diameter of pupil
1.1
Interorbital width
6.2
Posterior maxilla height
4.3
Postorbital length
16.9
Preanal length
46.6
Predorsal length
28.9
Body depth at origin of anal in
13.9
Pectoral in length
11.5
Pectoral in base height
5.4
ventral in length
17.8
Base ventral in – anal in origin 29.8
holotype +
3 paratypes
holotype
uSNM327954
n
Mean (range)
61.5 (49-72)
4
85.0 (84-86)
14.7 (14-15)
65.5 (63-68)
21.3 (21-22)
12
32.5 (32-33)
44.5 (44-45)
16.5 (15-18)
0.5 (0-1)
6.5 (6-7)
24.8 (23-27)
15.5 (15-17)
4
4
4
4
4
4
4
4
4
4
4
4
26.8 (24.4-28.7)
12.3 (10.3-14.1)
15.3 (12.9-16.5)
5.2 (4.4-6.3)
13.0 (12.1-14.1)
2.0 (1.8-2.4)
1.3 (1.1-1.6)
6.3 (5.8-6.8)
4.0 (3.4-4.3)
19.9 (16.9-21.4)
48.8 (46.6-50.3)
32.6 (28.9-35.2)
15.4 (13.9-17.5)
13.4 (11.5-15.0)
5.5 (5.2-5.9)
17.7 (14.4-19.8)
30.7 (29.8-32.0)
3
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
Standard length in mm
60
Meristic characters
Dorsal in rays
94
Caudal inrays
14
Anal in rays
71
Pectoral in rays
24
Precaudal vertebrae
12
Caudal vertebrae
33
Total vertebrae
45
rakers on anterior gill arch
16
Pseudobranchial ilaments
1
D/v
6
d/a
28
v/A
15
Morphometric characters in % of SL
head length
26.2
head width
11.9
head height
15.1
Snout length
5.5
upper jaw length
12.7
Diameter of pigmented eye
2.2
Diameter of pupil
1.2
Interorbital width
6.1
Posterior maxilla height
4.0
Postorbital length
19.0
Preanal length
47.0
Predorsal length
32.2
Body depth at origin of anal in
15.5
Pectoral in length
14.7
Pectoral in base height
5.1
ventral in length
19.1
Base ventral in – anal in origin 29.5
m, coll. D. hoese and party, 29 August 1984; ruSI 35938,
female, 70 mm Sl, Darwin harbour, Northern Territory,
Australia, coll. P. C. heemstra, 18 Feb 1988; uSNM 374172,
female, 66 mm Sl, Groote eylandt, Gulf of Carpentaria,
Northern Territory, Australia, depth 0–1 m, coll. r. Miller
et al., 25 April 1948; ZMuC P771618, 1 female, 69 mm Sl,
same data as AMS I.24676-050.
holotype +
6 paratypes
n
Mean (range)
65.3 (56-76)
7
95.0 (93-98)
14.7 (14-16)
71.3 (69-78)
23.0 (22-24)
12
32.7 (32-34)
44.7 (44-46)
15.7 (14-17)
0.7 (0-1)
6.3 (6-7)
27.0 (25-29)
14.9 (14-15)
7
6
7
7
6
7
7
7
7
7
7
7
26.4 (25.6-27.8)
13.6 (11.9-15.0)
16.1 (15.0-17.1)
5.5 (3.9-6.7)
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
5
7
2.0 (1.8-2.2)
1.3 (1.0-1.5)
6.5 (5.9-7.1)
4.0 (3.7-4.3)
19.4 (18.2-20.7)
49.0 (47.0-51.8)
33.5 (32.1-34.5)
17.5 (15.5-19.3)
14.5 (12.3-15.2)
5.4 (4.9-6.1)
18.1 (15.3-21.6)
31.7 (28.6-34.9)
Diagnosis. vertebrae 12+32-34=44-46, dorsal in rays
93–98, anal in rays 69–78, pectoral in rays 22–24, caudal
in rays 14–16, v in D 2.4–2.6; pseudobranchial ilaments
0–1; eyes small (1.8–2.2 % Sl); outer pseudoclasper wingshaped with backwardly directed tip; inner pseudoclasper
nearly as large as outer pseudoclasper, broadly connected
to it anteriorly, similarly wing-shaped; cheeks with 5–7
Fig. 4. Beaglichthys larsonae sp. nov., uSNM 327954, holotype, 60 mm Sl.
42
Dinematichthyine ishes of the Indo-west Paciic III
rows of scales on upper part and 1–2 rows on lower part;
otolith with undivided sulcus, its centre anteriorly of the
centre of the otolith, otolith length to otolith height = 2.2,
otolith length to sulcus length = 2.1.
Description (Figs 4, 5).The principal meristic and
morphometric characters are shown in Table 4. Body and
head slender, snout pointed. Mature at about 50 mm Sl.
head with scale patch on cheek containing 5–7 (7) vertical
rows of scales on the upper part and 1–2 vertical rows on
the lower part. horizontal diameter of scales on body about
1.5 % Sl, in 22 horizontal rows. Maxillary ending far
behind eyes, dorsal margin of maxillary covered by upper
lip dermal lobe, posterior end expanded, angular. Anterior
nostril positioned low, 1/5 the distance from upper lip to
aggregate distance to anterior margin of eye. Posterior
nostril small, about 1/5 the size of eye. Opercular spine
with free tip, pointed. Anterior gill arch with 14–17 (16)
rakers, thereof 3–4 (3) elongate rakers. Pseudobranchial
ilaments single or absent (1).
Head sensory pores (Fig. 5A). Supraorbital pores 2
to 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).
Preopercular pores: 3 lower, irst and second with separate
opening; third non-tubular; no upper preopercular pores.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to four
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 1/2 diameter
of pupil. vomer horseshoe-shaped, with two rows of
equally sized teeth up to 1/4 diameter of pupil. Palatine
with a single row of 12 teeth up to 1/4 diameter of pupil.
Dentary with three outer rows of granular teeth and one
inner row of larger teeth anteriorly, merging into one row
of larger teeth posteriorly, up to about size of 2/3 of pupil
diameter.
Otolith (Fig. 5e). elongate in shape, length to height 2.2
(79 mm Sl) and moderately thin (otolith height to otolith
thickness about 2.5). Anterior tip moderately pointed,
posterior rim expanded, rounded. Dorsal rim gently curved,
convex anteriorly, broadly concave posteriorly; ventral rim
gently and regularly curved. Inner face moderately convex,
outer face lat, both smooth. Otolith length to sulcus length
2.1. Sulcus positioned slightly towards anterior, with fused
Fig. 5. Beaglichthys larsonae sp. nov. A, lateral view of head, holotype; B, view of left pseudoclasper from inside, AMS I.24676-050, male,
56 mm Sl; C, view of left pseudoclasper from inside, holotype; D, inclined lateral view of male copulatory organ, holotype; E, median
view of right otolith, AMS I.24676-050, male, 56 mm Sl.
43
W. Schwarzhans and P. r. Møller
colliculi, not inclined to otolith. ventral furrow distinct,
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebra 5 inclined
and 6–8 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis.
Male copulatory organ (Fig. 5B–D). Two pairs of
large pseudoclaspers and of similar shape, outer pair
only about 50% larger than inner; outer pseudoclasper
moderately broadened base, wing-shaped, with tip directed
backward; inner pseudoclasper similarly wing-shaped
with backward directed tip, anteriorly broadly joined
to outer pseudoclasper, occasionally with small leshy
lap at joined. Penis curved, slightly longer than outer
pseudoclaspers.
Colour. live colour unknown. Preserved colour light
to medium grey-brown with darker back.
Comparison. For correlation of Beaglichthys larsonae
with B. bleekeri see above. From B. macrophthalmus it
differs in the lower vertebrae count (12+32-34 vs 14+37-42),
the lower dorsal and anal in ray counts (93–98 and 69–78
vs 111–113 and 83), the wing-shaped inner pseudoclaspers
(vs pad-shaped) and the short sulcus (otolith length to
sulcus length 2.1 vs 1.7).
Distribution. Beaglichthys larsonae so far is only
known from two locations, Darwin harbour and Groote
eylandt, Gulf of Carpentaria, both in the Northern
Territory of Australia (Fig. 1).
Ecology. The two locations, from which the species
has exclusively been obtained so far are both inshore
environments.
Etymology. Named in honour of helen K. larson,
Darwin, Australia, and her many contributions to the
knowledge of the ish fauna of the Northern Territory of
Australia.
Table 5. Meristic and morphometric characters of Beaglichthys
macrophthalmus Machida, 1993.
Beaglichthys macrophthalmus Machida, 1993
(Figs 1, 6, 7; Tables 2, 5)
Beaglichthys macrophthalmus Machida, 1993: 285;
Nielsen et al. 1999: 117.
Material examined. (3 specimens, 70–96+ mm Sl).
hOlOTyPe – NTM S.10395-001-1, female, 78 mm Sl,
Shoal Bay, Beagle Gulf, Northern Territory, Australia,
depth unknown, coll. N. T. Fisheries, 29 June 1973.
Additional specimens. WAM P.28155-019, male,
96+ mm Sl (damaged tail), 9°05’S, 143°15’e, off Daru
Island, Gulf of Papua, southern Papua New Guinea;
WAM P.31096-021, female, 70 mm Sl, 13°45’S, 126°48’e,
Stewart Island north of Kalumburu, Kimberleys, Western
Australia.
Diagnosis. vertebrae 14+37-42=51–56, dorsal in rays
111–113, anal in rays 83, pectoral in rays 20–22, caudal
in rays 12–14, v in D 2.3–2.7; pseudobranchial ilaments
2–3; eyes large (2.7–3.3 % Sl); outer pseudoclasper broad
wing-shaped with short, backward directed tip; inner
pseudoclasper about half the size of outer pseudoclasper,
broadly connected anteriorly to outer pseudoclasper with
expanded pad-shaped tip; cheeks with 10–11 rows of small
scales on upper part and lacking or up to three rows on
lower part; otolith with undivided sulcus, its centre anterior
to centre of otolith, otolith length to otolith height = 2.2–
2.3, sulcus long, otolith length to sulcus length = 1.7.
Description (Figs 6, 7).The principal meristic and
morphometric characters are shown in Table 5. Body and
head very slender; available specimens 70 to 95 mm Sl.
head with scale patch on cheek containing 10–11 vertical
rows of small scales on upper part and no scales or up
to three vertical scale rows on lower part. horizontal
diameter of scales on body about 0.8 % Sl, in 23 horizontal
rows (in 70 mm Sl female). Maxillary ending well behind
eyes, dorsal margin of maxillary covered by upper lip
dermal lobe, posterior end expanded, angular. Anterior
nostril positioned low, 1/5 distance from upper lip to
holotype
ntM
10395001-1
80
Standard length in mm
Meristic characters
Dorsal in rays
111
Caudal inrays
12
Anal in rays
83
Pectoral in rays
22
Precaudal vertebrae
14
Caudal vertebrae
37
Total vertebrae
51
rakers on anterior gill arch
12
Pseudobranchial ilaments
2
D/v
6
d/a
37
v/A
17
Morphometric characters in % of SL
head length
22.5
head width
13.3
head height
12.9
Snout length
4.1
upper jaw length
11.2
Diameter of pigmented eye
3.3
Diameter of pupil
2.0
Interorbital width
5.2
Posterior maxilla height
3.8
Postorbital length
15.7
Preanal length
49.7
Predorsal length
25.7
Body depth at origin of anal in
14.6
Pectoral in length
13.3
Pectoral in base height
5.0
ventral in length
16.8
Base ventral in – anal in origin 31.9
* One non-type with damaged tail.
44
holotype +
2 non-types
n
Mean (range)
70-96+
3*
112 (111-113)
13 (12-14)
83
21.3 (20-22)
14
39.5 (37-42)
53.5 (51-56)
15.0 (12-18)
2.3 (2-3)
6
36.3 (36-37)
17
2
2
2
3
3
2
2
3
3
3
3
3
22.8 (22.5-23.2)
12.1 (10.9-13.3)
13.2 (12.9-13.5)
4.4 (4.1-4.7)
11.2
3.0 (2.7-3.3)
1.7 (1.5-2.0)
4.9 (4.6-5.2)
3.5 (3.2-3.8)
16.2 (15.7-16.7)
47.5 (45.3-49.7)
26.2 (25.7-26.8)
14.0 (13.4-14.6)
13.4 (13.3-13.5)
4.9 (4.8-5.0)
16.9 (16.8-16.9)
30.1 (28.3-31.9)
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 6. Beaglichthys macrophthalmus Machida, 1993. A, fresh dead, B, preserved, WAM P.31096-021, female, 70 mm Sl.
aggregate distance to anterior margin of eye. Posterior
nostril small, about 1/4 size of eye. Opercular spine with
free tip, strong, pointed. Anterior gill arch with 12–18 (12)
rakers, thereof 3–5 (3) elongate rakers. Pseudobranchial
ilaments 2–3 (2).
Head sensory pores (Fig. 7A). Supraorbital pores 2
to 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).
Preopercular pores: 3 lower, irst and second with separate
opening; third non-tubular; no upper preopercular pores.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of a 70 mm Sl female non-type). Premaxilla
with up to four outer rows of granular teeth and one inner
row of larger teeth. Anteriormost teeth in inner row up to
1/3 diameter of pupil. vomer horseshoe-shaped, with two
rows of equally sized teeth up to 1/5 diameter of pupil.
Palatine teeth in two rows, teeth in inner row very small
– in outer row up to 1/5 diameter of pupil. Dentary with
four outer rows of granular teeth and one inner row of
larger teeth anteriorly, merging into one row of larger teeth
posteriorly, up to about size of 1/2 of pupil diameter.
Otolith (Fig. 7e). elongate in shape, length to height
2.2–2.3 (70–95 mm Sl) and moderately thin (otolith height
to otolith thickness about 2.5). Anterior tip moderately
pointed, posterior tip slightly expanded. Dorsal rim gently
curved to nearly lat, convex anteriorly; ventral rim gently
and evenly curved. Inner face moderately convex, outer
face lat, both smooth. Sulcus large, otolith length to sulcus
length 1.7. Sulcus positioned slightly towards anterior,
with fused colliculi, not or slightly inclined to otolith axis.
ventral furrow distinct, close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–10 depressed. Parapophyses present from vertebrae
7 to 14. Pleural ribs on vertebrae 2 to 13. First anal in
pterygophore elongated, but not reaching tip of last
precaudal parapophysis.
Male copulatory organ (Fig. 7B–D). Two pairs of
large pseudoclaspers, outer pair only about 50% larger
than inner; outer pseudoclasper broad wing-shaped, with
short backward-directed tip; inner pseudoclasper pad
shaped with expanded tip, anteriorly broadly joined to
outer pseudoclasper. Penis straight, slightly longer than
outer pseudoclaspers.
Colour. live colour known from a freshly caught
specimen (WAM 31096-021, Fig. 6A), which shows a violet
body colour, lighter ventrally and darker dorsally. The
vertical ins are bright translucent dark yellow. Preserved
colour medium brown.
Comparison. For comparison of Beaglichthys
macrophthalmus with B. bleekeri and B. larsonae see
above. From the co-occurring Brosmolus longicaudus it
differs in the lower dorsal and anal in ray counts (111–113
and 83 vs 124–129 and 90–94), the presence of two pairs
of pseudoclaspers (vs one pair) and the separate opening of
the irst and second lower preopercular pores (vs fused).
Distribution. Beaglichthys macrophthalmus is known
from few specimens from locations along the northern
Australian coast and one location from the southern coast
of New Guinea (Fig. 1).
45
W. Schwarzhans and P. r. Møller
Fig. 7. Beaglichthys macrophthalmus Machida, 1993. WAM P.28155-019, male, 96+ mm Sl. A, lateral view of head; B, view of left
pseudoclasper from ventral; C, view of left pseudoclasper from inside; D, inclined lateral view of male copulatory organ; E, median
view of right otolith.
origin 23.3–24.3 % Sl); precaudal vertebrae 14–15, total
vertebrae 56–59, dorsal in rays 124–129, anal in rays
90–94, pectoral in rays 23–24, branchiostegal rays 7, v in
D 2.4–2.5; head with scale patch on cheek only, no scales
on operculum; no upper preopercular pore; irst and second
lower preopercular pore with separate opening; otolith
elongate (otolith length to height 2.1), its sulcus straight,
short (otolith length to sulcus length 2.1), colliculi fused;
maxilla slightly expanded posterio-ventrally; anterior anal
in pterygiophore variable in length.
Remarks. Brosmolus was originally described by
Machida (1993) as a monospeciic genus and deined by
Brosmolus Machida, 1993
(Tables 1, 2, 6)
Brosmolus Machida, 1993: 281 (ty pe species
B. longicaudus Machida, 1993, by monotypy); Nielsen
and Cohen in Nielsen et al. 1999: 119.
Diagnosis. A genus of the Dinematichthyini with the
following combination of characters: Anterior nostril
placed low on snout; male copulatory organ with single,
simple and lap-like, small pair of (outer) pseudoclaspers;
probably a large species (> 59 mm Sl, as indicated from
a pre-adult male), very slender (body depth at origin
of anal in 10.9–11.7 % Sl); base ventral in to anal in
46
Dinematichthyine ishes of the Indo-west Paciic III
the following combination of characters: preanal length
42 % Sl; head and body covered with thin, transparent
skin; cheek scaly; developed rakers on irst gill arch
4; caudal in rays 16; precaudal vertebrae 15. As with
Beaglichthys (see above), these are all widespread
characters found in the Dinematichthyini and most of
them are not of particular diagnostic value. The preanal
length is towards the lower end of variation observed in
many dinematichthyine genera. The number of precaudal
vertebrae ranges from 14 to 15, resulting in an overlap
with Beaglichthys. In fact the respective type species
of Brosmolus and Beaglichthys are very similar, but
for the purpose of this review we have refrained from
synonymising both genera until the nature and distinction
of the two established genera can be analyzed from a
broader base of specimens than currently available.
Comparison. Brosmolus appears to be closely related
to Beaglichthys Machida, 1993, from which it differs in
being even more elongate with higher associated numbers
of dorsal and anal in rays (> 123 and > 89 vs < 114
and <84 respectively). The other main difference is the
presence of only one pair of pseudoclaspers vs two pairs
in Beaglichthys. In the case of Brosmolus, however, minute
pseudoclaspers are known from the single male holotype,
which appears to be pre-adult. Without the difference in
pseudoclaspers, we would suggest synonymising both
genera, in which case Brosmolus would gain priority.
Brosmolus, Beaglichthys, Dactylosurculus gen. nov.
and Zephyrichthys gen. nov. are unique amongst the
dinematichthyine genera of this part of the review in
exhibiting separate openings of the irst and the second
lower preopercular pores, a character otherwise only found
in the genera Dermatopsoides Smith, 1947, and Dipulus
Waite, 1905, which however differ in many signiicant
other characters (see Møller and Schwarzhans 2006).
Distribution. Brosmolus is known from few specimens
from northern Australia, from the Kimberleys to the Gulf
of Carpentaria.
Brosmolus longicaudus Machida, 1993
(Figs 1, 8, 9; Tables 2, 6)
Brosmolus longicaudus Machida, 1993: 282; Nielsen
et al. 1999: 120.
Material examined. (3 specimens, 37–61 mm Sl).
hOlOTyPe – NTM S.10623-001, male, 61 mm Sl,
11°50’S, 130°05’e, Camerons Beach, Shoal Bay, Beagle
Table 6. Meristic and morphometric characters of Brosmolus
longicaudus Machida, 1993.
holotype
ntM
10623001
61
Standard length in mm
Meristic characters
Dorsal in rays
129
Caudal inrays
16
Anal in rays
94
Pectoral in rays
24
Precaudal vertebrae
15
Caudal vertebrae
44
Total vertebrae
59
rakers on anterior gill arch
16
Pseudobranchial ilaments
2
D/v
6
d/a
40
v/A
18
Morphometric characters in % of SL
head length
21.5
head width
10.8
head height
11.0
Snout length
4.4
upper jaw length
10.5
Diameter of pigmented eye
2.2
Diameter of pupil
1.5
Interorbital width
4.8
Posterior maxilla height
2.5
Postorbital length
15.3
Preanal length
42.2
Predorsal length
24.4
Body depth at origin of anal in
10.9
Pectoral in length
12.9
Pectoral in base height
4.1
ventral in length
Base ventral in – anal in origin 24.3
holotype +
2 non-types
n
Mean (range)
47.0 (37-61)
3
126.0 (124-129)
16
92.0 (90-94)
23.7 (23-24)
14.7 (14-15)
43.0 (42-44)
57.3 (56-59)
15.7 (15-16)
2
6
39.7 (37-42)
17.3 (16-18)
3
1
3
3
3
3
3
3
3
3
3
3
23.4 (21.5-24.6)
10.3 (9.5-10.8)
12.2 (11.0-13.6)
4.7 (4.4-4.9)
11.2 (10.5-11.9)
2.3 (2.2-2.5)
1.3 (1.2-1.5)
4.6 (4.3-4.8)
2.7 (2.5-2.9)
16.0 (15.3-17.3)
41.9 (41.2-42.3)
25.8 (24.4-27.3)
11.4 (10.9-11.7)
13.0 (12.4-13.9)
4.5 (4.1-4.9)
18.1
23.8 (23.3-24.3)
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
1
3
Gulf, Northern Territory, Australia, depth unknown, coll.
N. T. Fisheries, 13 March 1974.
Additional specimens. AMS I.6905, female, 37 mm
Sl, 17°S, 139°e, off Sweers Island, Gulf of Carpentaria,
Queensland, Australia; AMS I.33461-034, female, 43 mm
Sl, 14°09’S, 126°38’e, north of Kalumburu, Kimberleys,
Western Australia.
Diagnosis. See generic diagnosis.
Description (Figs 8, 9).The principal meristic and
morphometric characters are shown in Table 6. Body
Fig. 8. Brosmolus longicaudus Machida, 1993. AMS I 33461-034, female, 43 mm Sl.
47
W. Schwarzhans and P. r. Møller
Fig. 9. Brosmolus longicaudus Machida, 1993. holotype, NTM S.10623-001. A, lateral view of head; B, ventral view of head; C, view of
left pseudoclasper from inside; D, inclined lateral view of male copulatory organ, holotype; E, median view of right otolith.
and head very slender; single male (holotype) of about
60 mm Sl probably pre-adult. head with scale patch
on cheek containing seven vertical rows of scales on the
upper part and three on the lower part (absent in 37 mm
Sl specimen). horizontal diameter of scales on body
about 0.9 % Sl, in 25 horizontal rows (In a 43 mm Sl
female). Maxillary ending far behind eyes, dorsal margin
of maxillary covered by upper lip dermal lobe, posterior
end slightly expanded, angular. Anterior nostril positioned
low, 1/6 the distance from upper lip to aggregate distance
to anterior margin of eye. Posterior nostril small, about
1/5 the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 15–16 (16) rakers, thereof 4 elongate
rakers. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 9A, B). Supraorbital pores
2. 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).
Preopercular pores: 3 lower, irst and second with separate
opening; third non-tubular; no upper preopercular pores.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of a 43 mm Sl non-type). Premaxilla with
up to four outer rows of granular teeth and one inner row
of larger teeth. Anteriormost teeth in inner row up to 1/3
diameter of pupil. vomer horseshoe-shaped, with two
rows of equally sized teeth up to 1/4 diameter of pupil.
Palatine teeth in two rows 1/4 diameter of pupil. Dentary
with four outer rows of granular teeth and one inner row of
larger teeth anteriorly, merging into one row of larger teeth
posteriorly, up to about size of 1/2 of pupil diameter.
Otolith (Fig. 9e). elongate in shape, length to height 2.1
(59 mm Sl) and moderately thin (otolith height to otolith
thickness about 2.5). Anterior tip moderately rounded,
posterior tip broadly expanded. Dorsal rim gently curved
anteriorly and posteriorly, slightly concave at its middle
part; ventral rim gently and evenly curved. Inner and
outer face moderately convex, both smooth. Sulcus short,
otolith length to sulcus length 2.1. Sulcus positioned at
centre of inner face, with fused colliculi, slightly inclined
to otolith axis. ventral furrow distinct, close to ventral
rim of otolith.
Axial skeleton. Neural spine of vertebra 4 inclined and
5–8 (9) depressed. Parapophyses present from vertebrae
48
Dinematichthyine ishes of the Indo-west Paciic III
7 to 15. Pleural ribs on vertebrae 2 to 14. First anal in
pterygophore usually not very elongated and not reaching
tip of last precaudal parapophysis.
Male copulatory organ (Fig. 9C, D). Single pair of
small, triangular, lap-like (outer) pseudoclaspers. Penis
curved, slightly longer than pseudoclaspers. The copulatory
organ of the single known male gives the impression of
being pre-adult by its small size and fragility.
Colour. live colour unknown. Preserved colour
medium greenish brown to brown.
Comparison. See comparison between Brosmolus
and other genera.
Distribution. See to genus Brosmolus (Fig. 1).
Dactylosurculus gen. nov.
(Tables 1, 2, 7)
Ogilbya (non Jordan and evermann in evermann and
Kendall, 1898) Kuiter 2000: 61 (photo).
Type species: Dactylosurculus gomoni sp. nov. Gender
masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed low
on snout; male copulatory organ with three pairs of
pseudoclaspers, all joined at base in narrow stalked stem,
outer pseudoclasper wing-shaped, v-shaped lap-like inner
pseudoclasper anteriorly joined to outer pseudoclasper,
short, second inner pseudoclasper inserted between the
two, intermediate in length, inger-like in shape, with
supporter; moderately large species, reaching about 70
mm Sl; precaudal vertebrae 13–14, total vertebrae 45–50;
pectoral in rays 22–26; 8–9 branchiostegal rays; head
with small scale patch on upper cheek only, no scales on
operculum; upper preopercular pore present; additional
small pore below eye; otolith elongate (otolith length to
height 2.1–2.3), sulcus inclined, colliculi fused; maxilla
expanded posterio-ventrally.
Comparison. Dactylosurculus is readily recognised by
a number of unique characters or characters unique in their
combination. First of all there is the peculiar arrangement
of the two inner pseudoclaspers, whereby one of them
could in fact could be called “middle” pseudoclasper.
The only other genus with two inner pseudoclaspers
is Ogilbichthys from America, but there, the two inner
pseudoclaspers are positioned along the axis, resulting
in an anterior and a posterior inner pseudoclasper. Other
signiicant characters are the eight branchiostegal rays
(also in some species of Beaglichthys) and the small extra
pore below the eye, elsewhere observed in a few species
of the genus Ogilbia from America.
Further distinguishing characters are the combination
of the presence of an upper preopercular pore, the high
number of precaudal and total vertebrae (13–14 and
45–50), the low degree of head squamation and the sulcus
of the otolith with fused colliculi.
Species. The genus is monotypic.
Etymology. Combined from dactylus (latin, from
Greek dactylos = inger) and the surculus (latin = sucker
of a grapevine tendril), referring to the functional analogy
with the pseudoclaspers, in this case also referring to the
speciic shape of the ‘middle’ pseudoclasper.
Dactylosurculus gomoni sp. nov.
(Figs 10–12; Tables 2, 7)
Ogilbya sp. Kuiter 2000: 61 (photo).
Material examined. (75 specimens, 22–74 mm Sl).
hOlOTyPe – WAM P.28290-002, male, 60 mm Sl,
32°16’S, 126°02’e, off Cocklebiddy, Western Australia,
Great Australian Bight, depth 2–3 m, coll. J. B. hutchins
et al., 9 April 1984. ParatyPeS – AMS I.17614-017,
1 male, 55 mm Sl, 3 females, 55–66 mm Sl, Spencer
Gulf, South Australia, depth 2 m, coll. D. hoese et al.,
25 Dec. 1973; AMS I.17614-033, 1 male, 52 mm Sl, 1
female, 42 1 juvenile, 26 mm Sl, 33°49’S, 137°40’e, off
Tickera, Spencer Gulf, South Australia, depth 2 m, coll.
Table 7. Meristic and morphometric characters of Dactylosurculus
gomoni sp. nov.
holotype
WaM
28290002
60
Standard length in mm
Meristic characters
Dorsal in rays
100
Caudal inrays
14
Anal in rays
74
Pectoral in rays
25
Precaudal vertebrae
13
Caudal vertebrae
35
Total vertebrae
48
rakers on anterior gill arch
13
Pseudobranchial ilaments
0
D/v
6
d/a
33
v/A
16
Morphometric characters in % of SL
head length
24.0
head width
12.2
head height
14.5
Snout length
5.5
upper jaw length
12.4
Diameter of pigmented eye
2.0
Diameter of pupil
1.2
Interorbital width
6.8
Posterior maxilla height
4.2
Postorbital length
16.8
Preanal length
46.9
Predorsal length
28.7
Body depth at origin of anal in
15.3
Pectoral in length
4.2
Pectoral in base height
12.0
ventral in length
6.0
Base ventral in – anal in origin 28.4
49
holotype +
74 paratypes
n
Mean (range)
22-74 (50.6)
75
101.3 (92-109)
14 (14-15)
74.0 (70-81)
24.1 (22-26)
13.1 (13-14)
34.0 (32-36)
47.0 (45-50)
13.2 (9-16)
1.0 (0-3)
6.2 (6-7)
31.3 (26-35)
15.5 (15-16)
39
11
39
21
43
43
43
34
34
39
39
39
24.5 (22.6-26.4)
12.2 (11.4-14.1)
14.4 (13.4-15.1)
5.8 (4.7-6.6)
12.1 (11.2-12.9)
2.2 (1.6-2.5)
1.4 (1.1-1.7)
6.5 (5.7-6.8)
4.1 (3.6-4.6)
17.2 (16.0-18.2)
47.3 (45.5-49.2)
29.5 (27.0-32.1)
14.4 (13.2-15.3)
12.9 (11.6-13.8)
5.8 (5.0-6.3)
18.2 (15.0-20.6)
29.2 (27.5-30.5)
18
9
9
9
9
18
12
9
9
9
9
16
9
9
9
16
9
W. Schwarzhans and P. r. Møller
Fig. 10. Sample sites of Dactylosurculus gomoni sp. nov., Lapitaichthys frickei sp. nov., Monothrix polylepis and Zephyrichthys barryi
sp. nov. One symbol may represent several samples.
hutchins, 30 June 1982; WAM P. 28290-015, 1 male, 54
mm Sl, 3 females, 55–66 mm Sl, same data as holotype;
WAM P.28296-015, 1 male, 34 mm Sl, 3 females, 49–66
mm Sl, 34°08’S, 122°16’e, Mondrian Island, Western
Australia, depth 5–6 m, coll. J. B. hutchins et al., 13
April 1984; WAM P.28513-004, 4 males, 50–55 mm Sl,
5 females, 51–63 mm Sl, 33°54’S, 122°37’e, Western
Australia, depth 8 m, coll. C. Bryce, 7 March 1985; WAM
P.28519-007, 1 male, 37 mm Sl, 3 females, 36–64 mm Sl,
33°32’S, 115°01’e, Cape Jervis, Fleurieu Peninsula, South
Australia, depth 6–7 m, coll. J. B. hutchins, 15 April 1986;
ZMuC P 771634-35, 1 male, 50 mm Sl, 1 female, 60 mm
Sl, same data as WAM P.28296-015.
Diagnosis. See generic diagnosis.
Description (Figs 11, 12). The principal meristic and
morphometric characters are shown in Table 7. Body
slender, about same height from neck to position at about
half of the anal in; mature at about 35 to 40 mm Sl.
head with small scale patch on upper cheek with 4 to
10 individual, small, non-imbricate scales. horizontal
diameter of scales on body about 1.2 % Sl, in 22 horizontal
rows. Maxillary ending far behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded, angular. Anterior nostril positioned moderately
low, 1/5 the distance from upper lip to aggregate distance
to anterior margin of eye. Posterior nostril small, about 1/4
the size of eye. Opercular spine with short free tip, pointed.
D. hoese et al., 25 Dec. 1973; AMS I.20216-010, 34°02’S,
122°14’e, off rob Island, Western Australia, coll. A.
Kuiter, 20 March 1978; NMv A3421, 1 female, 66 mm Sl,
38°19’S, 144°43’e, off Portsea, Port Phillip Bay, victoria,
Australia, depth 10 m, coll. r. h. Kuiter, 2 March 1984;
NMv A17793, 3 males, 54–57 mm Sl, 10 females, 43–66
mm Sl, 6 juveniles, 22–26 mm Sl, Mornington Peninsula,
victoria, Australia, depth 5 m, coll. M. lockett and r.
Ickeringill, 6 Feb. 1996; NMv A17794, 3 females, 65–74
mm Sl, 2 juveniles, 26–27 mm Sl, 38°09’S, 145°05’e, off
Frankston, Port Phillip Bay, victoria, Australia, depth 2
m, coll. M. F. Gomon et al., 6 Feb. 1996; NMv A17795, 1
female, 56 mm Sl, 37°59’S, 145°02’e, off rickets Point,
Port Phillip Bay, victoria, Australia, depth 3 m, coll. M.
F. Gomon et al., 7 Feb. 1996; NMv A17796, 1 female, 49
mm Sl, 37°59’S, 145°02’e, off rickets Point, Port Phillip
Bay, victoria, Australia, depth 3 m, coll. M. lockett and
r. Ickeringill, 9 Dec. 1995; NMv A18141, 1 male, 52 mm
Sl, 38°07’S, 144°41’e, off Portarlington, Port Phillip Bay,
victoria, Australia, depth 2 m, coll. M. F. Gomon et al.,
29 Oct. 1996; WAM P.4790, 1 male 51 mm Sl, 1 female
60 mm Sl, Cape Jarvis, South Australia, coll. T. D. Scott,
6 Nov. 1956; WAM P.27138-001, 3 females, 36–62 mm
Sl, 34°44’S, 135°52’e, off Port lincoln, South Australia,
coll. J. B. hutchins, 8 April 1981; WAM P.27643-005, 4
males, 33–45 mm Sl, 6 females, 38–57 mm Sl, 33°32’S,
115°02’e, Cape Naturaliste, Western Australia, coll. J. B.
50
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 11. Dactylosurculus gomoni sp. nov. A, fresh dead, WAM P.28296-015, female, 49 mm Sl; B, WAM P.28290-002, holotype, 60 mm Sl.
Anterior gill arch with 9–16 (13) rakers, thereof 0–2 (0)
elongate rakers (usually 1). Pseudobranchial ilaments
0–3 (0), usually 1.
Head sensory pores (Fig. 12A, B). Supraorbital pores
3. Infraorbital pores 7 (3 anterior, 3 posterior and 1 small
pore below eye): three posterior pores about 1/3 the size
of three anterior pores. Mandibular pores 6 (3 anterior and
3 posterior); irst anterior mandibular pore with cirrus.
Preopercular pores: 3 lower, irst and second with separate
opening; third tubular; tubular upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to four
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 2/3 diameter
of pupil. vomer horseshoe-shaped, with two rows of
equally sized teeth up to 1/4 diameter of pupil. Palatine
with an outer row of small teeth up to 1/5 diameter of
pupil and inner row of larger teeth up to 1/4 diameter of
pupil. Dentary with four outer rows of granular teeth and
one inner row of larger teeth anteriorly, merging into one
row of larger teeth posteriorly, up to about size of 2/3 of
pupil diameter.
Otolith (Fig. 12F, G). elongate in shape, length to height
2.1–2.3 (24–74 mm Sl) and thin (otolith height to otolith
thickness 2.2–2.5). Anterior tip sharply pointed, posterior
tip expanded, massive. Dorsal rim with broad predorsal
angle and expanded, sharply pointed postdorsal angle,
long section in between straight, marked concavity above
anterior tip and sometimes also behind postdorsal angle;
ventral rim gently and regularly curved, deepest at about
its middle. Inner face convex, outer face lat to slightly
concave, both smooth. Otolith length to sulcus length
2.0–2.1. Sulcus positioned anterior of centre of inner face,
slightly inclined, single undivided colliculum, its ventral
margin more convex than dorsal margin. ventral furrow
long, distinct, broad, moderately close to ventral rim of
otolith, slightly turning upwards towards its tips. Dorsal
depression long, deep. A low degree of sexual dimorphism
seems to be relected in the way that otoliths of males
(Fig. 12F) tend to be slightly more compressed and with a
slightly wider sulcus than those of females (Fig. 12G).
Otoliths described by Nolf (1980) and Schwarzhans
(1981) as Dermatopsis macrodon (plate 13, ig. 13 and
ig. 145 respectively) and Dermatopsis multiradiatus
(plate 13, ig. 14 and ig. 146 respectively) likely represent
Dactylosurculus gomoni.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–10 depressed. Parapophyses present from vertebrae
6 (7) to 13. Pleural ribs on vertebrae 2 to 12. First anal
in pterygophore elongated, reaching tip of last precaudal
parapophysis in males but not in females.
Male copulatory organ (Fig. 12 C–e). Three pairs of
pseudoclaspers, all joined at base in narrow stalked stem,
outer pseudoclasper wing shaped, v-shaped lap-like inner
pseudoclasper anteriorly joined to outer pseudoclasper,
short, second inner pseudoclasper inserted between the
two and intermediate in length, inger-like in shape, with
supporter; isthmus narrow. Penis curved, slightly longer
than pseudoclasper, with broad basis.
Colour. live colour uniformly median brown (WAM
28296-015, Fig. 11A). vertical fins translucent, with
yellow-brownish base. Preserved colour mostly light
brown, rarely medium brown.
51
W. Schwarzhans and P. r. Møller
Fig. 12. Dactylosurculus gomoni sp. nov. A, lateral view of head, WAM P.27643-005, male, 45 mm Sl; B, ventral view of head, WAM
P.27643-005, male, 45 mm Sl; C, view of left pseudoclasper from inside, WAM P.28290-015); D, view of left pseudoclasper from inside,
holotype; E, inclined lateral view of male copulatory organ, holotype; F, median view of right otolith, NMv A 17793, male, 59 mm Sl;
G, median view of right otolith, NMv A17794, female, 66 mm Sl.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed low on
snout; male copulatory organ with single pair of (outer)
pseudoclaspers with two equally long supporters; Sl up
to 66 mm Sl, females pregnant at 30 mm Sl; precaudal
vertebrae 11–13 (mostly 12), total vertebrae 40–45,
branchiostegal rays 6–7; head with scale patch on cheek
only, no scales on operculum; upper preopercular pore
present; otolith elongate (otolith length to height 2.1–2.3),
sulcus not inclined or inclined (0–10°), colliculi fused;
maxilla expanded postero-ventrally; anterior anal in
pterygiophore long.
Comparison. See comparison between Dactylosurculus
and other genera.
Distribution. Dactylosurculus gomoni is widely
distributed along the southern shores of Australia from
Cape Naturaliste in the west to the Port Phillip Bay in the
east (Fig. 10).
Etymology. In honour of Martin F. Gomon, Melbourne,
NMv, and his many contributions to the knowledge of the
ish fauna of Australia.
Didymothallus gen. nov.
(Tables 1, 2, 8–10)
Type species: Didymothallus criniceps sp. nov. Gender
masculine.
52
Dinematichthyine ishes of the Indo-west Paciic III
Comparison. Didymothallus is best characterised by
its single (outer) pair of pseudoclaspers with two slender,
almost equally long supporters, a character otherwise
only found in two American Dinematichthyini, namely
Gunterichthys Dawson, 1966, and a single species of
the genus Ogilbia Jordan and evermann, 1898 (Ogilbia
davidsmithi Møller, Schwarzhans and Nielsen, 2005)
in which the anterior supporter however is only about
half the length of the posterior supporter. Didymothallus
differs from Gunterichthys in the free opercular spine (vs
hidden) and the posterior-ventral expansion of the maxilla
(vs shallow). From Ogilbia it differs further in the lack of
inner pseudoclaspers and the fused colliculi of the sulcus
(vs separated).
In the absence of males, female specimens of
Didymothallus can be confused in the Indo-west Paciic
with other genera with which they share the presence of
the upper preopercular pore and fused colliculi in the
sulcus of the otolith, such as Diancistrus Ogilby, 1899,
and Monothrix Ogilby, 1897, which partly overlap with
Didymothallus in their geographic distribution or occur
in adjacent areas, Brotulinella Schwarzhans, Møller
and Nielsen, 2005, endemic to the northern Philippines
and Taiwan and Majungaichthys gen. nov., endemic to
Madagascar. Fishes of Didymothallus are usually markedly
more slender than those of Diancistrus (body depth at
origin of anal in 12.0–16.0 vs 14.5–23.0). Monothrix is
distinguished by high vertebrae and dorsal in ray counts,
which only slightly overlap with those of Didymothallus
(total vertebrae 45–47 vs 40–45, dorsal in rays 93–104 vs
69–97). Monothrix is a genus with only one pair of (outer)
pseudoclaspers and with one supporter, not two supporters
as is the case in Didymothallus.
Biology. As already stated in the introduction, many
Dinematichthyini are noted for their narrow distribution
ranges, which might be related to their exceptionally
low fecundity. In the case of Didymothallus gen. nov.,
which is also known from a rather restricted geographic
distribution along Australian and New Caledonian shores,
exceptionally low numbers (1 to 3, usually 2) of (large)
embryos have been found in gravid females of D. mizolepis
(Fig. 16C). Another species, D. criniceps, is remarkable for
bearing eggs in the body cavity (Fig. 13B), an indication of
a (secondary) oviparous reproduction and, if substantiated
by life observations, the irst known in the Bythitoidei.
Species. Two species from the northern Australian
shelf, D. criniceps sp. nov. from the Great Barrier reef and
D. mizolepis (Günther, 1867) from north-western Australia
to Cape york and one species from New Caledonia, D.
pruvosti sp. nov.
Etymology. Combined from the Greek didymos (=
double, twofold) and thallus (= branch in plants), referring
to the two supporters of nearly equal length in the single
pair of pseudoclaspers.
Didymothallus criniceps sp. nov.
(Figs 13–15; Tables 2, 8)
Material examined. (30 specimens, 19–60 mm Sl).
hOlOTyPe – AMS IA.2611, male, 48 mm Sl, 23°S,
152°e, One Tree Island, Great Barrier reef, Queensland,
Australia, coll. G. P. Whitley, date unknown. PArATyPeS
– AMS IA. 2611-002, 1 male, 42 mm Sl, 1 female, 53 mm
Sl, same data as holotype; AMS IA.6793, 1 female, 47
mm Sl, 3 juveniles, 31–32 mm Sl, 20°27’S, 149°02’e,
lindeman Island, Great Barrier reef, Queensland,
Australia, coll. G. P. Whitley, date unknown; AMS I.
20063-004, 2 males, 31–35 mm Sl, 1 female, 35 mm Sl,
2 juveniles, 19–21 mm Sl, 23°30’S, 152°05’e, One Tree
Island, Great Barrier reef, Queensland, Australia, coll.
F. Talbot and party, 24 Nov. 1969; AMS I.20205-060, 1
male, 54 mm Sl, 23°30’S, 152°05’e, One Tree Island,
Great Barrier reef, Queensland, Australia, coll. F. Talbot
and party, 27 Sept. 1968; AMS I.27746-006, 2 males,
45 mm Sl, 20°27’S, 149°02’e, lindeman Island, Great
Table 8. Meristic and morphometric characters of Didymothallus
criniceps sp. nov.
holotype
aMS
IA.2611
Standard length in mm
48
Meristic characters
Dorsal in rays
71
Caudal inrays
15
Anal in rays
53
Pectoral in rays
18
Precaudal vertebrae
12
Caudal vertebrae
31
Total vertebrae
43
rakers on anterior gill arch
14
Pseudobranchial ilaments
2
D/v
6
d/a
23
v/A
15
Morphometric characters in % of SL
head length
23.9
head width
13.4
head height
14.6
Snout length
5.0
upper jaw length
11.8
Diameter of pigmented eye
1.8
Diameter of pupil
1.1
Interorbital width
6.4
Posterior maxilla height
3.4
Postorbital length
17.2
Preanal length
50.3
Predorsal length
31.2
Body depth at origin of anal in
16.5
Pectoral in length
13.5
Pectoral in base height
6.0
ventral in length
Base ventral in – anal in origin 31.8
53
holotype +
29 paratypes
n
Mean (range)
39.7 (19-60)
30
73.1 (69-77)
14.6 (13-16)
55.0 (52-61)
19.2 (18-22)
12.0
31.3 (30-33)
43.2 (42-45)
14.5 (12-16)
2
6.6 (6-7)
23.8 (23-26)
15.2 (15-16)
26
19
26
19
25
26
26
23
22
26
26
26
26.1 (23.6-28.4)
12.0 (10.5-14.0)
14.8 (14.2-15.5)
5.2 (3.9-5.9)
11.9 (11.2-13.5)
2.3 (1.8-2.8)
1.3 (1.1-1.6)
6.0 (5.4-6.9)
3.4 (3.0-3.9)
18.3 (17.2-19.6)
49.9 (44.0-53.0)
32.1 (30.0-35.0)
14.2 (12.5-16.8)
13.9 (9.6-16.9)
5.3 (4.4-6.0)
19.8 (16.8-25.8)
32.9 (29.4-35.6)
17
13
13
13
13
17
14
13
13
13
13
17
13
12
13
20
12
W. Schwarzhans and P. r. Møller
Fig. 13. Didymothallus criniceps sp. nov. A, AMS IA.2611, holotype, 48 mm Sl; B, x-ray, uSNM 366587, female, 46 mm Sl.
Barrier reef, Queensland, Australia, M. Ward, 1934; AMS
I. 34311-010, 1 female, 58 mm Sl, 22°14’S, 150°19’e,
Cannibal Group, Queensland, Australia, coll. AMS party,
15 Sept. 1993; uSNM 366587, 4 males, 37–50 mm Sl, 3
females 45–46 mm Sl, One Tree Island, Great Barrier
reef, Queensland, Australia, depth 0–1 m, coll. v.G.
Springer et al., 27 Nov. 1966; uSNM 374187, 1 male, 36
mm Sl, One Tree Island, Great Barrier reef, Queensland,
Australia, depth 0–5 m, coll. v.G. Springer et al., 25 Nov.
1966; uSNM 374202, 2 females, 32 mm Sl, One Tree
Island, Great Barrier reef, Queensland, Australia, depth
0–2 m, coll. v.G. Springer et al., 20 Nov. 1966; ZMuC
P771629-30, 1 male, 40 mm Sl, 1 female, 60 mm Sl, same
data as AMS IA.6793.
Additional specimens. AMS I. 19441-021, 1 female,
46 mm Sl, 14°44’S, 145°31’e, lizard Island, Great Barrier
reef, Queensland, Australia.
Tentatively assigned specimens: AMS I.21315-001,
1 male, 29 mm Sl, 14°06’S, 121°56’e, Scott reef, off
Western Australia.
Diagnosis. vertebrae 12+30-33=42-45, dorsal in rays
69–77, anal in rays 52–61, D/A 23–26 v in D 1.9–2.1;
head with multiple small cirri on occiput; single (outer)
pseudoclasper wing-shaped with two slender supporters,
slightly bent, particularly the posterior one, of exact
same length; cheeks with 3–5 rows of scales on upper
part and 2 rows on lower part; otolith with moderately
pointed anterior tip and pronounced postdorsal angle,
with undivided short sulcus, its centre anterior to centre
of otolith, inclined with 5 to 10°, otolith length to otolith
height = 2.2, otolith length to sulcus length = 2.2–2.4.
Description (Figs 13, 14). The principal meristic
and morphometric characters are shown in Table 8.
Body slender; mature at about 35 mm Sl. head with
scale patch on cheek containing 3–5 (3) vertical rows of
scales on upper part and 2 vertical rows on lower part.
horizontal diameter of scales on body about 1.4 % Sl, in
23 horizontal rows. Occiput with many thin, short, hairlike cirri (Fig. 14A, C). Maxillary ending far behind eyes,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, angular. Anterior nostril
positioned low, 1/6 the distance from upper lip to aggregate
distance to anterior margin of eye. Posterior nostril small,
about 1/4 the size of eye. Opercular spine with long free tip,
pointed. Anterior gill arch with 12–16 (14) rakers, thereof
3 elongated. Pseudobranchial ilaments always 2.
Head sensory pores (Fig. 14A–C). 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); irst
anterior mandibular pore without cirrus. Preopercular
pores: 3 lower, irst and second with joined opening;
third non-tubular; tubular upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to three
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 2/3 diameter
of pupil. vomer horseshoe-shaped, with three rows of
equally sized teeth up to 1/4 diameter of pupil. Palatine
with two rows of teeth up to 1/4 diameter of pupil. Dentary
with four outer rows of granular teeth and one inner row of
54
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 14. Didymothallus criniceps sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, dorsal view of head,
holotype; D, ventral view of male copulatory organ with hood not bent forward, uSNM 366587, 49 mm Sl; E, ventral view of male
copulatory organ with hood bent forward, uSNM 366587, 49 mm Sl; F, inclined lateral view of male copulatory organ, uSNM 366587,
49 mm Sl; G, view of left pseudoclasper from inside, holotype; H, view of left pseudoclasper from inside, uSNM 366587, 49 mm Sl;
I, view of left pseudoclasper from inside, uSNM 366587, 44 mm Sl; J, (tentatively assigned specimen) view of left pseudoclasper from
inside, AMS I.21315-001, 29 mm Sl; K, median view of right otolith, uSNM 366587, male, 37 mm Sl; L, median view of right otolith,
uSNM 366587, female, 44 mm Sl.
55
W. Schwarzhans and P. r. Møller
Biology. Several female specimens were observed
to have 9 to 18 eggs in the body cavity (Fig. 13 B). Two
females of 44 mm Sl (uSNM 366587) were opened and
found to contain orange-coloured eggs measuring 1.4–1.7
mm in diameter. No female specimens have been reported
with embryos inside. This observation could indicate
that D. criniceps is oviparous rather than viviparous, an
assumed diagnostic character for all Bythitoidei. however,
this irst indication will need further substantiation by
in-life observation.
Etymology. From crinis (latin = hair) and cephalos
(Greek = head), refering to the many hair-like cirri on the
occiput.
larger teeth anteriorly, merging into one row of larger teeth
posteriorly, up to about size of 3/4 of pupil diameter.
Otolith (Fig. 14K–l). elongate in shape, length to
height 2.2 (36–60 mm Sl) and moderately thin (otolith
height to otolith thickness about 2.0). Anterior tip pointed,
posterior rim expanded. Dorsal rim with rounded pre- and
postdorsal angles, the latter more pronounced, section in
between straight, concave towards anterior and posterior
tips; ventral rim gently and regularly curved. Inner and
outer faces moderately convex, smooth. Otolith length to
sulcus length 2.2–2.4. Sulcus positioned slightly towards
anterior, with fused colliculi, inclined 5–10° towards
otolith axis. ventral furrow faint, close to ventral rim of
otolith, anteriorly turned upward to meet tip of sulcus. A
slight sexual dimorphism is observed in that otoliths of
females show a more rounded anterior tip while those from
males are sharply pointed.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 (9) depressed. Parapophyses present from
vertebrae 6 to 12. Pleural ribs on vertebrae 2 to 11. First
anal fin pterygophore elongated, reaching tip of last
precaudal parapophysis in males but not in females.
Male copulatory organ (Fig. 14D–J). A single pair
of large (outer) pseudoclaspers, wing-shaped, slightly
bent outwards, base and tip similarly wide, with two
almost exactly equally long and thin supporters, slightly
bent, posterior supporter usually somewhat more so
than anterior. Penis slightly curved, slightly longer than
pseudoclaspers, pointed, with broad base.
Colour. live colour unknown. Preserved colour light
to medium brown, sometimes reddish brown.
Comparison. Didymothallus criniceps closely
resembles D. pruvosti sp. nov. from New Caledonia, with
which it probably forms an allopatric pair. It differs in the
posterior supporter of the pseudoclasper being slender
throughout (vs with expanded club-like tip), the higher D/A
(23–26 vs 20–22), the lower number of scale rows on the
cheeks (5–6 on upper and 2 on lower cheek vs 7–8 on upper
and 3–5 on lower cheeks) and details of the otolith outline
and otolith length to sulcus length (2.2–2.4 vs 2.6). Also,
D. pruvosti sp. nov. does not have the cirri on the occiput
as observed in D. criniceps. Didymothallus criniceps is
distinguished from D. mizolepis (Günther, 1867) in the
lower number of dorsal in rays (69–77 vs 78–94), the
lower v in D (1.9–2.1 vs 2.2–2.6), the equal length of the
supporters in the pseudoclasper (vs the anterior being
slightly shorter than the posterior), the curved penis (vs
straight), the short and inclined sulcus (otolith length to
sulcus length 2.2–2.4 vs 1.9–2.1 and not inclined) and
several aspects of the otolith outline.
Distribution. Didymothallus criniceps is known from
the Great Barrier reef of eastern Australia from about
14°S to 23°30’S (Fig. 15). A single pre-adult male recorded
from the offshore Scott reef off north-western Australia is
only tentatively placed in the species due to its geographic
remote location.
Didymothallus mizolepis (Günther, 1867)
(Figs 15–17; Tables 2, 9)
Dinematichthys mizolepis Günther, 1867: 66.
Monothrix mizolepis. – Cohen & Nielsen 1978: 60;
Paxton et al. 1989: 317; larson and Williams 1997: 349;
Nielsen et al. 1999: 134.
Material examined. (177 specimens, 14–66 mm Sl).
hOlOTyPe – BMNh 1867.5.13.17, female, 47 mm Sl,
Cape york, Queensland, Australia, purchased by Mr.
Damel, date unknown.
Additional specimens. AMS I.7792, 1 male, 48 mm
Sl and 1 female, 56 mm Sl, Melville Island, Northern
Territory, Australia; AMS I.17060-035, 1 male, 33 mm
Sl and 1 female, 44 mm Sl, 22°15’S, 114°15’e, exmouth
Gulf, Western Australia; AMS I.24676-044, 22 males, 28–
34 mm Sl, 31 females, 26–41 mm Sl, 17 juveniles, 18–26
mm Sl, 12°29’S, 130°53’e, Darwin harbour, Northern
Territory, Australia; AMS I. 24678-053, 1 male, 49 mm
Sl, 1 female, 50 mm Sl, 12°29’S, 130°53’e, Darwin
harbour, Northern Territory, Australia; AMS I.27743-002,
1 male, 39 mm Sl, Melville Island, Northern Territory,
Australia; BPBM 17405, 1 female, 45 mm Sl, 20°S, 116°e,
Kendrew Island, Dampier Archipelago, Western Australia;
MNhN 1890-0299, 2 females, 44–49 mm Sl, 10°37’S;
142°10’e, Thursday Island, Queensland, Australia ; NTM
S.12886-002, 1 female, 47 mm Sl, 3 juveniles, 17–24
mm Sl, 17°59’S, 122°11’e, Broome, Western Australia;
NTM S.14472-007, 3 females, 30–35 mm Sl, 12°04’S,
132°19’e, Cunningham Channel, Northern Territory,
Australia; QM 33829, 1 female, 50 mm Sl, 1 juvenile,
17 mm Sl, 17°08’S, 139°36’e, Sweers Island, Gulf of
Carpentaria, Queensland, Australia; ruSI 35938, 5 males,
27–35 mm Sl, 3 females, 30–33 mm Sl, Darwin harbour,
Northern Territory, Australia; SMNS 14557, 1 male, 44
mm Sl, 21°39’S, 115°08’e, Onslow, Western Australia;
SMNS 18421, 1 female, 56 mm Sl, 17°00’S, 122°e,
Gantheaume Bay, Broome, Western Australia; SMNS
18495, 1 female, 61 mm Sl, 12°19’S, 130°50’e, Darwin
harbour, Northern Territory, Australia; SMNS 18551, 1
female, 47 mm Sl, 12°19’S, 130°50’e, Darwin harbour,
Northern Territory, Australia; uSNM 327949, 4 males,
25–34 mm Sl, 9 females, 23–34 mm Sl, 1 juvenile, 15
56
Dinematichthyine ishes of the Indo-west Paciic III
mm Sl, 12°19’S, 130°53’e, Darwin harbour, Northern
Territory, Australia; WAM P.25111-033, 1 female, 51 mm
Sl, 20°00’S, 116°00’e, Dampier Archipelago, Western
Australia; WAM P.27967-038, 1 male, 37 mm Sl, 24°29’S,
113°25’e, off Beagle hill, Western Australia; WAM
P.27980-065, 1 male, 42 mm Sl, 20°26’S, 115°35’e,
Montebello Islands, Western Australia; WAM P.30307013, 1 female, 62 mm Sl, 1 juvenile, 21 mm Sl, 13°48’S,
125°47’e, Western Australia; WAM P.30849-010, 1 male,
33 mm Sl, 15°00’S, 125°00’e, Western Australia; WAM
P.30929-003, 1 male, 33 mm Sl, 16°16’S, 123°30’e,
hidden Island, Western Australia; WAM P.31013-010, 6
specimens, mm Sl, 22°06’S, 114°31’e, exmouth Gulf,
Western Australia; WAM P.31015-031, 1 female, 51 mm
Sl, 22°07’S, 114°29’e, exmouth Gulf, Western Australia;
WAM P.31078-031, 4 males, 33–47 mm Sl, 2 females,
57–62 mm Sl, 2 juveniles, 22–24 mm Sl, 13°45’S,
126°22’e, eclipse Islands, Western Australia; WAM
P.31085-032, 2 females, 43–55 mm Sl, 13°59’S, 126°20’e,
vansittart Bay, Western Australia; WAM P.31092-008, 1
male, 51 mm Sl, 5 females, 44–63 mm Sl, 3 juveniles,
22–23 mm Sl, 13°45’S, 126°45’e, Stewart Islands,
Western Australia; WAM P.31205-016, 2 males, 29 mm
Sl, 1 female, 32 mm Sl, 16°22’S, 123°03’e, Swan Island,
Western Australia; WAM P.31236-003, 3 females, 36–40
mm Sl, 15°32’S, 124°25’e, Camden Sound, Western
Australia; WAM P.31239-002, 1 male, 48 mm Sl, 14°46’S,
125°01’e, york Sound, Western Australia; WAM P.31243-
005, 3 specimens, 27–50 mm Sl, 14°00’S, 125°00’e,
Western Australia; WAM P.31250-031, 3 females, 42–46
mm Sl, 4 juveniles, 14–28 mm Sl, 15°17’S, 124°10’e,
Champagny Islands, Western Australia; WAM P.31251056, 2 males, 27–53 mm Sl, 3 females, 27–43 mm Sl,
15°55’S, 124°03’e, viney Island, Western Australia; WAM
P.31506-001, 1 male, 51 mm Sl, 1 female, 66 mm Sl,
20°28’S, 116°52’e, Dolphin Island, Western Australia;
WAM P.31516-006, 1 female, 47 mm Sl, 1 juvenile, 16
mm Sl, Dampier Shelf, Western Australia; ZMuC P
771636-37, 1 male, 40 mm Sl, 1 female, 60 mm Sl, same
data as WAM P.31092-008.
Remarks. The holotype is a female specimen from
Cape york, at the far end of the known distribution range
of the species. In fact, it represents the furthest record to
the east (together with two female specimens from the
Thursday Islands), the next being one from the southern
tip of the Gulf of Carpentaria. The meristics exclude it
from representing a specimen of D. criniceps (see below),
which has been recorded from the Great Barrier reef,
but somewhat further south of Cape york. eusurculus
pistillum gen. nov. sp. nov., which is described later, is in
good agreement in all relevant meristic and morphometric
values and is distributed throughout the range of both
Didymothallus species. The pseudoclasper pattern (single
outer pseudoclasper with two supporters vs two pairs of
pseudoclaspers, the inner sucker-disk-like) and the sulcus
morphology (fused colliculi vs separated) distinguish
Fig. 15. Sample sites of Didymothallus criniceps sp. nov., D. mizolepis and D. pruvosti sp. nov. One symbol may represent several
samples.
57
W. Schwarzhans and P. r. Møller
and thicker than anterior one; penis straight; cheeks with
4–6 rows of scales on upper part and 0–2 rows on lower
part; otolith with rounded anterior tip and with undivided
long sulcus, its centre anterior of centre of otolith, not
inclined, otolith length to otolith height = 2.2–2.3, otolith
length to sulcus length = 1.9–2.1.
Description (Figs 16, 17). The principal meristic and
morphometric characters are shown in Table 9. head and
body slender, deepest markedly behind head; a small
species, mature at about 30 mm Sl. head with scale
patch on cheek containing 4 to 6 vertical rows of scales on
upper part, lower part of cheek without scales or with 0–2
vertical rows (Fig. 17 A,C). horizontal diameter of scales
on body about 1.1 % Sl, in 20 horizontal rows. Maxillary
ending far behind eyes, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded, with
small knob anterior of largest expansion. Anterior nostril
positioned very low, 1/5–1/6 the distance from upper lip
to aggregate distance to anterior margin of eye. Posterior
nostril small, about 1/4 the size of eye. Opercular spine
with free tip, pointed. Anterior gill arch with 9–16
(11) rakers, thereof 2–4 (3) elongate. Pseudobranchial
ilaments 0–2 (usually 1).
Head sensory pores (Fig. 17A–C). Supraorbital pores
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 mandibular pore with cirrus. Preopercular pores:
3 lower, irst and second with joined opening; third nontubular; tubular upper preopercular pore. [See description
of Beaglichthys bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with up to four
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 1/2 diameter of
pupil. vomer horseshoe-shaped, with two rows of equally
sized teeth up to 1/3 diameter of pupil. Palatine with one
row of teeth up to 1/3 diameter of pupil. Dentary with
three outer rows of granular teeth and one inner row of
larger teeth anteriorly, merging into one row of larger teeth
posteriorly, up to about size of 3/4 of pupil diameter.
Otolith (Fig. 17I–K). elongate in shape, length to height
2.2–2.3 (51–66 mm Sl) and moderately thin (otolith height
to otolith thickness about 2.2). Anterior tip rounded to
slightly pointed, posterior rim expanded. Dorsal rim with
rounded pre- and post-dorsal angles, the latter sometimes
more pronounced, section in between long and straight,
small concavity towards anterior tip; ventral rim gently
and regularly curved. Inner face moderately convex, outer
face lat to slightly concave, both smooth. Otolith length to
sulcus length 1.9–2.1. Sulcus positioned slightly towards
anterior, with fused colliculi, not inclined towards otolith
axis. ventral furrow distinct, close to ventral rim of otolith,
curved upwards anteriorly and posteriorly to nearly meet
sulcus tips.
Axial skeleton. Neural spine of vertebra 4 (-5) inclined
and (5) 6–8 (9) depressed. Parapophyses present from
Table 9. Meristic and morphometric characters of Didymothallus
mizolepis (Günther, 1867).
holotype
bMnh
1867.9.13
Standard length in mm
47
Meristic characters
Dorsal in rays
92
Caudal inrays
16
Anal in rays
68
Pectoral in rays
21
Precaudal vertebrae
12
Caudal vertebrae
30
Total vertebrae
42
rakers on anterior gill arch
11
Pseudobranchial ilaments
1
D/v
6
d/a
30
v/A
14
Morphometric characters in % of SL
head length
24.1
head width
11.7
head height
14.2
Snout length
5.3
upper jaw length
11.7
Diameter of pigmented eye
2.2
Diameter of pupil
1.1
Interorbital width
5.3
Posterior maxilla height
3.5
Postorbital length
16.8
Preanal length
50.0
Predorsal length
30.1
Body depth at origin of anal in
14.1
Pectoral in length
12.5
Pectoral in base height
5.6
ventral in length
16.4
Base ventral in – anal in origin 32.1
holotype +
non-types
n
Mean (range)
35.0 (14-66)
177
85.6 (78-97)
15.8 (15-17)
63.5 (57-73)
20.4 (18-22)
12.1 (12-13)
30.2 (28-32)
42.2 (40-45)
11.8 (9-16)
1.0 (0-2)
6.1 (5-7)
28.2 (24-33)
14.7 (14-16)
134
18
133
20
138
138
138
25
24
134
134
134
24.6 (23.0-26.3)
12.2 (10.6-13.9)
14.9 (13.7-16.0)
5.2 (4.7-6.0)
12.3 (11.6-13.3)
2.1 (1.8-2.4)
1.3 (1.0-2.1)
5.4 (4.5-6.4)
3.1 (2.1-4.2)
17.9 (16.8-19.0)
50.0 (46.3-52.0)
29.9 (28.3-33.7)
14.7 (12.0-16.5)
14.0 (12.5-16.2)
5.5 (4.7-5.8)
19.8 (15.1-27.3)
32.2 (28.9-34.1)
19
12
12
12
12
21
13
12
19
12
12
12
12
12
12
25
12
species of Didymothallus well from those of eusurculus
gen. nov., but both characters are not available in the
case of the holotype of D. mizolepis. The only remaining
character for distinction then remains the position of the
anterior nostril, which is very low above the upper lip in
the case of D. mizolepis (1/5 to 1/6 the distance from upper
lip to aggregate distance to anterior margin of eye) and
much higher in e. pistillum (1/3 to 1/3.5), a value, already
close to the genus Dinematichthys (< 1/1.3).
In conclusion, the nature of the female type specimen of
D. mizolepis can not be regarded as ultimately ascertained.
The following redeinition of the species therefore is based
on well preserved male specimens now attributed to the
species and based on the complete review of the available
material from the area.
Diagnosis. vertebrae 12-13+28-32=40-45, dorsal in
rays 78–97, anal in rays 57–73, v in D 2.2–2.6; single
(outer) pseudoclasper wing-shaped with two slender,
straight supporters, the posterior one being slightly longer
58
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 16. Didymothallus mizolepis (Günther, 1867). A, fresh dead, WAM P.27967-038, male, 37 mm Sl; B, BMNh 1867.5.13, holotype,
female, 47 mm Sl; C, x-ray, WAM P.31078-031, female, female 57 mm Sl.
vertebrae 6 (7) to 12. Pleural ribs on vertebrae 2 to 11
(12). First anal in pterygophore elongate, sometimes
reaching tip of last precaudal parapophysis in males but
not in females.
Male copulatory organ (Fig. 17D–h). Single pair
of large (outer) pseudoclaspers, wing-shaped, straight
and not bent outwards, tips wider than base, with two
long supporters, posterior being slightly longer and
thicker than anterior. Penis straight, slightly longer than
pseudoclaspers, pointed, with broad base. Both supporters
of pseudoclaspers occasionally bent backwards at their
tips; penis slightly curved.
Colour. live colour known from two specimens (WAM
P.31015-031 and WAM P.27967-038, Fig. 16A), which both
show a uniform dusky red to reddish-violet body colour,
lighter ventrally and darker dorsally. The vertical ins bear
the same colour but lighter and translucent. Preserved
colour is variable brown to greyish-brown, often rather
dark.
Comparison. Didymothallus mizolepis is easily
distinguished from the two other species of the genus,
D. criniceps and D. pruvosti sp. nov. in the higher number
of dorsal in rays (78–97 vs 69–77), higher v in D (2.2–2.6
vs 1.9–2.1), the anterior supporter of the pseudoclasper
being slightly shorter than the posterior (vs being equally
long), the straight penis (vs curved), the long sulcus (otolith
length to sulcus length 1.9–2.1 vs 2.6) and several aspects
of the otolith outline.
Variability. The range of variation seen in D. mizolepis
is unusually high, in fact amongst the highest observed
in species described in this volume; for example the total
number of vertebrae (40 to 45), dorsal in rays (78–97) or
anal in rays (57–73). Other remarkable variations concern
squamation of the cheeks, ranging from a relatively small
patch on the upper cheek to a continuous patch over the
upper and lower cheek, and variation in the pseudoclaspers
with straight or slightly backward bent supporters. It
appears that the more western specimens (exmouth Gulf)
(Fig. 17 C,h,K) tend to show lower meristic numbers,
lesser cheek squamation and bent supporters in the
pseudoclaspers, whereas the most eastern specimens
(Darwin area) (Fig. 17A–B, D–G, I–J) have higher meristic
counts, more extended cheek squamation and straight
supporters in the pseudoclaspers. Geographic overlap
as well as meristic and morphologic overlap, however,
is complete and without an indication of a bimodal
59
W. Schwarzhans and P. r. Møller
Fig. 17. Didymothallus mizolepis (Günther, 1867). A, lateral view of head, AMS I. 24678-053, 1 male, 49 mm Sl; B, ventral view of
head, AMS I. 24678-053,1 male, 49 mm Sl; C, lateral view of head, WAM P.31015-031, 1 female, 51 mm Sl; D, inclined lateral view of
male copulatory organ, SMNS 14557, 44 mm Sl; E, ventral view of male copulatory organ, WAM P.31251-056), 53 mm Sl; F, view of
left pseudoclasper from inside, SMNS 14557, 44 mm Sl; G, view of left pseudoclasper from inside, AMS I. 24678-053, 49 mm Sl; H,
view of left pseudoclasper from inside, AMS I.17060-035, 33 mm Sl; I, median view of right otolith, WAM P.31092-008, male, 51 mm
Sl; J, ventral view of right otolith, WAM P.31092-008, male, 51 mm Sl; K, median view of right otolith, WAM P.31506-001, female,
66 mm Sl.
60
Dinematichthyine ishes of the Indo-west Paciic III
distribution pattern, so that separation into two species is
not justiied at the present stage of knowledge.
Distribution. Didymothallus mizolepis is known from
northern and western Australia from 24°S and 113°e to
the Cape york, the north-easternmost tip of Australia
(Fig. 15). The species is associated with crevices of silty
coralline rock.
Ecology. This species appears to be most common in
inshore and back reef environments.
Biology. Several female specimens are known
from D. mizolepis, and they are remarkable for having
unusually low numbers of large embryos, namely 1 to 3
(predominantly 2) (Fig. 16C). A 32 mm Sl female (uSNM
327949) contained one embryo of 14 mm Sl in length.
This is among the lowest degree of fecundity observed in
Dinematichthyini.
Didymothallus pruvosti sp. nov.
(Figs 15, 18–19; Tables 2, 10)
Material examined. (5 specimens, 28–44 mm Sl).
hOlOTyPe – MNhN 1980-0961, male, 44 mm Sl, ca.
22°40’S, 166°37’e, Passe Mato, New Caledonia, coll. M.l. Bauchot and l. A. Maugé, 13 January 1979, depth 6–10
m. PArATyPeS – SMNS 19687, 1 female, 28 mm Sl,
22°19’S, 166°49’e, Prony Bay, New Caledonia, depth 0–1
m, coll. r. Fricke, 7 Nov. 1998; SMNS 22636, 1 male, 34
mm Sl, 22°19’S, 166°49’e, Prony Bay, New Caledonia,
depth 0–2 m, coll. r. Fricke, 8 May 2000; SMNS 22809,
2 females, 31 and 42 mm Sl, 21°04’S, 165°28’e, Mou
Gulf, eSe of Ponérihuen, New Caledonia, depth 0.3–2.8
m, coll. r. Fricke, 13 May 2000.
Diagnosis. vertebrae 12+31-33=43-45, dorsal in rays
70–73, anal in rays 52–58, D/A 20–22, v in D 1.9; head
without cirri on occiput; single (outer) pseudoclasper
wing shaped with two supporters, the anterior slightly
longer and bent backward, the posterior straight, with
club-like expanded tip; cheeks with 5–6 rows of scales
on upper part and 3–5 rows on lower part; otolith with
sharply pointed anterior tip and moderately pronounced
post-dorsal angle, with undivided short sulcus, its centre
anteriorly of the centre of the otolith, inclined 5°–10°,
otolith length to otolith height = 2.2–2.3, otolith length to
sulcus length = 2.6.
Description (Figs 18, 19). The principal meristic and
morphometric characters are shown in Table 10. Body
Table 10. Meristic and morphometric characters of Didymothallus
pruvosti sp. nov.
holotype
Mnhn
19800961
44
Standard length in mm
Meristic characters
Dorsal in rays
73
Caudal inrays
15
Anal in rays
54
Pectoral in rays
19
Precaudal vertebrae
12
Caudal vertebrae
32
Total vertebrae
44
rakers on anterior gill arch
15
Pseudobranchial ilaments
2
D/v
6
d/a
22
v/A
15
Morphometric characters in % of SL
head length
24.9
head width
10.8
head height
14.2
Snout length
5.3
upper jaw length
11.6
Diameter of pigmented eye
2.3
Diameter of pupil
1.2
Interorbital width
5.5
Posterior maxilla height
3.6
Postorbital length
17.7
Preanal length
51.9
Predorsal length
31.8
Body depth at origin of anal in
12.4
Pectoral in length
14.5
Pectoral in base height
4.9
ventral in length
Base ventral in – anal in origin 34.5
holotype +
4 paratypes
n
Mean (range)
35.2 (28-44)
5
71.8 (70-73)
15.5 (15-16)
54.5 (52-58)
19.3 (19-20)
12
32.0 (31-33)
44.0 (43-45)
14.8 (14-15)
2
6.5 (6-7)
20.8 (20-22)
14.5 (14-15)
4
2
4
3
4
4
4
5
5
4
4
4
25.7 (24.9-26.4)
11.7 (10.8-12.9)
15.0 (14.1-15.9)
5.2 (5.0-5.3)
11.8(11.1-12.4)
2.2 (1.8-2.3)
1.1 (1.0-1.2)
5.8 (5.4-7.0)
3.5 (3.2-3.8)
18.6 (17.7-19.1)
46.4 (43.3-51.9)
31.2 (30.6-32.1)
13.2 (11.5-15.1)
13.9 (12.4-15.5)
5.1 (3.9-5.8)
19.7 (16.6-22.6)
28.6 (26.4-34.5)
5
5
5
5
5
5
5
5
5
4
5
5
5
5
5
5
5
slender; small sized, mature at about 30 mm Sl. head with
scale patch on cheek containing 5–6 (5) vertical rows of
small scales on upper part and 3–5 vertical rows on lower.
horizontal diameter of scales on body about 1.0 % Sl,
in 18 horizontal rows. Occiput without cirri. Maxillary
ending far behind eyes, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded, angular.
Anterior nostril positioned low, 1/4 the distance from
Fig. 18. Didymothallus pruvosti sp. nov., MNhN 1980-0961, holotype, 44 mm Sl.
61
W. Schwarzhans and P. r. Møller
Fig. 19. Didymothallus pruvosti sp. nov. A, lateral view of head, SMNS 22809, female, 42 mm Sl; B, ventral view of head, SMNS 22809,
female, 42 mm Sl; C, inclined lateral view of male copulatory organ, holotype; D, view of left pseudoclasper from inside, holotype;
E, median view of right otolith, SMNS 22636, male, 34 mm Sl.
upper lip to aggregate distance to anterior margin of eye.
Posterior nostril small, about 1/4 the size of eye. Opercular
spine with long free tip, pointed. Anterior gill arch with
14–15 (15) rakers, thereof 3 elongate. Pseudobranchial
ilaments always 2.
Head sensory pores (Fig. 19A, B). Supraorbital pores
2–3. Infraorbital pores 6 (3 anterior and 3 posterior): three
posterior pores about half size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior); irst
anterior mandibular pore without cirrus. Preopercular
pores: 3 lower, irst and second with joined or separate
opening; third tubular; tubular upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to two outer
rows of granular teeth and one inner row of larger teeth.
Anteriormost teeth in inner row up to 2/3 diameter of pupil.
vomer horseshoe-shaped, with two rows of equally sized
teeth up to 2/3 diameter of pupil. Palatine with two rows
of densely set teeth up to 2/3 diameter of pupil. Dentary
with two outer rows of granular teeth and one inner row of
larger teeth anteriorly, merging into one row of larger teeth
posteriorly, up to about size of 3/4 of pupil diameter.
Otolith (Fig. 19e). elongate in shape, length to height
2.2–2.3 (34–54 mm Sl) and moderately thin (otolith height
to otolith thickness about 2.3–2.5). Anterior tip sharply
pointed, posterior rim expanded. Dorsal rim with rounded
pre- and more pronounced postdorsal angles, section in
between straight or slightly curved, markedly concave
towards anterior and less towards posterior tip; ventral
rim gently and regularly curved. Inner face moderately
convex, outer face lat to slightly convex, both smooth.
Otolith length to sulcus length 2.6. Sulcus positioned
slightly towards anterior, with fused colliculi, inclined
5–10° towards otolith axis. ventral furrow faint, close
to ventral rim of otolith, anteriorly terminating at ridge,
connecting rostrum with anterior tip of sulcus.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–9 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis in males but not in females.
62
Dinematichthyine ishes of the Indo-west Paciic III
Male copulatory organ (Fig. 19C, D). Single pair
of large (outer) pseudoclaspers, wing-shaped, slightly
bent outwards, base wider than tip, with two supporters,
anterior slightly longer and bent backward, posterior
straight, with club-like expanded tip. Penis slightly curved,
slightly longer than pseudoclaspers, pointed, with broad
base.
Colour. live colour whitish yellow (SMNS 22809) or
greyish yellow (SMNS 22636). Preserved colour light to
medium brown.
Comparison. Didymothallus pruvosti closely
resembles D. criniceps from the Great Barrier reef, with
which it probably forms an allopatric pair. For distinction
from the other two species of Didymothallus see respective
discussions above.
Distribution. Didymothallus pruvosti is geographically
restricted to the shores of New Caledonia (Fig. 15). The
species is associated with crevices of silty coralline rock;
the presence of live corals is not essential.
Etymology. In honour of Patrice Pruvost, Paris,
MNhN, and his support of our review.
Eusurculus gen. nov.
(Tables 1, 2, 11–13)
Type species: eusurculus pistillum sp. nov. Gender
masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed
low on snout; male copulatory organ with two pairs
of pseudoclaspers, outer pseudoclasper wing shaped
with massive supporter, distally much expanded
and with anteriorly pointing hook below ligament
cover, inner pseudoclasper free, slightly shorter than
outer pseudoclasper, massive, distally sucker-disk-like
expanded; small sized, not exceeding 55 mm Sl; precaudal
vertebrae mostly 12–13 (ranging from 11 to 14), total
vertebrae 41–47, branchiostegal rays 6–7; head with
scale patch on cheek only, no scales on operculum; upper
preopercular pore present; otolith moderately elongate to
elongate (otolith length to height 2.0–2.4), sulcus not or
slightly inclined (0–5°), colliculi separated to partly fused,
but ostium and cauda always distinguishable; maxilla
expanded posteroventrally.
Comparison. With males, eusurculus is well
characterised by the morphology of the pseudoclaspers,
the outer pseudoclasper with a massive supporter with its
anterior hook and free inner pseudoclasper with the suckerdisk-like expanded tip. very similar outer pseudoclaspers
are found in Ungusurculus gen. nov. but their inner
pseudoclasper is claw-like instead of sucker-disk-like.
Otherwise, eusurculus differs from Ungusurculus
gen. nov. in the presence of an upper preopercular pore
(vs absent) and distinguishable ostium and cauda of the
sulcus, usually with separated colliculi (vs fused). It is
assumed though from the pseudoclasper pattern that both
genera are closely related.
Other genera with two pairs of pseudoclaspers, an
upper preopercular pore and distinctive ostium and cauda
of the sulcus of the otolith are Dinematichthys (in part) and
Mascarenichthys gen. nov. from the Indo-west Paciic and
Ogilbia from the Americas. even though Mascarenichthys
gen. nov. and Ogilbia show specialised features of the
inner pseudoclaspers, they are never comparable to the
sucker-disk-like forms of eusurculus. eusurculus tends
to have higher counts of precaudal vertebrae (12–14, but
also regularly 11 in one species) than Mascarenichthys
gen. nov. and Ogilbia (predominantly 11, sometimes 12).
Dorsal in ray counts also distinguish eusurculus from
Mascarenichthys gen. nov. (74–92 vs 62–73).
Species. Three species covering a wide range of
distribution – e. andamanensis sp. nov. from the Andaman
Islands and the south coast of Sumatra, e. pistillum sp. nov.
from the northern coast of Australia and the Coral Sea
and e. pristinus sp. nov. from the Bismarck Sea to the
Solomons and vanuatu.
Etymology. Combined from the latin and Greek eu (=
good, real) and the latin surculus (= grapevine tendril),
referring to the functional analogy with the pseudoclaspers
and the speciic shape of the inner pseudoclasper.
Eusurculus andamanensis sp. nov.
(Figs 20–22; Tables 1, 2, 11)
Material examined. (5 specimens, 24–44 mm Sl).
hOlOTyPe – uSNM 366471, male, 44 mm Sl, Aves
Island, Andaman Islands, India, coll. A.G.K. Menon, 2
Dec. 1970. PArATyPeS – uSNM 263709, 1 female, 24
mm Sl, Smith Island, Andaman Islands, India, coll. A.G.
K. Menon, 22 Feb. 1970; uSNM 366217, 1 male, 35 mm
Sl, 2°00’S, 99°35’e, Siburu Island north of Sipura Island,
Mentawai Islands, north-western Sumatra, Indonesia, coll.
r. Bolin et al., 30 Nov. 1963; uSNM 374167, 1 male, 33
mm Sl, 4°01’S, 101°01’e, south of Pagai Selatan Island,
Mentawai Islands, north-western Sumatra, Indonesia, coll.
r. Bolin et al., 3 Dec. 1965; ZMuC P 771631, 1 male, 35
mm Sl, same data as uSNM 263709.
Diagnosis. vertebrae 12+30-31=42-43, dorsal in rays
78–84, anal in rays 59–63, D/A 21–27, v in D 2.2–2.3;
anterior nostril positioned 1/4.5 to 1/5 the distance from
upper lip to aggregate distance to anterior margin of eye;
two pairs of free pseudoclaspers, outer pseudoclasper
wing-shaped with broad base and distally widened
supporter with anterior hook (covered by ligament), inner
pseudoclasper stalked, with slightly widened base and
distally with concave sucker-disk; cheek with 5–6 rows of
scales on upper part and 3–4 rows on lower part.
Description (Figs 21, 22). The principal meristic and
morphometric characters are shown in Table 11. Body
slender, head with pointed snout; small sized, mature
at about 30 mm Sl. head with scale patch on cheek
containing 5–6 (6) vertical rows of small scales on upper
part and 3–4 vertical rows on lower. horizontal diameter
of scales on body about 1.3 % Sl, in 21 horizontal rows.
63
W. Schwarzhans and P. r. Møller
vomer horseshoe-shaped, with two rows of equally sized
small teeth up to 1/3 diameter of pupil. Palatine with two
rows of equally sized teeth up to 1/3 diameter of pupil.
Dentary with four outer rows of granular teeth and one
inner row of larger teeth anteriorly, merging into one row
of larger teeth posteriorly, up to about size of 3/4 of pupil
diameter.
Otolith. Not known.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–9 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongate, reaching tip of last precaudal
parapophysis in males but not in females.
Male copulatory organ (Fig. 22C-G). Two pairs of large,
free pseudoclaspers, outer pseudoclasper wing-shaped
with broad base and distally widened supporter with
anterior hook (covered by ligament), inner pseudoclasper
stalked, with slightly widened base and distally with
concave sucker-disk. Penis slightly curved, slightly longer
than pseudoclaspers, pointed, with broad base.
Colour. live colour unknown. Preserved colour
medium brown.
Maxillary ending far behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded, angular. Anterior nostril positioned low, 1/4.5
to 1/5 the distance from upper lip to aggregate distance to
anterior margin of eye. Posterior nostril small, about 1/5
the size of eye. Opercular spine with free tip, moderately
pointed. Anterior gill arch with 15–18 (18) rakers, thereof
3 elongate rakers. In holotype and one paratype row of
elongate rakers interrupted by short raker. Pseudobranchial
ilaments 1–2 (1).
Head sensory pores (Fig. 22A, B). Supraorbital
pores 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 mandibular pore with cirrus. Preopercular
pores: 3 lower, irst and second with joined or separate
opening; third tubular; tubular upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to ive outer
rows of granular teeth and one inner row of larger teeth.
Anteriormost teeth in inner row up to 2/3 diameter of pupil.
Fig. 20. Sample sites of eusurculus andamanensis sp. nov., e. pistillum sp. nov. and e. pristinus sp. nov. One symbol may represent
several samples.
64
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 21. eusurculus andamanensis sp. nov., uSNM 366471, holotype, 44 mm Sl.
Fig. 22. eusurculus andamanensis sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male
copulatory organ with hood not bent forward, uSNM 374167, 33 mm Sl; D, ventral view of male copulatory organ with hood bent forward,
uSNM 374167, 33 mm Sl; E, inclined lateral view of male copulatory organ, holotype; F, view of left pseudoclasper from inside, holotype;
G, view of left pseudoclasper from inside, uSNM 366217, 35 mm Sl.
65
W. Schwarzhans and P. r. Møller
treated in the forthcoming part of our review dealing
with that genus.]
Comparison. eusurculus andamanensis resembles
e. pistillum sp. nov. in the pseudoclasper morphology,
except for that in e. andamanensis the sucker-disk of
the inner pseudoclasper is more simple in shape than the
broader and folded disk of e. pistillum sp. nov.. eusurculus
andamensis further differs from e. pistillum sp. nov. in
the lower position of the anterior nostril (the distance from
upper lip to aggregate distance to anterior margin of eye
1/4.5 to 1/5 vs 1/3.5 to 1/4), lower number of precaudal (12
vs predominantly 13) and total vertebrae (42–43 vs 43–47)
and lower D/A (21–27 vs 25–32).
The third species, e. pristinus sp. nov., shows a tip of
the inner pseudoclasper that looks like a miniature suckerdisk with two small hooks, i.e. intermediate between the
eusurculus and the Ungusurculus pattern. eusurculus
andamanensis otherwise resembles e. pristinus sp. nov.
well.
Distribution. eusurculus andamanensis occurs along
the shores of the Andaman Islands to north-western
Sumatra (Fig. 20).
Etymology. Named after the type area, the Andaman
Islands.
Table 11. Meristic and morphometric characters of eusurculus
andamanensis sp. nov.
holotype
uSNM
366471
Standard length in mm
44
Meristic characters
Dorsal in rays
82
Caudal inrays
14
Anal in rays
62
Pectoral in rays
21
Precaudal vertebrae
12
Caudal vertebrae
30
Total vertebrae
42
rakers on anterior gill arch
18
Pseudobranchial ilaments
1
D/v
6
d/a
25
v/A
14
Morphometric characters in % of SL
head length
26.8
head width
12.9
head height
15.0
Snout length
5.5
upper jaw length
14.2
Diameter of pigmented eye
2.7
Diameter of pupil
1.6
Interorbital width
5.4
Posterior maxilla height
4.4
Postorbital length
18.8
Preanal length
53.0
Predorsal length
32.6
Body depth at origin of anal in
14.9
Pectoral in length
13.6
Pectoral in base height
5.5
ventral in length
Base ventral in – anal in origin 31.4
holotype +
4 paratypes
n
Mean (range)
34.0 (24-44)
5
81.2 (78-84)
14
61.4 (59-63)
20.8 (20-22)
12
30.4 (30-31)
42.4 (42-43)
16.4 (15-18)
1.8 (1-2)
6.2 (6-7)
24.6 (21-27)
14.6 (14-15)
5
2
5
5
5
5
5
5
5
5
5
5
26.7 (25.4-28.7)
11.7 (11.1-12.9)
14.8 (14.2-15.2)
5.0 (4.1-5.5)
12.9 (11.8-14.2)
2.3 (2.2-2.7)
1.4 (1.1-1.6)
5.6 (5.4-6.2)
3.8 (3.1-4.4)
19.2 (18.7-20.3)
50.0 (45.7-53.0)
32.1 (30.5-32.8)
14.2 (12.8-14.9)
14.9 (13.4-16.1)
5.8 (5.5-6.2)
19.4 (17.5-20.9)
30.8 (28.0-33.0)
5
5
5
5
5
5
5
5
5
5
5
5
5
5
4
3
4
Eusurculus pistillum sp. nov.
(Figs 20, 23–24; Tables 2, 12)
Material examined. (118 specimens, 13–55 mm
Sl). hOlOTyPe – AMS I. 44520-001, male, 48 mm
Sl, 19°08’S 146°52’e, Queensland, Australia, Magnetic
Island, 21 km northeast of Townsville, 0–3 m depth in
sublittoral; rocks, mud, corals, brown algae; ronald
Fricke; 3 June 1993. PArATyPeS – AMS IA.5089, 1
female, 46 mm Sl, 10°40’S, 142°07’e, Prince of Wales
Island, Australia, coll. G. P. Whitley, May 1931; AMS
IA.5090, 1 female, 45 mm Sl, 10°40’S, 142°07’e, Prince
of Wales Island, Australia, coll. G. P. Whitley, May 1931;
AMS IA.5091, 1 male, 41 mm Sl, 10°40’S, 142°07’e,
Prince of Wales Island, Australia, coll. G. P. Whitley,
May 1931; AMS IB.6233, 2 males, 46–47 mm Sl, 4
females, 26–52 mm Sl, 21°00’S, 152°00’e, Swain reefs,
Great Barrier reef, Australia, coll. Australian Museum
exp., Oct. 1962; AMS IA.6793-002, 1 female, 46 mm
Sl, lindeman Island, Great Barrier reef, Australia,
coll. G. P. Whitley, date unknown; AMS I.19108-146, 2
males, 41–42mm Sl, 2 females, 26–39 mm Sl, 14°40’S,
145°28’e, lizard Island, Great Barrier reef, Australia,
coll. D. hoese and party, 17 Nov. 1975; AMS I.20201062, 4 males, 45–50 mm Sl, 8 females, 31–51 mm Sl,
3 juveniles, 21–28 mm Sl, 23°30’S, 152°05’e, One Tree
Island, Great Barrier reef, Australia, depth 0–2 m, coll.
D. hoese, 29 Sept. 1971; AMS I.20770-015, 1 male, 46
mm Sl, 11°55’S, 143°27’e, Sir Charles hardy Islands,
Great Barrier reef, Australia, coll. AMS-AIMS party,
14 Feb. 1979; AMS I.22579-057, 2 males, 35–42 mm Sl,
1 female, 37 mm Sl, 15°49’S, 145°50’e, Great Barrier
Remarks. Alcock (1890) described as Dinematichthys
piger from Great Coco Island, Andaman Islands, a
dinematichthyine ish, which is of potential relevance
to e. andamanensis. The unique holotype of D. piger
(ZSI F12939) was not available for study. 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. Both observations point to a species of the genus
Dinematichthys, not eusurculus andamanensis, which
does not seem to exceed 50 mm Sl, has higher dorsal
and anal in ray counts (78–84 and 59–63) and has a low
anterior nostril at 1/4.5 to 1/5 the distance from upper lip
to aggregate distance to anterior margin of eye. [There
are two species of the genus Dinematichthys observed
along the shores of the Andaman Sea, which will be
66
Dinematichthyine ishes of the Indo-west Paciic III
reef, Australia, depth 2–4 m, AMS party, 28 Oct. 1981;
AMS I.33739-041, 4 males, 32–41 mm Sl, 2 females,
36–39 mm Sl, 5 juveniles, 13–21 mm Sl, 10°09’S,
144°35’e, Ashmore reef, Coral Sea, Australia, depth
6–9 m, coll. FQN party, 25 Jan. 1993; AMS I.34311-052,
1 male, 42 mm Sl, 22°S, 150°e, Northumberland Isles,
Great Barrier reef, Australia, AMS party, 15 Sept. 1993;
ANSP 120589, 1 male, 40 mm Sl, 3 females, 40–49 mm
Sl, 15°45’S, 145°42’e, endeavour reef, Great Barrier
reef, Australia, depth 2–5 m, coll. J. C. Tyler and C.
Smith, 16 Jan. 1969; ANSP 120593, 2 males, 29–40 mm
Sl, 3 females, 39–49 mm Sl, 3 juveniles, 21–27 mm
Sl, 15°45’S, 145°42’e, endeavour reef, Great Barrier
reef, Australia, depth 1–2.5 m, coll. J. C. Tyler and C.
Smith, 13 Jan. 1969; BPBM 14420, 1 male, 45 mm Sl,
1 female, 51 mm Sl, One Tree Island, Great Barrier
reef, Australia, depth 0–1 m, coll. J. e. randall and J.
h. Choat, 14 Jan. 1974; SMNS 14840, 1 male, 51 mm Sl,
18°59’S, 146°55’e, Great Barrier reef, Australia, depth
0.0–0.5 m, coll. r. Fricke, 1 June 1993; SMNS 26365, 3
males, 36–48 mm Sl, 3 females, 30–54 mm Sl, same
data as holotype; WAM P.25111-046, 2 males, 44–47
mm Sl, 4 females, 48–55 mm Sl, 20°00’S, 116°00’e,
Dampier Archipelago, Western Australia, G. r. Allen
et al., 3 Nov. 1974; WAM P.27980-066, 2 males, 43–44
mm Sl, 2 females, 34–55 mm Sl, 20°26’S, 115°35’e,
Montebello Islands, Western Australia, depth 1–3 m, coll.
G. r. Allen, 25 May 1983; WAM P.29081-004, 2 females,
41–50 mm Sl, 1 juvenile, 19 mm Sl, 15°31’S, 123°09’e,
Adele Island, Western Australia, depth 0.1–2.0 m, coll.
G. r. Allen, 22 Sept. 1986; WAM P.30305-006, 1 female,
50 mm Sl, 1 juvenile, 25 mm Sl, 13°52’S, 126°56’e, Sir
Graham Moore Islands, Western Australia, depth 0–1
m, coll. G. r. Allen, 15 Aug 1991; WAM P.30311-017, 1
male, 42 mm Sl, 1 female, 39 mm Sl, 14°15’S, 125°38’e,
Bonaparte Archipelago, Western Australia, depth 3–4
m, coll. G. r. Allen, 20 Aug. 1991; WAM P.30320-056,
2 males, 34 mm Sl, 1 female, 37 mm Sl, 1 juvenile, 14
mm Sl, 16°05’S, 123°27’e, Powerful Island, Buccaneer
Archipelago, Western Australia, depth 0.1–1.0 m, coll.
G. r. Allen, 26 Aug. 1991; WAM P.30652-062, 6 males,
38–43 mm Sl, 2 females, 30–32 mm Sl, 1 juvenile, 25
mm Sl, 22°33’S, 113°39’e, Norwegian Bay, Western
Australia, depth 1–2 m, coll. B. hutchins et al., 19 June
1993; WAM P.30842-011, 1 male, 40 mm Sl, 2 females,
30–46 mm Sl, 12°10’S, 123°07’e, Ashmore reef,
Western Australia, coll. G. r. Allen and C. Bryce, 18
Sept. 1994; ZMuC P 771638-39, 1 male, 43 mm Sl, 1
female, 31 mm Sl, same data as WAM P.30652-062.
Additional material. uSNM 377263, 1 specimen,
One Tree Island, Great Barrier reef; uSNM 263753, 1
specimen, 40 mm Sl, One Tree Island, Great Barrier reef;
uSNM 366569, 1 male, 42 mm Sl, heron Island, Great
Barrier reef; uSNM 366604, 2 females, 46–50 mm Sl,
One Tree Island, Great Barrier reef; uSNM 366681, 2
males and 2 females, 40–48 mm Sl, heron Island, Great
Barrier reef; uSNM 366683, 2 males and 3 females,
31–46 mm Sl, heron Island, Great Barrier reef; uSNM
366684, 1 male, 46 mm Sl, heron Island, Great Barrier
reef; uSNM 366722, 2 males, 47–48 mm Sl, One Tree
Island, Great Barrier reef.
Diagnosis. vertebrae (12) 13-14+30-34=43-47,
dorsal in rays 80–92, anal in rays 60–70, D/A 25–32,
v in D 2.1–2.4; anterior nostril positioned 1/3 to 1/3.5
distance from upper lip to aggregate distance to anterior
margin of eye; two pairs of free pseudoclaspers, outer
pseudoclasper wing-shaped with broad base and distally
widened supporter with indistinct anterior hook (covered
by ligament), inner pseudoclasper stalked, with narrow
base and distally with large, folded sucker-disk; cheek
with 4–6 rows of scales on upper part and 2–3 rows on
lower; otoliths with distinct ostium and cauda, colliculi
separate or partly fused, otolith length to otolith height
2.2–2.4, ostial colliculum length to caudal colliculum
length 3–4.
Description (Figs 23, 24). The principal meristic and
morphometric characters are shown in Table 12. Body
slender; mature at about 35 mm Sl. head with scale patch
on cheek containing 4–6 vertical rows of scales on upper
part and 2–3 vertical rows on lower. horizontal diameter
of scales on body about 1.8 % Sl, in 24 horizontal rows.
Maxillary ending far behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded, angular. Anterior nostril positioned moderately
low, 1/3.5 to 1/4 the distance from upper lip to aggregate
distance to anterior margin of eye. Posterior nostril small,
about 1/5 to 1/7 the size of eye. Opercular spine with
Fig. 23. eusurculus pistillum sp. nov., AMS I. 44520-001, holotype, 48 mm Sl.
67
W. Schwarzhans and P. r. Møller
Fig. 24. eusurculus pistillum sp. nov. A, lateral view of head, WAM P.30842-011, male, 40 mm Sl; B, ventral view of head, WAM P.30842011, male, 40 mm Sl; C, ventral view of male copulatory organ with hood not bent forward, holotype; D, lateral view of male copulatory
organ with hood not bent forward, holotype; E, ventral view of male copulatory organ with hood bent forward, WAM P.30842-011, 40
mm Sl; F, ventral view of right pseudoclasper, WAM P.30842-011, 40 mm Sl; G, inclined lateral view of male copulatory organ, WAM
P.30842-011, 40 mm Sl; H, view of left pseudoclasper from inside, holotype; I, view of left pseudoclasper from inside, SMNS 26365,
48 mm Sl; J, median view of right otolith, holotype; K, ventral view of right otolith, holotype.
68
Dinematichthyine ishes of the Indo-west Paciic III
Table 12. Meristic and morphometric characters of eusurculus
pistillum sp. nov.
holotype
holotype +
AMS I. 101 paratypes
44520Mean (range)
001
48
38.7 (13-55)
Standard length in mm
Meristic characters
Dorsal in rays
87
Caudal inrays
16
Anal in rays
64
Pectoral in rays
20
Precaudal vertebrae
13
Caudal vertebrae
31
Total vertebrae
44
rakers on anterior gill arch
13
Pseudobranchial ilaments
2
D/v
6
d/a
29
v/A
15
Morphometric characters in % of SL
head length
24.7
head width
13.3
head height
15.3
Snout length
5.2
upper jaw length
12.2
Diameter of pigmented eye
2.2
Diameter of pupil
1.4
Interorbital width
7.0
Posterior maxilla height
4.3
Postorbital length
18.1
Preanal length
49.8
Predorsal length
29.4
Body depth at origin of anal in
14.8
Pectoral in length
13.9
Pectoral in base height
5.3
ventral in length
23.9
Base ventral in – anal in origin 34.9
n
102
86.5 (80-92)
15.7 (15-16)
65.2 (60-70)
21.1 (20-22)
13.0 (12-14)
31.5 (30-34)
44.5 (43-47)
13.4 (8-17)
1.9 (1-2)
5.9 (5-7)
28.3 (25-32)
15.0 (14-16)
69
14
69
10
69
69
69
11
11
69
69
69
24.9 (23.1-26.0)
12.6 (11.3-13.8)
14.7 (13.6-15.8)
5.5 (4.8-6.2)
12.4 (11.1-13.5)
2.4 (2.1-2.7)
1.5 (1.4-1.8)
6.6 (6.0-7.2)
4.0 (3.4-4.5)
17.8 (16.6-18.7)
48.9 (44.7-51.4)
29.8 (28.5-30.9)
15.1 (13.8-17.1)
13.7 (12.2-14.7)
5.4 (4.9-6.1)
23.6 (22.3-26.8)
31.4 (28.2-34.9)
10
10
10
10
10
10
10
10
10
10
9
10
10
10
10
9
10
free pointed tip. Anterior gill arch with 8–17 (13) rakers,
thereof 3 elongate rakers. Pseudobranchial ilaments 1–2
(usually 2).
Head sensory pores (Fig. 24A, 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).
Preopercular pores: 3 lower, irst and second with joined
opening; third tubular; tubular upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to ive outer
rows of granular teeth and one inner row of larger teeth.
Anteriormost teeth in inner row up to 1/2 diameter of
pupil. vomer horseshoe-shaped, with three rows of equally
sized small teeth up to 1/5 diameter of pupil. Palatine with
three rows of equally sized small teeth up to 1/5 diameter
of pupil. Dentary with four outer rows of granular teeth
and one inner row of larger teeth anteriorly, merging into
one row of larger teeth posteriorly, up to about size of 3/4
of pupil diameter.
Otolith (Fig. 24J–K). elongate in shape, length to
height 2.2–2.4 (37–48 mm Sl) and thin (otolith height
to otolith thickness about 2.5). Anterior tip moderately
pointed, posterior rim expanded, mostly pointed. Dorsal
rim with broadly rounded predorsal angle and sharp
postdorsal angle, section in between straight or slightly
concave, also concave in front of predorsal angle and
behind postdorsal angle; ventral rim gently and regularly
curved, deepest anterior to middle. Inner face moderately
convex, outer face concave, both smooth. Otolith length to
sulcus length 1.6–1.8. Sulcus positioned slightly towards
anterior, inclined about 5° towards otolith axis, colliculi
separated or partly joined. Ostium much longer and wider
than cauda (length of ostial colliculum to length of caudal
colliculum 3–4, height of ostial colliculum to height of
caudal colliculum 1.8–2.2). ventral furrow long, distinct,
close to ventral rim of otolith, anteriorly and posteriorly
turning upward.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 (9) depressed. Parapophyses present from
vertebrae (6) 7 to 13. Pleural ribs on vertebrae 2 to 12.
First anal in pterygophore elongate in males, but not
reaching tip of last precaudal parapophysis, less elongated
in females.
Male copulatory organ (Fig. 24C–I). Two pairs of
large, free pseudoclaspers, outer pseudoclasper wingshaped with broad base and distally widened supporter
with small, indistinct anterior hook (covered by ligament),
inner pseudoclasper stalked, with narrow base and distally
with large, folded sucker-disk. Penis slightly curved,
longer than pseudoclaspers, pointed, with broad base.
Colour. live colour reported yellow (BPBM 14420).
Preserved colour light to medium greyish-brown, often
with darker back including the dorsal in.
Comparison. eusurculus pistillum resembles
e. andamanensis (see description for differentiation).
From e. pristinus sp. nov. it differs in the sucker-shaped
inner pseudoclasper (vs miniature sucker-disk with two
small hooks), the somewhat higher position of the anterior
nostril (distance from upper lip to aggregate distance
to anterior margin of eye 1/3.5 – 1/4 vs 1/4 – 1/5), the
generally higher number of precaudal vertebrae (mostly
13–14 vs 11–12) and the more elongate otoliths (otolith
length to otolith height 2.2–2.4 vs 2.0–2.1).
Female and juvenile specimens of e. pistillum
can possibly be confused with members of the genus
Dinematichthys Bleeker, 1855, because of the relatively
high position of the anterior nostril. however, eusurculus
pistillum is easily recognised by the higher number of
precaudal vertebrae of 13 to 14 (one in 118 specimens with
12) vs mostly 11, rarely 12 in Dinematichthys. Another
source of confusion of females could be with Didymothallus
mizolepis, with which e. pistillum shares a large area of
distribution, although not occurring syntopically, possibly
69
W. Schwarzhans and P. r. Møller
due to adaption to differing environments, i.e. more
offshore, outer reef fringes in the case of e. pistillum
and more inshore, backreef environments in the case
of D. mizolepis. Female specimens of e. pistillum are
distinguished best from those of D. mizolepis by their
distinctly higher position of the anterior nostril (see
D. mizolepis for further discussion).
Distribution. eusurculus pistillum shows a disjunctive
distribution pattern along the northern shores of Australia,
in the west from exmouth to the Kimberleys and to the
offshore ashmore reef in the timor Sea and in the east
from the Coral Sea and Cape york southwards to One Tree
Island of the southern reaches of the Great Barrier reef
(Fig. 20). It has not been obtained yet from the shores of
the Northern Territory or the Gulf of Carpentaria. The
holotype and other SMNS materials were found associated
with crevices of silty rock.
Ecology. This species appears to be most common
in reef environments, often on outer reefs, and avoid of
inshore and back reef areas.
Etymology. From pistillum (latin = pistil) referring
to the sucker-disk shape of the inner pseudoclasper,
which resembles the shape of a lower’s pistil. A noun in
apposition.
Table 13. Meristic and morphometric characters of eusurculus
pristinus sp. nov.
holotype
uSNM
374171
Standard length in mm
43
Meristic characters
Dorsal in rays
80
Caudal inrays
14
Anal in rays
60
Pectoral in rays
19
Precaudal vertebrae
12
Caudal vertebrae
30
Total vertebrae
42
rakers on anterior gill arch
14
Pseudobranchial ilaments
2
D/v
6
d/a
27
v/A
15
Morphometric characters in % of SL
head length
26.3
head width
12.3
head height
16.5
Snout length
5.0
upper jaw length
12.9
Diameter of pigmented eye
2.3
Diameter of pupil
1.4
Interorbital width
5.7
Posterior maxilla height
4.3
Postorbital length
19.1
Preanal length
47.8
Predorsal length
31.4
Body depth at origin of anal in
16.6
Pectoral in length
16.1
Pectoral in base height
5.6
ventral in length
21.8
Base ventral in – anal in origin 29.2
Eusurculus pristinus sp. nov.
(Figs 20, 25–26; Tables 2, 13)
Material examined. (30 specimens, 12–46 mm Sl).
hOlOTyPe – uSNM 374171, male, 45 mm Sl, 4°46’S,
146°09’e, Bagabag Island, off Madang, northern Papua
New-Guinea, B. B. Collette, 19 June 1979. PArATyPeS
– AMS IA.793, 1 male, 30 mm Sl, 1 female, 46 mm Sl,
16°37’S, 168°09’e, epi Island, vanuatu, coll. McCulloch
and party, date unknown; AMS I.33732-026, 1 female, 41
mm Sl, 16°49’S, 168°22’e, epi Island, vanuatu, coll. J.
Williams and party, 11 June 1996; uSNM 189954, 1 male,
32 mm Sl, New Georgia, Solomons, coll. W. Chapman,
1944; uSNM 189956, 1 female, 40 mm Sl, New Georgia,
Solomons, coll. W. Chapman and Cheyne, 15 June 1944;
uSNM 356207, 1 male, 39 mm Sl, 2 juveniles, 24–26
mm Sl, 2 larvae, 12 mm Sl, 16°49’S, 168°22’e, Namuka
Island, vanuatu, coll. J. T. Williams et al., 3 June 1996;
holotype +
29 paratypes
n
Mean (range)
28.9 (12-46)
30
81.0 (74-86)
14.8 (14-15)
62.6 (58-70)
19.3 (18-20)
11.5 (11-12)
31.3 (30-33)
42.8 (41-44)
15.3 (14-18)
1.3 (0-2)
6.1 (6-7)
22.4 (18-27)
14.0 (13-15)
24
4
24
9
24
24
24
13
13
24
24
24
26.0 (23.9-28.3)
11.8 (10.8-12.7)
14.9 (12.7-16.5)
5.2 (4.4-5.8)
12.5 (11.7-13.3)
2.3 (1.6-3.1)
1.4 (0.9-1.7)
6.1 (5.6-6.6)
3.6 (3.3-4.3)
19.1 (17.8-21.0)
47.2 (42.3-49.7)
30.1 (30.1-33.0)
15.0 (12.3-17.2)
14.8 (11.9-16.5)
5.6 (4.7-6.2)
22.3 (19.3-26.2)
29.2 (25.5-32.5)
13
13
13
13
13
25
14
13
13
13
13
13
13
13
13
11
12
uSNM 366219, 1 male, 29 mm Sl, 1 female, 23 mm Sl,
4°14’S, 152°26’e, Mioko Island, New Britain, Bismarck
Archipelago, Papua New Guinea, coll. D. Cohen and W.
Davis, 25 Feb. 1965; uSNM 374185, 1 male, 27 mm Sl,
8 females, 23–29 mm Sl, 2 juveniles, 20–21 mm Sl,
Fig. 25. eusurculus pristinus sp. nov., uSNM 374171, holotype, 45 mm Sl.
70
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 26. eusurculus pristinus sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male copulatory
organ with hood not bent forward, uSNM 356207, 39 mm Sl; D, inclined lateral view of male copulatory organ, uSNM 356207, 39 mm
Sl; E, view of left pseudoclasper from inside, uSNM 356207, 39 mm Sl; F, view of left pseudoclasper from inside, holotype; G, median
view of right otolith, holotype; H, ventral view of right otolith, holotype.
Bougainville, Solomons, coll. D. Cohen et al., 11 March
1965; uSNM 384195, 1 female, 40 mm Sl, 1 juvenile,
23 mm Sl, same data as holotype. ZMuC P771632-33,
1 male, 38 mm Sl, 1 female, 34 mm Sl, same data as
uSNM 374185.
Additional specimens. uSNM 189951, 2 females, 35–
42 mm Sl, 8°52’S, 158°30’e, New Georgia, Solomons.
Diagnosis. vertebrae 11-12+30-33=41-44, dorsal in
rays 74–86, anal in rays 58–70, D/A 18–27, v in D 2.0–2.3;
anterior nostril positioned 1/4 to 1/5 distance from upper
lip to aggregate distance to anterior margin of eye; two
pairs of free pseudoclaspers, outer pseudoclasper wingshaped with moderately broad base and distally widened
supporter with strong anterior hook (covered by ligament),
inner pseudoclasper stalked, with slightly widened base
and miniature sucker-disk with two small hooks; cheek
with 5–8 rows of scales on upper part and 3–4 rows on
lower; otoliths with distinct ostium and cauda, colliculi
71
W. Schwarzhans and P. r. Møller
partly fused, otolith length to otolith height 2.0–2.1, ostial
colliculum length to caudal colliculum length 3.5–5.
Description (Figs 25, 26). The principal meristic and
morphometric characters are shown in Table 13. Body
slender; small sized, mature at about 25 mm Sl. head with
scale patch on cheek containing 5–8 (6) vertical rows of
scales on upper part and 3–4 vertical rows on lower part.
horizontal diameter of scales on body about 1.8 % Sl,
in 20 horizontal rows. Maxillary ending far behind eyes,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, angular. Anterior nostril
positioned low, 1/4 to 1/5 the distance from upper lip to
aggregate distance to anterior margin of eye. Posterior
nostril small, about 1/6 the size of eye. Opercular spine
with free tip, moderately pointed. Anterior gill arch
with 14–18 (14) rakers, thereof 2–4 (4) elongate rakers.
Pseudobranchial ilaments 0–2 (2).
Head sensory pores (Fig. 26A, B). Supraorbital pores
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
mandibular pore with or without cirrus. Preopercular pores:
3 lower, irst and second with joined opening; third nontubular; tubular upper preopercular pore. [See description
of Beaglichthys bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with up to ive
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 1/2 diameter
of pupil. vomer horseshoe-shaped, with two rows of
equally sized teeth up to 1/4 diameter of pupil. Palatine
with two rows of equally sized teeth up to 1/3 diameter
of pupil. Dentary with four outer rows of granular teeth
and one inner row of larger teeth anteriorly, merging into
one row of larger teeth posteriorly, up to about size of 3/4
of pupil diameter.
Otolith (Fig. 26G, h). Compact, length to height
2.0–2.1 (39–45 mm Sl) and moderately thin (otolith height
to otolith thickness about 2.0). Anterior tip moderately
pointed, posterior broad, not much expanded. Dorsal rim
with broadly rounded predorsal angle and sharp postdorsal
angle, section in between slightly concave to slightly
convex, markedly concave in front of predorsal angle,
less concave behind postdorsal angle; ventral rim gently
and regularly curved, deepest anterior to middle. Inner
face moderately convex, outer face slightly concave, both
smooth. Otolith length to sulcus length 2.0–2.2. Sulcus
positioned slightly towards anterior, slightly inclined 5°
towards otolith axis, ostium and cauda separated, colliculi
partly fused. Ostium much longer and wider than cauda
(length of ostial colliculum to length of caudal colliculum
3.5–5, height of ostial colliculum to height of caudal
colliculum 2.0–3.0). ventral furrow long, not very distinct,
close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebrae 4–5
inclined and 6–8 (9) depressed. Parapophyses present
from vertebrae 7 to 12. Pleural ribs on vertebrae 2 to 11.
First anal in pterygophore elongate, reaching tip of last
precaudal parapophysis in males but not in females.
Male copulatory organ (Fig. 26C–F). Two pairs of
large, free pseudoclaspers; outer pseudoclasper wingshaped with moderately broad base and distally much
widened supporter with strong anterior hook (covered
by ligament); inner pseudoclasper stalked, with slightly
widened base and distally with miniature sucker-disk with
two small hooks, i.e. intermediate between the eusurculus
and the Ungusurculus pattern. Penis curved, slightly
longer than pseudoclaspers, with broad base.
Colour. live colour unknown. Preserved colour
medium brown.
Comparison. See e. andamanensis and e. pistillum
for detailed comparison.
The shape of the inner pseudoclasper with the miniature
sucker-disk with two small hooks resembles the pattern
found in species of Ungusurculus gen. nov., and appears
intermediate between eusurculus and Ungusurculus
gen. nov. There is one species of Ungusurculus gen. nov.
geographically co-occurring with e. pristinus, U. collettei
sp. nov. eusurculus pristinus, however differs from
all Ungusurculus species in the presence of an upper
preopercular pore (vs absent) and clearly separated ostium
and cauda (vs fused, although sometimes with a small step
at the ventral margin of the sulcus indicating a former
separation of ostium and cauda).
Distribution. eusurculus pristinus is distributed from
the Bismarck Archipelago and Bagabag Island off the New
Guinea mainland to the Solomon Islands and vanuatu
(Fig. 20), an offshore oceanic island distribution pattern
similar to Diancistrus eremitus Schwarzhans, Møller and
Nielsen, 2005.
Etymology. From pristinus (latin = ancient, original)
referring to the simpler pattern of the inner pseudoclasper
when compared to the sucker-disk observed in the other
two species of the genus. A noun in apposition.
Lapitaichthys gen. nov.
(Tables 1, 2, 14)
Type species: Lapitaichthys frickei sp. nov. Gender
masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed
moderately low on snout (1/3.5 the distance from upper
lip to aggregate distance to anterior margin of eye);
male copulatory organ with single pair of small (outer)
pseudoclaspers with single supporter and small hook
at anterior rim; small sized, not exceeding 50 mm Sl;
precaudal vertebrae 12–13, total vertebrae 43–45; dorsal in
rays 92–99, branchiostegal rays 6–7; v in D 2.4–2.6; head
with scale patch on cheek only, no scales on operculum;
upper preopercular pore present; otolith moderately
elongate (otolith length to height 2.0–2.1), sulcus slightly
inclined (5°), colliculi separated, ostium length to cauda
length 2.5–2.8; maxilla expanded posterio-ventrally.
72
Dinematichthyine ishes of the Indo-west Paciic III
Compar i son. T he combi n at ion of a si ngle
pseudoclasper, the presence of an upper preopercular
pore and the separation of the colliculi of the sulcus of
the otolith distinguishes Lapitaichthys from all other
dinematichthyine genera. Monothrix resembles it in
the single pseudoclasper and the presence of an upper
preopercular pore, but has fused colliculi in the undivided
sulcus. eusurculus and Mascarenichthys gen. nov. as well
as the New World Ogilbia and Ogilbichthys all share the
presence of the upper preopercular pore and the separated
colliculi (or at least ostium and cauda with partly fused
colliculi) with Lapitaichthys, but all have two (or three)
pairs of pseudoclaspers with the inner pseudoclaspers
usually being highly specialised. The position of the
anterior nostril is moderately low, intermediate between
the situation in Dinematichthys (less than 1/3 the distance
from upper lip to aggregate distance to anterior margin
of eye) and most other Dinematichthyini (more than
1/4 the distance from upper lip to aggregate distance to
anterior margin of eye). A similar intermediate position
of the anterior nostril is found in some species of the
genus Diancistrus (see Schwarzhans et al. 2005) and in
eusurculus pistillum (see above).
Species. The genus is monotypic.
Etymology. Named in recognition of the lapita
culture, the early indigenous pottery culture of Polynesia,
which was irst discovered in New Caledonia, the region
to which Lapitaichthys appears to be endemic. The name
is based on the local word ‘xaapeta’, meaning ‘dig a hole’,
which was misheard and became ‘lapita’.
Lapitaichthys frickei sp. nov.
(Figs 10, 27–28; Tables 2, 14)
Material examined. (48 specimens, 16–50 mm Sl).
hOlOTyPe – SMNS 23029, male, 43 mm Sl, 22°09.2’S,
166°14.7’e, New Caledonia, coastal reefs about 27 km to
the west-north-west of Nouméa, depth 0.2–2.5 m, coll. r.
Fricke, 23 May 2000. PArATyPeS – MNhN 1980-1005,
2 males, 47–49 mm Sl, 1 females, 45 mm Sl, Anse vata,
Nouméa, New Caledonia, depth 0–2 m, coll. J. C. randall
and l. A. Mauge, 14 Jan. 1979; NMNZ P.029586, 1 male,
46 mm Sl, 22°38.6’S, 166°38.6’e, 6 miles off Goelands
Kay, New Caledonia, depth 1–2 m, coll. C. D. roberts
and C. Paulin, 27 Oct. 1992; rOM 65316, 2 males, 45–47
mm Sl, 4 females, 38–43 mm Sl, 1 juvenile, 22 mm
Sl, 22°16.3’S, 166°23.0’e, Isle Nou, New Caledonia,
coll. G. Klassen and M. Kulbicki, 29 Aug. 1991; rOM
65318, 2 males, 43–47 mm Sl, 4 females, 39–45 mm Sl,
3 juveniles, 16–18 mm Sl, 22°21’S, 166°21’e, N of Isle
De Crouy, New Caledonia, coll. G. Klassen and P. Tirard,
16 Sept. 1991; SMNS 18238, 2 females, 46–48 mm Sl,
22°12.3’S, 166°21.3’e, coastal reefs about 10 km to the
west-north-west of Nouméa, New Caledonia, depth 0–1
m, coll. r. Fricke, 22 July 1996; SMNS 22849, 1 female,
49 mm Sl, 22°34.1’S, 167°25.4’e, Ile des Pins, New
Caledonia, depth 0–2.5 m, coll. r. Fricke, 16 May 2000;
Table 14. Meristic and morphometric characters of Lapitaichthys
frickei sp. nov.
holotype
SMnS
23029
Standard length in mm
43
Meristic characters
Dorsal in rays
93
Caudal inrays
16
Anal in rays
66
Pectoral in rays
22
Precaudal vertebrae
12
Caudal vertebrae
32
Total vertebrae
44
rakers on anterior gill arch
16
Pseudobranchial ilaments
1
D/v
6
d/a
32
v/A
15
Morphometric characters in % of SL
head length
23.9
head width
12.2
head height
14.8
Snout length
5.4
upper jaw length
12.3
Diameter of pigmented eye
2.7
Diameter of pupil
1.9
Interorbital width
6.8
Posterior maxilla height
3.8
Postorbital length
16.9
Preanal length
49.3
Predorsal length
28.8
Body depth at origin of anal in
16.4
Pectoral in length
14.2
Pectoral in base height
4.8
ventral in length
22.9
Base ventral in – anal in origin 33.7
holotype +
47 paratypes
n
Mean (range)
39.5 (16-50)
48
95.0 (89-99)
15.6 (14-16)
68.7 (62-74)
22.3 (22-23)
12.0 (12-13)
31.8 (30-33)
43.9 (43-45)
16.4 (15-22)
1.0 (0-2)
6
31.4 (28-35)
15.2 (14-17)
31
31
31
8
34
34
34
10
10
32
32
31
24.6 (22.7-27.8)
11.5 (9.1-13.1)
14.4 (12.1-16.2)
5.4 (4.5-7.1)
12.2 (11.1-14.1)
2.6 (2.3-3.3)
1.4 (1.2-1.9)
6.5 (5.5-7.6)
4.0 (3.1-4.6)
17.0 (15.6-20.4)
48.4 (46.4-50.5)
29.1 (27.5-32.5)
15.1 (11.8-16.4)
13.3 (11.7-15.0)
5.6 (4.8-7.2)
22.0 (19.5-26.6)
31.1 (28.0-33.7)
10
10
10
10
10
10
10
10
10
10
8
9
10
10
10
10
8
SMNS 26367, 1 male, 41 mm Sl, 3 females, 32–43 mm Sl,
same data as holotype; SMNS 25443, 1 male, 45 mm Sl, 4
females, 38–50 mm Sl, 1 juvenile, 23 mm Sl, 22°36.4’S,
167°24.8’e, Ilot Mwéré, Iles des Pins, New Caledonia,
0–1.5 m, coll. r. Fricke, 27 Oct. 2006; uSNM 319897, 6
males, 27–45 mm Sl, 7 females, 36–45 mm Sl, 22°15’S,
166°22’e, northwest off Nouméa, New Caledonia, depth
2 m, coll. J. T. Williams and G. Mhou Tham, 8 Nov. 1991;
ZMuC P771619-20, same data as uSNM 319897, 1 male,
43 mm Sl, 1 female, 40 mm Sl.
Diagnosis. See generic diagnosis.
Description (Figs 27, 28). The principal meristic and
morphometric characters are shown in Table 14. Body and
head slender, snout pointed; small sized, mature at about
25 mm Sl. head with scale patch on cheek containing 6–7
vertical rows of scales on the upper part and 4 vertical rows
on the lower part. horizontal diameter of scales on body
about 1.5 % Sl, in 22 horizontal rows. Maxillary ending
far behind eyes, dorsal margin of maxillary covered by
73
W. Schwarzhans and P. r. Møller
Fig. 27. Lapitaichthys frickei sp. nov., SMNS 23029, holotype, 43 mm Sl.
upper lip dermal lobe, posterior end expanded, angular
with knob. Anterior nostril positioned moderately low,
1/3.5 the distance from upper lip to aggregate distance to
anterior margin of eye. Posterior nostril small, about 1/5
size of eye. Opercular spine with short free tip, pointed.
Anterior gill arch with 15–22 (16) rakers, thereof 2–3 (2)
elongate rakers. Pseudobranchial ilaments 0–2 (1).
Head sensory pores (Fig. 28A, B). Supraorbital pores
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 mandibular pore without cirrus. Preopercular
pores: 3 lower, irst and second with joined opening;
third non-tubular; tubular upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with up to three
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 1/2 diameter
of pupil. vomer horseshoe-shaped, with two rows of
Fig. 28. Lapitaichthys frickei sp. nov. A, lateral view of head, uSNM 319897, male, 45 mm Sl; B, ventral view of head, uSNM 319897,
male, 45 mm Sl; C, inclined lateral view of male copulatory organ, holotype; D, view of left pseudoclasper from inside, holotype; E,
median view of right otolith, uSNM 319897, male, 45 mm Sl; F, ventral view of right otolith, uSNM 319897, male, 45 mm Sl.
74
Dinematichthyine ishes of the Indo-west Paciic III
equally sized teeth up to 1/3 diameter of pupil. Palatine
with two rows of equally sized teeth up to 1/4 diameter of
pupil. Dentary with two outer rows of granular teeth and
one inner row of larger teeth anteriorly, merging into one
row of larger teeth posteriorly, up to about size of 1/2 of
pupil diameter.
Otolith (Fig. 28e, F). Moderately elongate in shape,
length to height 2.0–2.1 (36–49 mm Sl) and thick (otolith
height to otolith thickness about 2). Anterior tip slightly
pointed; posterior tip more expanded and pointed. Dorsal
rim with sharp predorsal angle and sharp projecting
postdorsal angles, section in between concave, also
markedly concave in front of predorsal angle and behind
postdorsal angle; ventral rim gently and evenly curved,
deepest about at middle. Inner face convex, outer face
slightly concave, both smooth. Otolith length to sulcus
length 1.6–1.8. Sulcus positioned slightly towards anterior,
slightly inclined 5° towards otolith axis, markedly
deepened, colliculi clearly separated. Ostium longer and
wider than cauda (length of ostial colliculum to length of
caudal colliculum 2.5–2.8, height of ostial colliculum to
height of caudal colliculum 1.6–1.9). ventral furrow long,
distinct, very close to ventral rim of otolith, posteriorly
turning upwards; dorsal depression narrow, bent, almost
resembling ventral furrow.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–9 depressed. Parapophyses present from vertebrae
6 to 12. Pleural ribs on vertebrae 2 to 11 (12). First anal
in pterygophore elongate, but usually not reaching tip of
last precaudal parapophysis in males and even shorter in
females.
Male copulatory organ (Fig. 28C–D). Single pair of
small (outer) pseudoclaspers, simple wing-shaped, with
single supporter, often with little claw-like hook at middle
of anterior rim of pseudoclasper. Penis stout, not much
longer than pseudoclasper, curved, with broad basis.
Colour. live colour yellow (SMNS 23029, SMNS
22849). Preserved colour usually very light to light
brown.
Comparison. See comparison between Lapitaichthys
and other genera.
Distribution. Known exclusively from south-eastern
New Caledonia, from the vicinity of Nouméa to the Isle
des Pins (Fig. 10).
Ecology. Lapitaichthys frickei is restricted to lagoonal
and back reef environments. The holotype and most SMNS
paratypes were found in crevices of coralline rock of
fringing lagoonal reefs associated with live corals. SMNS
22849 from the Ile des Pins was found in crevices of the
coralline rock of a lagoonal patch reef, also associated
with live corals.
Etymology. Named in honour of ronald Fricke,
Stuttgart, SMNS, in recognition of his many contributions
to the ishes of the south-western West Paciic and his
support in making available the valuable material from
the SMNS collection.
Majungaichthys gen. nov.
(Tables 1, 2, 15)
Type species: Majungaichthys simplex sp. nov. Gender
masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed low on
snout (1/4 the distance from upper lip to aggregate distance
to anterior margin of eye); male copulatory organ with
two pairs of small pseudoclaspers, outer pseudoclasper
simple, lap-like, not much larger than simple lap-like
inner pseudoclasper; small sized ish, not much exceeding
40 mm Sl; precaudal vertebrae 12, total vertebrae 43–44;
dorsal in rays 71–79; v in D 1.9–2.1; head with scale patch
on cheek only, no scales on operculum; upper preopercular
pore present; otolith moderately elongate (otolith length
to height 2.1–2.2), sulcus short (otolith length to sulcus
length 1.9) slightly inclined (5°), colliculi fused (sometimes
incomplete with indication of distinction of ostial and
caudal colliculi); maxilla expanded posteroventrally.
Comparison. Majungaichthys shares the combination
of two pairs of pseudoclasper, the presence of an upper
preopercular pore and the (sometimes incompletely)
fused colliculi of the undivided sulcus of the otolith with
a number of dinematichthyine genera, i.e. Brotulinella,
Diancistrus, Paradiancistrus and Ungusurculus n. gen.
from the Indo-west Paciic and Pseudogilbia from american
waters. They are, however, all distinguished by their
respective specializations of their pseudoclaspers – the
anteriorly joined inner and outer pseudoclaspers found in
Brotulinella, Diancistrus and Paradiancistrus, the claw-like
specialization of the inner pseudoclasper and the hooked
outer pseudoclasper in Ungusurculus gen. nov. – and in the
case of Paradiancistrus and Pseudogilbia by the presence
of a single lower preopercular pore (vs 3 pores). In terms
of the pseudoclasper morphology, Majungaichthys (and
Mascarenichthys gen. nov., see later) is the most similar
of all the Indo-west Paciic Dinematichthyini to the most
common American genus Ogilbia. Majungaichthys differs
from both these genera in the otolith showing an undivided
sulcus with fused colliculi (vs divided and separated
colliculi). From Mascarenichthys it further differs in the
higher number of precaudal vertebrae (12 vs 11) and total
vertebrae (43–44 vs 39–42).
Species. The genus is monotypic.
Etymology. Named after the Majunga (Mahajanga)
province of Madagascar, where the type locality is
located.
Majungaichthys simplex sp. nov.
(Figs 29–31; Tables 2, 15)
Material examined. (7 specimens, 26–40 mm Sl).
hOlOTyPe – uSNM 374170, male, 37 mm Sl, 16°21’S,
43°59’e, Ile Chesterield, off Cape Saint-Andre, Majunga
province, Madagascar, l. W. Knapp et al., 16 Oct. 1964.
PArATyPe – uSNM 384196, 1 female, 40 mm Sl, same
data as holotype.
75
W. Schwarzhans and P. r. Møller
Table 15. Meristic and morphometric characters of Majungaichthys
simplex sp. nov.
holotype Paratype Tentatively
uSNM uSNM
assigned
374170 384196 non-types n = 4
Standard length in mm
37
Meristic characters
Dorsal in rays
79
Caudal inrays
14
Anal in rays
58
Pectoral in rays
20
Precaudal vertebrae
12
Caudal vertebrae
31
Total vertebrae
43
rakers on anterior gill arch
13
Pseudobranchial ilaments
2
D/v
6
d/a
24
v/A
15
Morphometric characters in % of SL
head length
25.6
head width
11.6
head height
15.0
Snout length
5.6
upper jaw length
12.9
Diameter of pigmented eye
2.5
Diameter of pupil
1.6
Interorbital width
7.0
Posterior maxilla height
4.2
Postorbital length
18.2
Preanal length
49.5
Predorsal length
32.8
Body depth at origin of
16.4
anal in
Pectoral in length
16.5
Pectoral in base height
5.4
ventral in length
Base ventral in –
30.1
anal in origin
Fig. 29. Sample sites of Majungaichthys simplex sp. nov.,
Mascarenichthys heemstrai sp. nov., M. microphthalmus sp.
nov. and Mascarenichthys sp. One symbol may represent several
samples.
Tentatively assigned specimens. uSNM 374164,
1 female, 26 mm Sl, 10°19’S, 56°35’e, Agalega Island
(north island), Mauritius; uSNM 374174, 3 females, 30–35
mm Sl, 10°19’S, 56°35’e, Agalega Island (north island),
Mauritius; uSNM 374175, 1 female, 32 mm Sl, 10°21’S,
56°35’e, Agalega Island (north island), Mauritius.
Diagnosis. See generic diagnosis.
Description (Figs 30, 31). The principal meristic and
morphometric characters are shown in Table 15. Body
moderately elongate with blunt snout; small sized, mature
at about 35 mm Sl. head with scale patch on cheek
containing 4–5 (5) 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 20 horizontal rows.
Maxillary ending well behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
40
Mean (range)
32.4 (26-35)
78
14
59
20
12
32
44
15
2
6
24
15
72.8 (71-75)
14.3 (14-15)
59.0 (56-62)
18.3 (18-19)
12
31.5 (31-32)
43.5 (43-44)
14.0 (12-17)
2
6
20.8 (20-21)
13.8 (13-14)
24.2
12.5
15.5
5.4
12.6
2.4
1.7
6.3
4.3
17.9
46.8
30.3
26.2 (24.7-26.9)
12.6 (11.9-13.5)
15.1 (14.6-16.2)
5.8 (5.3-6.3)
13.5 (12.7-14.1)
2.4 (1.9-2.8)
1.5 (1.2-1.8)
6.6 (6.1-7.3)
4.0 (3.6-4.4)
18.3 (18.0-19.1)
44.1 (41.9-45.3)
32.1 (31.4-32.9)
16.8
15.1 (13.8-15.8)
15.2
5.7
25.9
15.2 (13.6-16.2)
5.3 (4.3-5.9)
22.2 (19.7-24.9)
30.3
26.7 (25.5-27.5)
expanded, angular. Anterior nostril positioned moderately
low, 1/4 the distance from upper lip to aggregate distance
to anterior margin of eye. Posterior nostril small, about
Fig. 30. Majungaichthys simplex sp. nov., uSNM 374170, holotype, 37 mm Sl.
76
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 31. Majungaichthys simplex sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male
copulatory organ, holotype; D, inclined lateral view of male copulatory organ, holotype; E, view of left pseudoclasper from inside,
holotype; F, median view of right otolith, holotype; G, median view of right otolith, uSNM 374174, female, 35 mm Sl.
1/5 the size of eye. Opercular spine with short free tip,
pointed. Anterior gill arch with 12–17 (15) rakers, thereof
3 elongate rakers. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 31A, B). Supraorbital pores
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 mandibular pore with cirrus. Preopercular pores:
3 lower, irst and second with joined opening; third nontubular; tubular upper preopercular pore. [See description
of Beaglichthys bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with up to four
outer rows of granular teeth and one inner row of larger
teeth. Anteriormost teeth in inner row up to 2/3 diameter
of pupil. vomer horseshoe-shaped, with two rows of
equally sized teeth up to 1/4 diameter of pupil. Palatine
with a single row of teeth, up to 1/4 diameter of pupil.
Dentary with three outer rows of granular teeth and one
inner row of larger teeth anteriorly, merging into one row
of larger teeth posteriorly, up to about size of 3/4 of pupil
diameter.
Otolith (Fig. 31F, G). Moderately elongate in shape,
length to height 2.1 (35–37 mm Sl) and moderately thick
(otolith height to otolith thickness about 2.1). Anterior tip
angular, rather blunt; posterior tip incomplete, probably
somewhat expanded. Dorsal rim with rounded pre- and
postdorsal angles, section in between slightly concave, also
concave behind postdorsal angle; ventral rim gently and
evenly curved, somewhat deeper in front of middle. Inner
face convex, outer face lat, both smooth. Otolith length
to sulcus length 1.9. Sulcus positioned slightly towards
anterior, slightly inclined 5° towards otolith axis, ostium
and cauda undivided, colliculi fused. ventral furrow long,
distinct, close to ventral rim of otolith, posteriorly turning
upward to meet posterior tip of sulcus.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 depressed. Parapophyses present from vertebrae 7
to 12. Pleural ribs on vertebrae 2 to 10 (11 in the specimens
77
W. Schwarzhans and P. r. Møller
from Agalega). First anal fin pterygophore elongate,
reaching tip of last precaudal parapophysis in males but
not in females.
Male copulatory organ (Fig. 31C-e). Two pairs of
moderately large, simple, lap-like pseudoclaspers, the
outer being just slightly longer than the inner; isthmus
wide. Penis curved, slightly longer than pseudoclasper,
with broad basis.
Colour. live colour unknown. Preserved colour
medium reddish brown.
Comparison. See comparison between Majungaichthys
and other genera.
Distribution. exclusively known from the small Ile
Chesterield, Majunga province, off the north-western
coast of Madagascar (Fig. 29). Five female specimens
from the Agalega Islands to the north-east of Madagascar
are only tentatively assigned to the species. They differ
slightly from the two specimens from Madagascar in the
lower dorsal in ray count (71–75 vs 78–79), lower D/A
(20–21 vs 24), 1 loin vertebrae (vs 2) and the distance from
the base of the ventral in to the anal in origin (25.5–27.5
vs 30.1–30.3). It is expected that these small differences
represent intraspeciic variation, but it is also possible
that, once pseudoclaspers from males from Agalega have
become known, they could qualify for a distinct species.
Etymology. From simplex (latin = simple) referring
to the simple pattern of the pseudoclaspers. A noun in
apposition.
the penis distinguish Mascarenichthys from eusurculus
with its stalked inner pseudoclasper terminating in
a sucker-disk. The low dorsal fin ray count (62–73)
further distinguishes Mascarenichthys from eusurculus
(74–92).
For distinction from Majungaichthys, with which it
co-occurs in the general area, see description of the genus
(above).
Of all I ndo -west Pacif ic Di nemat icht hy i n i,
Mascarenichthys (and Majungaichthys, see above) appears
to be closest to the American genus Ogilbia, with which
it shares many diagnostically important characters (see
above). The main difference is the pocket on the isthmus,
in which the inner pseudoclasper is hidden in the resting
position. This has never been observed in Ogilbia, but is a
common character in another american dinematichthyine
genus (Ogilbichthys) which has two inner pseudoclaspers
(like Dactylosurculus) of which the anterior one in some
species is partly hidden in a pocket of the isthmus. Whether
this rather unique development represents a synapomorphy
or is a functional analogy remains uncertain at present. In
Mascarenichthys the inner pseudoclasper is so well hidden
in the pocket of the isthmus in such a way that without
speciic unearthing with the use of forceps it may easily
go unrecognised.
Species. The genus includes two new species plus a
possible further species currently left in open nomenclature
due to the lack of adequate male specimens.
Etymology. Named in reference to the Mascarene
plate, from where most of the specimens observed so far
have been obtained.
Mascarenichthys gen. nov.
(Tables 1, 2, 16, 17)
Type species: Mascarenichthys heemstrai sp. nov.
Gender masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed low on
snout (1/4 to 1/5 the distance from upper lip to aggregate
distance to anterior margin of eye); male copulatory organ
with two pairs of pseudoclaspers, the inner broad, with
three ridges in anterior, posterior and inward direction
and hidden in pocket of wide isthmus in resting position;
tip of penis bent, hook-like, at 90° angle; small sized, not
much exceeding 50 mm Sl; precaudal vertebrae 11, total
vertebrae 39–42; dorsal in rays 62–73, branchiostegal
rays 6–7; v in D 1.8–2.1; head with scale patch on cheek
only, no scales on operculum; upper preopercular pore
present; otolith very elongate (otolith length to height
2.3–2.5), sulcus slightly inclined (5°), colliculi separated,
ostium length to cauda length 3.5–4.5; maxilla expanded
posteroventrally, rounded.
Comparison. Mascarenichthys shares the combination
of two pairs of pseudoclaspers, the presence of an upper
preopercular pore and the separated colliculi of the clearly
divided sulcus of the otolith, with eusurculus from the
Indo-west Paciic and Ogilbia from American waters. The
broad inner pseudoclasper with the three ridges, its itting
in a pocket of the isthmus and the hook-like bent tip of
Mascarenichthys heemstrai sp. nov.
(Figs 29, 32, 33; Tables 2, 16)
Material examined. (77 specimens, 19–51 mm Sl).
hOlOTyPe – ANSP 138387, male, 44 mm Sl, 4°39’S,
55°31’e, off eastern Mahe, Anonyme Island, large
sandy-bottom tidepool, Seychelles, depth 0–2 m, coll.
D. Dockins and r. rosenblatt, 4 Feb. 1964. PArATyPeS
– Seychelles: ANSP 138385, 1 male, 36 mm Sl, 5°25’S,
53°18’e, Amirante Islands, D’Arros Island, depth 0–2.5
m, coll. J. e. Boehlke et al., 6 March 1964; ANSP 187333,
5 males, 28–42 mm Sl, 14 females 23–51 mm Sl, same
data as holotype; ANSP 179178, 4 males, 28–41 mm Sl,
8 females, 26–48 mm Sl, 2 juveniles, 19–22 mm Sl,
4°39’S, 55°30’e, off eastern Mahe, depth 0–3 m, coll.
J. e. Boehlke et al., 10 Feb. 1964; BMNh 1932.7.29.48,
1 female, 51 mm Sl, Cerf Island, north of Mahe, coll.
Wood, date unknown; BPBM 21598, 2 males, 30–37 mm
Sl, 2 females, 36–41 mm Sl, la Digue Island, depth
0–1 m, coll. J. e. and h. A. randall and D. Woodland, 1
June 1977; BPBM 22979, 1 female, 24 mm Sl, la Digue
Island, depth 0–1 m, coll. J. e. and h. A. randall and D.
Woodland, 1 June 1977; uSNM 267205, 5 males, 27–34
mm Sl, 4 females, 24–33 mm Sl, Mahe, depth 0–3 m,
coll. J. T. Williams at al., 1 May 1984; Cargados Carajos:
78
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 32. Mascarenichthys heemstrai sp. nov., ANSP 138387, holotype, 44 mm Sl.
uSNM 366232, 2 males, 21–36 mm Sl, 6 females, 24–36
mm Sl, 1 juvenile, 22 mm Sl, 16°43’S, 59°35’e, veronge
Bay, depth 0–1 m, coll. v. Springer et al., 11 April 1976;
uSNM 374165, 1 juvenile, 20 mm Sl, 16°15’S, 59°33’e,
Albatross Island, depth 0–18 m, coll. v. Springer et al.,
6 April 1976; uSNM 374168, 2 males, 5 females, 31–42
Table 16. Meristic and morphometric characters of Mascarenichthys
heemstrai sp. nov.
holotype
anSP
138387
Standard length in mm
44
Meristic characters
Dorsal in rays
68
Caudal inrays
15
Anal in rays
52
Pectoral in rays
18
Precaudal vertebrae
11
Caudal vertebrae
28
Total vertebrae
39
rakers on anterior gill arch
15
Pseudobranchial ilaments
2
D/v
6
d/a
23
v/A
14
Morphometric characters in % of SL
head length
26.4
head width
14.3
head height
16.3
Snout length
5.7
upper jaw length
14.1
Diameter of pigmented eye
2.1
Diameter of pupil
1.3
Interorbital width
6.6
Posterior maxilla height
3.3
Postorbital length
19.2
Preanal length
51.5
Predorsal length
32.8
Body depth at origin of anal in
17.3
Pectoral in length
14.5
Pectoral in base height
6.4
ventral in length
22.3
Base ventral in – anal in origin 31.9
holotype +
47 paratypes
n
Mean (range)
35.0 (19-51)
77
67.4 (62-73)
15.6 (15-16)
51.5 (46-56)
17.5 (16-18)
11
29.1 (28-31)
40.1 (39-42)
14.6 (13-18)
2
6.1 (5-7)
20.9 (17-25)
13.7 (13-15)
69
16
69
37
68
68
69
22
20
69
69
69
25.7 (23.4-27.5)
12.7 (11.9-14.3)
14.5 (12.8-16.3)
5.6 (5.1-6.5)
12.8 (11.0-14.3)
2.1 (1.5-2.7)
1.3 (0.8-1.9)
5.8 (5.1-6.6)
3.4 (3.0-4.4)
18.5 (16.1-20.7)
48.0 (44.1-51.5)
32.2 (29.4-35.5)
15.2 (13.0-18.6)
13.8 (11.5-16.1)
5.4 (5.0-6.4)
21.9 (17.1-26.2)
30.1 (27.1-33.1)
44
25
25
25
25
47
40
24
26
25
41
42
26
26
25
43
23
mm Sl, 16°25’S, 59°36’e, raphael Island, depth 6–11
m, coll. v. Springer et al., 6 April 1976; uSNM 374169, 1
male, 5 females, 23–36 mm Sl, 16°28’S, 59°40’e, raphael
Island, depth 0–6 m, coll. v. Springer et al., 5 Apr 1976;
ZMuC P771621-22, 1 male, 30 mm Sl, 1 female, 34 mm
Sl, same data as uSNM 366232; Mauritius: AMS I.28101033, 1 male, 34 mm Sl, 20°30.8’S, 57°33.7’e, south coast
at Senneville, riviere des Anguilles, depth 0.2–1.5 m,
coll. J. Paxton, O. Grifiths and M. Welshman, 1 Nov.
1988; BPBM 20202, 2 females, 39–51 mm Sl, between
Trou d’eau Douce and Palmar, depth 0.5–1.5 m, coll. J.
e. randall, 7 November 1973; reunion: SMNS 21084, 1
female, 42 mm Sl, 21°09.2’S, 55°16.3’e, SSe Pointe des
Chateaux, west coast of Island, depth 0–1.5 m, coll. r.
Fricke, 31 Dec. 1998.
Additional material. uSNM 349826, 1 female, 39 mm
Sl, 20°30’S, 57°34’e, south-eastern coast of Mauritius.
Diagnosis. vertebrae 11+28-30 (rarely 31) = 39–41
(rarely 42), dorsal in rays 62–73, anal in rays 46–56,
pectoral fin rays 16–18, D/A 17–25, v in D 1.8–2.1;
anterior nostril positioned 1/4–1/5 the distance from
upper lip to aggregate distance to anterior margin of eye;
eyes moderately large (1.5–2.7 % Sl); two pairs of free
pseudoclaspers, outer pseudoclasper simple, lap-shaped
with pointed tip, broad at base, short, slightly longer than
inner pseudoclasper, inner pseudoclasper broad, with
three ridges in anterior, posterior and inward direction
and hidden in pocket of wide isthmus (in resting position);
cheek with 5–7 rows of scales on upper part and 3–4 rows
on lower part; upper preopercular pore present; otolith very
elongate (otolith length to height 2.3–2.5), sulcus slightly
inclined (5°), colliculi separated, ostium length to cauda
length 3.5–4.5.
Description (Figs 32, 33). The principal meristic and
morphometric characters are shown in Table 16. Body
moderately elongate with moderately pointed snout; small
sized, mature at about 25 mm Sl. head with scale patch
on cheek containing 5–7 (7) vertical scale rows on upper
part and 3–4 vertical rows on lower. horizontal diameter
of scales on body about 1.2 % Sl, in 21 horizontal rows.
Maxillary ending well behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior
end expanded, angular. Anterior nostril positioned low,
79
W. Schwarzhans and P. r. Møller
Fig. 33. Mascarenichthys heemstrai sp. nov. A, lateral view of head, BMNh 1932.7.29.48, female, 51 mm Sl; B, lateral view of head,
SMNS 21084, female, 42 mm Sl; C, ventral view of head, BPBM 20202, female, 51 mm Sl; D, lateral view of head, uSNM 366232,
male, 36 mm Sl; E, ventral view of head, uSNM 366232, male, 36 mm Sl; F, ventral view of male copulatory organ with hood not bent
forward, uSNM 366232, male, 36 mm Sl; G, ventral view of male copulatory organ with hood bent forward and inner pseudoclaspers
submerged in pocket of isthmus, holotype; H, ventral view of male copulatory organ with hood bent forward and inner pseudoclaspers
excavated, holotype; I, inclined lateral view of male copulatory organ, AMS I.28101-033, male, 34 mm Sl; J, view of left pseudoclasper
from inside, holotype; K, view of left pseudoclasper from inside, BPBM 21598, male, 37 mm Sl; L, view of left pseudoclasper from
inside, uSNM 366232, male, 36 mm Sl; M, median view of right otolith, BMNh 1932.7.29.48, female, 51 mm Sl; N, ventral view of
right otolith, BMNh 1932.7.29.48, female, 51 mm Sl; O, median view of right otolith, SMNS 21084, female, 42 mm Sl; P, median view
of right otolith, uSNM 366232, male, 36 mm Sl.
80
Dinematichthyine ishes of the Indo-west Paciic III
1/4–1/5 distance from upper lip to aggregate distance to
anterior margin of eye. Posterior nostril small, about 1/4
size of eye. Opercular spine with free tip, pointed. Anterior
gill arch with 13–18 (15) rakers, thereof 3 elongate rakers.
Pseudobranchial ilaments 2.
Head sensory pores (Fig. 33A–e). 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);
irst anterior mandibular pore with or without cirrus.
Preopercular pores: 3 lower, irst and second with joined
opening; third non-tubular; tubular upper preopercular
pore. [See description of Beaglichthys bleekeri for position
of pores.]
Dentition (of holotype). Premaxilla with four outer
rows of granular teeth and one inner row of larger teeth.
Anteriormost teeth in inner row up to 2/3 of diameter of
pupil. vomer horseshoe-shaped, with two rows of equally
sized teeth up to 1/2 diameter of pupil. Palatine with two
rows of equally sized teeth up to 1/2 diameter of pupil.
Dentary with three outer rows of granular teeth and one
inner row of larger teeth anteriorly, merging into one row
of larger teeth posteriorly, up to about size of 3/4 of pupil
diameter.
Otolith (Fig. 33M–P). very elongate in shape, length
to height 2.3–2.5 (30–51 mm Sl) and moderately thin
(otolith height to otolith thickness about 2.5). Anterior tip
moderately pointed, more pointed and longer in smaller
specimens, posterior tip expanded, moderately pointed.
Dorsal rim with broadly rounded predorsal angle and
slightly more pronounced postdorsal angle (particularly
in smaller specimens, Fig. 33P), small concavity above
anterior tip in large specimens, almost straight in small
specimens, concavity behind postdorsal angle only in
small specimens; ventral rim gently and regularly curved,
deepest in front of its middle. Inner face moderately
convex, outer face concave, both smooth. Otolith length to
sulcus length 1.8–2.1 in large specimens, 2.1–2.25. Sulcus
positioned slightly anteriorly, inclined about 5° toward
otolith axis, colliculi separate, ostium much longer and
wider than cauda, ostium length to cauda length 3.5–4.5.
ventral furrow long, close and parallel to ventral rim of
otolith.
Axial skeleton. Neural spine of vertebra 4 inclined
and 5–8 depressed. Parapophyses present from vertebrae
(6) 7 to 11. Pleural ribs on vertebrae 2 to 10. First anal in
pterygophore elongated, but usually not reaching tip of
last precaudal parapophysis in males and less elongated
in females.
Male copulatory organ (Fig. 33F–l). Two pairs of
moderately large, free pseudoclaspers. Inner pseudoclasper
well hidden in pocket of broad isthmus, often unrecognised
when hood bent forward, only visible when freed with
tweezers from pocket (Fig. 33G–h). Outer pseudoclasper
simple, flap-shaped with pointed tip, broad at base,
short, only slightly longer than inner pseudoclasper;
inner pseudoclasper broad, with three ridges in anterior,
posterior and inward directions. Penis with tip strongly
bent at about 90°, slightly longer than pseudoclaspers,
with broad base.
Colour. live colour dull yellow (BPBM 20202).
Preserved colour light brown.
Comparison. Mascarenichthys heemstrai co-occurs
with Dinematichthys indicus Machida, 1994 (identiication
subject to review of the genus Dinematichthys), which
in fact is much more common. It is distinguished by the
lack of scales on the operculum (vs head entirely scaled
in D. indicus, albeit incomplete in juveniles, which are
similar in size with adults of M. heemstrai), presence
of an upper preopercular pore (vs absent) and the very
speciic male copulatory organ. From Majungaichthys
simplex, which also occurs along the Mascarene chain
(Agalega Islands), it differs in the number of precaudal
vertebrae (11 vs 12), the male copulatory organ and the
elongate otolith (otolith length to height 2.3–2.5 vs 2.1)
with separate colliculi (vs fused).
It differs from the South African M. microphthalmus
sp. nov. in the larger eye (1.5–2.7 % Sl vs 0.8–1.2 % Sl)
and the fact that the outer pseudoclasper is slightly longer
than the inner pseudoclasper (vs shorter than the inner
pseudoclasper).
Remarks. The specimens of M. heemstrai from the
four main areas of distribution (Seychelles, Cargados
Carajos, Mauritius and reunion) seem to differ statistically
from each other, but not to the extent that would warrant
separation into individual species in our opinion. For
instance, the specimens from Cargados Carajos (Fig. 33D–
F, l, P) seem to mature at smaller sizes and also do not
seem to grow to the same size as those from the other
areas. Their head proile is often more blunt and the ish
appear more slender. Specimens from Mauritius and
reunion (Fig. 33B, C, O) appear more thick-headed than
those from the Seychelles (Fig. 33A, G–K, M–N) and with
their otoliths tending to be more compressed. The single
specimen from reunion (Fig. 33B) is further characterised
by a very wide maxilla.
Distribution. Seychelles, Cargados Carajos, Mauritius
and reunion (Fig. 29); not known from the Agalega
Islands.
Ecology. Mascarenichthys heemstrai is most commonly
observed in catches from the back reef lagoonal and tide
pool environments, whereas it is rare to nearly absent from
the main and outer reef environments, which in fact is the
domain of Dinematichthys indicus.
Etymology. Named in honour of Phillip C. heemstra,
Grahamstown, SAIAB, for his many contributions to the
knowledge of the ishes of South and east Africa.
Mascarenichthys microphthalmus
(Figs 29, 34, 35; Tables 2, 17)
Material examined. (2 specimens, 38–40 mm Sl).
hOlOTyPe – ruSI, 8548, male, 40 mm Sl, 33°59’S,
81
W. Schwarzhans and P. r. Møller
25°40’e, Bird Island, Algoa Bay, South Africa, coll. J.l.B.
Smith, 11 May 1965. PArATyPe – SAIAB 80185, 1
female, 38 mm Sl, same data as holotype.
Diagnosis. vertebrae 11+29-30 = 40-41, dorsal in
rays 67–69, anal in rays 52–55, pectoral in rays 19, D/A
19–22, v in D 2.0; anterior nostril positioned 1/4 the
distance from upper lip to aggregate distance to anterior
margin of eye; eyes small (0.8–1.2 % Sl); two pairs of free
pseudoclaspers, outer pseudoclasper simple, lap-shaped
with pointed tip, broad at base, short, slightly shorter than
inner pseudoclasper, inner pseudoclasper broad, with
three ridges in anterior, posterior and inward directions
and hidden in pocket of wide isthmus (in resting position);
cheeks with six rows of scales on upper part and three rows
on lower part; upper preopercular pore present.
Description (Figs 34, 35).The principal meristic and
morphometric characters are shown in Table 17. Body
slender; mature at about 35 to 40 mm Sl. head with
scale patch on cheek containing six vertical scale rows on
upper part and three vertical rows on lower. horizontal
diameter of scales on body about 0.8 % Sl, in 15 horizontal
rows. Maxillary ending far behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded, angular. Anterior nostril positioned low, 1/4 the
distance from upper lip to aggregate distance to anterior
margin of eye. Posterior nostril small, about 1/3 the size
of eye. Opercular spine with free tip, pointed. Anterior
gill arch with 14–15 (15) rakers, thereof 3 elongate rakers.
Pseudobranchial ilaments 2.
Head sensory pores (Fig. 35A). Supraorbital pores
2. 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);
irst anterior mandibular pore with or without cirrus.
Preopercular pores: 3 lower, irst and second with joined
opening; third non-tubular; tubular upper preopercular
pore. [See description of Beaglichthys bleekeri for position
of pores.]
Dentition (of holotype). Premaxilla with one outer
row of granular teeth and two inner rows of larger teeth.
Anteriormost teeth in inner row up to size of diameter of
pupil. vomer horseshoe-shaped, with two rows of equally
sized teeth up to 1 1/4 diameter of pupil. Palatine with
two rows of equally sized teeth up to size of diameter of
Table 17. Meristic and morphometric characters of Mascarenichthys
microphthalmus sp. nov. (South Africa) and Mascarenichthys sp.
(Tanzania).
M. microphthalmus Mascarenichthys
sp. nov.
sp.
holotype Paratype
2 non-types
ruSI SAIAB
uSNM 366528
8548
80185
40
38
41-49
Standard length in mm
Meristic characters
Dorsal in rays
67
69
Caudal inrays
16
16
Anal in rays
52
55
Pectoral in rays
19
19
Precaudal vertebrae
11
11
Caudal vertebrae
29
30
Total vertebrae
40
41
rakers on anterior gill arch
15
14
Pseudobranchial ilaments
2
2
D/v
7
7
d/a
22
19
v/A
14
14
Morphometric characters in % of SL
head length
25.4
24.3
head width
9.9
9.6
head height
13.1
11.8
Snout length
4.5
5.0
upper jaw length
12.5
12.4
Diameter of pigmented eye
0.8
1.2
Diameter of pupil
0.5
0.9
Interorbital width
5.7
5.9
Posterior maxilla height
3.7
3.7
Postorbital length
19.7
18.1
Preanal length
47.7
47.9
Predorsal length
30.6
31.1
Body depth at origin
12.7
12.2
of anal in
Pectoral in length
13.9
13.3
Pectoral in base height
5.4
5.7
ventral in length
24.5
21.5
Base ventral in –
30.1
29.0
anal in origin
69-70
16
53
18
11
29
40
13
2
6
19-21
13-14
24.6-25.7
11.4-12-5
14.6-15.6
5.6-6.1
12.9-13.2
1.6-2.0
1.1
6.0-6.2
4.1-4.3
17.6-19.3
48.3-49.3
32.4-34.0
15.7-16.2
14.5-15.4
5.7-5.9
20.1-23.7
30.0-30.2
pupil. Dentary with three outer rows of granular teeth and
one inner row of larger teeth anteriorly, merging into one
row of larger teeth posteriorly, up to about size of 1 1/2
of pupil diameter.
Fig. 34. Mascarenichthys microphthalmus sp. nov., ruSI 8548, holotype, 40 mm Sl.
82
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 35. Mascarenichthys microphthalmus sp. nov. holotype. A, lateral view of head; B, view of left pseudoclasper from inside.
Otolith. Not known.
Axial skeleton. Neural spine of vertebra 4 inclined
and 5–8 depressed. Parapophyses present from vertebrae
(6) 7 to 11. Pleural ribs on vertebrae 2 to 10. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis in males but not in females.
Male copulatory organ (Fig. 35B). Two pairs of
moderately large, free pseudoclaspers. Inner pseudoclasper
well hidden in pocket of broad isthmus, often unrecognised
until hood bent forward and only visible when freed with
tweezers from pocket. Outer pseudoclasper simple lapshaped with pointed tip, broad at base, short, slightly
shorter than inner pseudoclasper, inner pseudoclasper
broad, with three ridges in anterior, posterior and inward
direction. Penis with tip strongly bent at about 90°, slightly
longer than pseudoclaspers, with broad base.
Colour. live colour unknown. Preserved colour in
somewhat dried-up specimen dark brown.
Comparison. Mascarenichthys microphthalmus
resembles M. heemstrai, differing merely in the small eye
size (0.8–1.2 % Sl vs 1.5–2.7 % Sl) and the short outer
pseudoclasper (outer pseudoclasper shorter than inner
pseudoclasper vs longer than inner pseudoclasper).
Distribution. Known from two specimens from Bird
Island in the Algoa Bay of South Africa (Fig. 29).
Etymology. From micros (Greek = small) and
ophthalmos (Greek = eye), referring to the small eye size
of the species. A noun in apposition.
Mascarenichthys sp.
(Figs 29, 36; Tables 2, 17)
Material examined. (2 specimens, 41–49 mm Sl).
Non-types: uSNM 366528, 2 females, 41–49 mm Sl,
6°54’S, 39°55’e, latham Island between Zanzibar and
Maia Islands, Tanzania, h. A. Fehlmann, 20 Nov. 1964.
Remarks. Two female specimens from Tanzania
likely represent an undescribed species characterized
by an interrupted scale patch on the cheeks with six
vertical rows of scales on the large patch on the upper
cheek and two rows on the small patch on the lower
cheek and an otolith with a deepened sulcus in which
ostial and caudal colliculi are fused, but distinction of
Fig. 36. Mascarenichthys sp. uSNM 366528, female, 49 mm Sl. A, lateral view of head; B, ventral view of head; C, ventral view of right
otolith; D, median view of right otolith.
83
W. Schwarzhans and P. r. Møller
both is still possible through an indention of the ventral
sulcus margin.
WAM P.29685-001, 2 males, 48–49 mm Sl, 35°01’S,
150°46’e, Jervis Bay, New South Wales, Australia; ZMuC
P771623-24, 1 male, 44 mm Sl, 1 female, 51 mm Sl, same
data as AMS I.19103-008.
Diagnosis. See generic diagnosis.
Description (Figs 37, 38). The principal meristic and
morphometric characters are shown in Table 18. Body
and head very slender, highest position far behind head;
mature at about 40 mm Sl. head with scale patch on cheek
containing four vertical rows of scales on the upper part
and three on the lower part. horizontal diameter of scales
on body about 1.0 % Sl, in 22 horizontal rows (in 49 mm
Sl male non-type). Maxillary ending far behind eyes,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, angular. Anterior nostril
positioned low, 1/4 the distance from upper lip to aggregate
distance to anterior margin of eye. Posterior nostril small,
about 1/4 the size of eye. Opercular spine with free tip,
sharply pointed. Anterior gill arch with 8–15 (11) rakers,
Monothrix Ogilby, 1897
(Tables 1, 2, 18)
Monothrix Ogilby, 1897: 87 (type-species M. polylepis
Ogilby, 1897, by monotypy); Cohen and Nielsen, 1978:
60; Paxton et al. 1989: 317; Nielsen and Cohen in Nielsen
et al. 1999: 116, 133.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed low
on snout; male copulatory organ with single, simple and
small lap-like pair of (outer) pseudoclaspers; small-sized
(<55 mm Sl), predorsal length short (26.5–30.7 % Sl);
precaudal vertebrae 13–14, total vertebrae 45–47, dorsal
in rays 93–104, anal in rays 64–76, branchiostegal rays
6–7, v in D 2.4–2.6; head with scale patch only on cheek,
no scales on operculum; upper preopercular pore present;
irst and second lower preopercular pore with joined
opening; otolith elongate (otolith length to height 2.3–2.4),
its sulcus with nearly straight dorsal and convex ventral
rim, short (otolith length to sulcus length 2.0), colliculi
fused; maxilla expanded posteroventrally; anterior anal
in pterygiophore short.
Comparison. Monothrix belongs with those genera
having a single pair of (outer) pseudoclaspers and otoliths
with fused colliculi like Brosmolus, Didymothallus and
the New World Gunterichthys. however, the latter two are
characterised by the presence of two almost equally long
supporters in their single pseudoclaspers, a fairly unique
character in Dinematichthyini. Monothrix differs from
Brosmolus in the presence of an upper preopercular pore
(vs absent) and lower vertebrae and dorsal in ray counts
(45–47 and 93–104 vs 56–59 and 124–129).
Distribution. Monothrix is monotypic, distributed
along the shores of south-eastern Australia, from the Port
Phillip Bay to Sydney (Fig. 10).
Table 18. Meristic and morphometric characters of Monothrix
polylepis Ogilby, 1897.
holotype
aMS
I.3654
Standard length in mm
50
Meristic characters
Dorsal in rays
103
Caudal inrays
14
Anal in rays
76
Pectoral in rays
22
Precaudal vertebrae
14
Caudal vertebrae
44
Total vertebrae
47
rakers on anterior gill arch
11
Pseudobranchial ilaments
1
D/v
6
d/a
33
v/A
16
Morphometric characters in % of SL
head length
23.8
head width
13.5
head height
13.8
Snout length
5.4
upper jaw length
11.8
Diameter of pigmented eye
2.5
Diameter of pupil
1.5
Interorbital width
6.2
Posterior maxilla height
4.3
Postorbital length
16.7
Preanal length
48.2
Predorsal length
27.5
Body depth at origin of anal in
14.2
Pectoral in length
13.0
Pectoral in base height
4.5
ventral in length
Base ventral in – anal in origin
26.2
Monothrix polylepis Ogilby, 1897
(Figs 10, 37, 38; Tables 2, 18)
Monothrix polylepis Ogilby, 1897: 88; Cohen and
Nielsen, 1978: 60; Paxton et al. 1989: 317; Nielsen and
Cohen in Nielsen et al. 1999: 134.
Material examined. (31 specimens, 29–53 mm Sl).
hOlOTyPe – AMS I.3654, male, 50 mm Sl, 33°57’S,
151°16’e, Maroubra, Sydney, New South Wales, Australia,
depth unknown, coll. T. Whitelegge, 1897.
Additional specimens. AMS I.15912-048, 1 female, 32
mm Sl, 35°01’S, 150°46’e, Jervis Bay, New South Wales,
Australia; AMS I.16849-034, 1 male, 46 mm Sl, 35°08’S,
150°45’e, Jervis Bay, Australia; AMS I.19103-008, 1 male,
46 mm Sl, 21 females, 29–53 mm Sl, 33°51’S, 151°16’e,
Sydney harbour, New South Wales, Australia; NMv
A3208, 1 female, 34 mm Sl, Portsea, Port Phillip Bay,
victoria, Australia; NMv A17791, 1 female, 33 mm Sl,
38°13’S, 145°01’e, Port Phillip Bay, victoria, Australia;
84
holotype +
non-types
n
Mean (range)
29-53 (41.3)
31
97.8 (93-104)
14.1 (13-15)
70.3 (64-76)
22.8 (22-24)
13.5 (13-14)
32.6 (31-34)
46.1 (45-47)
10.7 (8-15)
0.6 (0-1)
6.3 (6-7)
31.3 (28-35)
15.9 (15-17)
31
10
31
12
31
31
31
11
11
31
31
31
24.4 (23.0-27.5)
11.0 (8.7-13.5)
13.4 (12.1-14.8)
5.4 (4.9-6.6)
11.8 (11.0-13.1)
2.6 (2.0-3.5)
1.4 (1.1-1.7)
6.3 (5.4-7.1)
3.9 (3.5-4.3)
16.8 (16.1-18.8)
47.5 (44.6-50.2)
28.9 (26.5-30.7)
13.0 (11.5-14.2)
13.4 (12.1-15.0)
5.2 (4.5-6.0)
18.6 (15.0-21.3)
28.8 (24.1-31.7)
12
11
11
11
11
11
12
11
11
11
11
12
11
11
11
10
11
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 37. Monothrix polylepis Ogilby, 1897. WAM P.29685-001, male, 50 mm Sl.
Fig. 38. Monothrix polylepis Ogilby, 1897. A, lateral view of head, WAM P.29685-001, male, 49 mm Sl; B, ventral view of head, WAM
P.29685-001, male, 49 mm Sl; C, median view of right otolith, WAM P.29685-001, male, 49 mm Sl; D, ventral view of right otolith,
WAM P.29685-001, male, 49 mm Sl; E, view of left pseudoclasper from inside, AMS I.16849-034, 46 mm Sl; F, inclined lateral view
of male copulatory organ, WAM P.29685-001, male, 49 mm Sl.
thereof 1–2 (2) elongate rakers. Pseudobranchial ilaments
0–1 (1).
Head sensory pores (Fig. 38A, B). Supraorbital
pores 3. Infraorbital pores 6 (3 anterior and 3 posterior):
three posterior pores nearly the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior).
Preopercular pores: 3 lower, irst and second with joined
opening; third tubular; upper preopercular pore tubular.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of a 49 mm Sl male non-type). Premaxilla
with two outer rows of granular teeth and one inner rows
of larger teeth. Anteriormost teeth in inner row up to 1/2
diameter of pupil. vomer horseshoe-shaped, with two rows
of equally sized teeth up to 1/2 diameter of pupil. Palatine
teeth in a single up to 1/3 diameter of pupil. Dentary with
two outer rows of granular teeth and one inner row of
larger teeth anteriorly, merging into one row of larger teeth
posteriorly, up to 3/4 of pupil diameter.
Otolith (Fig. 38C, D). elongate in shape, length to
height 2.3–2.4 (48–53 mm Sl) and moderately thin
(otolith height to otolith thickness about 2.3). Anterior
tip moderately rounded, posterior tip broadly expanded.
Dorsal rim gently and rather regularly curved with rounded
predorsal and slightly more pronounced postdorsal angles;
ventral rim gently and regularly curved, deepest at about
its middle. Inner face moderately convex, outer face lat,
both smooth. Sulcus short, otolith length to sulcus length
2.0. Sulcus positioned anterior of the centre of inner face,
with fused colliculi, not inclined to otolith axis; its dorsal
rim nearly straight and ventral rim convex. ventral furrow
distinct, close to ventral rim of otolith.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–9 depressed. Parapophyses present from vertebrae 7
85
W. Schwarzhans and P. r. Møller
the functional analogy with the pseudoclaspers and the
speciic shape of the inner pseudoclasper.
to 13 (14). Pleural ribs on vertebrae 2 to 13 (14). First anal
in pterygophore not or only slightly elongated.
Male copulatory organ (Fig. 38e–F). A single pair of
small, triangular, lap like (outer) pseudoclaspers widely
connected around penis. Penis curved, about twice as long
as pseudoclaspers.
Colour. live colour unknown. Preserved colour
medium to dark brown.
Comparison. See comparison between Monothrix
and other genera.
Distribution. See genus Monothrix (Fig. 10).
Ungusurculus collettei sp. nov.
(Figs 39–41; Tables 2, 19)
Material examined. (2 specimens, 29–34 mm Sl).
hOlOTyPe – uSNM 374166, male, 34 mm Sl, 6°41’S,
147°53’e, Tami Islands, off Finschhafen, huon Peninsula,
south-eastern New Guinea, coll. B. B. Collette et al., 19
June 1979. PArATyPe – uSNM 365821, 1 male, 29
mm Sl, 1°09’S, 144°22’e, Ninigo Atoll, hermit Island
Group, Papua New Guinea, coll. v. Springer et al., 26
Oct. 1978.
Diagnosis. vertebrae 12+29=41, dorsal in rays 76–78,
anal in rays 54–61, D/A 25–27, v in D 2.2; anterior nostril
positioned 1/6 the distance from upper lip to aggregate
distance to anterior margin of eye; two pairs of free
pseudoclaspers, outer pseudoclasper wing-shaped with
broad base and distally widened supporter with strong
anterior hook (covered by ligament), inner pseudoclasper
stalked, with slightly widened base and distally bifurcate
claw-like with two strong outwardly directed spines;
cheeks with ive rows of scales on upper part and three
rows on lower part; upper preopercular pore absent; otolith
compressed, otolith length to otolith height = 1.8, with
undivided short sulcus, its centre anterior of centre of
otolith, inclined about 5°, otolith length to sulcus length
= 2.25.
Description (Figs 40, 41). The principal meristic and
morphometric characters are shown in Table 19. Body
moderately elongate with pointed snout; small, mature at
at less than 30 mm Sl. head with scale patch on cheek
containing ive vertical scale rows on upper part and three
Ungusurculus gen. nov.
(Tables 1, 2, 19–23)
Type species: Ungusurculus riauensis sp. nov. Gender
masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed low
on snout; male copulatory organ with two pairs of
pseudoclaspers, outer pseudoclasper wing-shaped with
massive supporter, often distally expanded with anteriorly
pointing hook below ligament cover, inner pseudoclasper
free, slightly shorter than outer pseudoclasper, massive,
bifurcate to claw-like; small sized, not much exceeding
55 mm Sl; precaudal vertebrae mostly 12 (12–13), total
vertebrae 40–44, branchiostegal rays 6–7; head with
scale patch only on cheek, no scales on operculum; upper
preopercular pore absent; otolith moderately compressed
to elongate (otolith length to height 1.8–2.4), sulcus
inclined (5–10°), colliculi more or less completely fused,
sulcus short (otolith length to sulcus length 2.2–2.4);
maxilla expanded posteroventrally.
Comparison. In males, Ungusurculus is well
characterised by the morphology of the pseudoclaspers,
the outer with a massive supporter and its anterior hook
and the free inner pseudoclasper with the bifurcate,
claw-like shape. very similar outer pseudoclaspers are
found in eusurculus, but their inner pseudoclaspers are
sucker-disk-like. For other differences see discussion of
eusurculus.
The only other genus sharing with Ungusurculus the
presence of two pairs of pseudoclaspers, the absence of an
upper preopercular pore and fused colliculi of the sulcus
of the otolith is Beaglichthys, from which it differs in the
free, claw-like inner pseudoclasper (vs anteriorly joined to
outer pseudoclasper), the mostly lower number of dorsal
in rays (70–87 vs 84–113) and the short sulcus (otolith
length to sulcus length 2.2–2.4 vs 1.7–2.2).
Species. Six species – U. collettei sp. nov., U. komodoensis
sp. nov., U. philippinensis sp. nov., U. riauensis sp. nov.,
U. sundaensis sp. nov. and U. williamsi sp. nov. – mostly
from Philippine and Indonesian waters, but one species
(U. collettei sp. nov.) from south-eastern New Guinea.
Etymology. Combined from ungulus (latin = claw)
and surculus (latin = grapevine tendril), referring to
Fig. 39. Sample sites of Ungusurculus collettei sp. nov.,
U. komodoensis sp. nov., U. philippinensis sp. nov., U. riauensis sp.
nov., U. sundaensis sp. nov. and U. williamsi sp. nov. One symbol
may represent several samples.
86
Dinematichthyine ishes of the Indo-west Paciic III
.Fig. 40. Ungusurculus collettei sp. nov. uSNM 374166, holotype, 34 mm Sl.
Fig. 41, Ungusurculus collettei sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male
copulatory organ with hood not bent forward, uSNM 365821, 29 mm Sl; D, inclined lateral view of male copulatory organ, holotype;
E, ventral view of male copulatory organ with hood bent forward, uSNM 365821, 29 mm Sl; F, view of left pseudoclasper from inside,
uSNM 365821, 29 mm Sl; G, view of left pseudoclasper from inside, holotype; H, median view of right otolith, holotype; I, ventral
view of right otolith, holotype.
87
W. Schwarzhans and P. r. Møller
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with three outer
rows of granular teeth and one inner row of larger teeth.
Anteriormost teeth in inner row up to 1/2 diameter of
pupil. vomer horseshoe-shaped, with one row of teeth up
to 1/4 diameter of pupil. Palatine with one row of teeth up
to 1/3 diameter of pupil. Dentary with three outer rows of
granular teeth and 1 inner row of larger teeth anteriorly,
merging into one row of larger teeth posteriorly, up to
about size of 2/3 of pupil diameter.
Otolith (Fig. 41h, I). Compressed in shape, length to
height 1.8 (34 mm Sl) and moderately thin (otolith height
to otolith thickness about 2.4). Anterior tip moderately
pointed, posterior tip slightly expanded. Dorsal rim
with broadly rounded predorsal angle and slightly more
pronounced postdorsal angle, section in between straight,
small concavity above anterior tip and behind postdorsal
angle; ventral rim gently and regularly curved, deepest
at about its middle. Inner face moderately convex, outer
face lat, both smooth. Otolith length to sulcus length 2.25.
Sulcus positioned slightly anteriorly, inclined about 5°
towards otolith axis, sulcus undivided with fused colliculi.
ventral furrow short, distinct, moderately close to ventral
rim of otolith, anteriorly and posteriorly turning upward
towards tips of sulcus.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 depressed. Parapophyses present from vertebrae
6 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis in males (no in females known).
Male copulatory organ (Fig. 41C–G). Two pairs of
large, free pseudoclaspers, outer pseudoclasper wingshaped with broad base and distally widened supporter
with strong anterior hook (covered by ligament); inner
pseudoclasper stalked, with slightly widened base
and distally bifurcate and claw-like with two strong,
moderately long outwards directed spines. Penis slightly
curved, slightly longer than pseudoclaspers, pointed, with
broad base.
Colour. live colour unknown. Preserved colour medium
brown with darker back in front of the dorsal in.
Comparison. Ungusurculus collettei resembles
U. komodoensis sp. nov., U. riauensis sp. nov. and
U. sundaensis sp. nov. in the bifurcate claw-like inner
pseudoclasper. It differs from U. komodoensis sp. nov. in the
longer spines of the claw-like inner pseudoclasper and the
low number of vertebrae (41 vs 44). Ungusurculus riauensis
sp. nov. and U. sundaensis sp. nov. both bear a prominent
knob on the inner face of the outer pseudoclasper that its
into the opening of the claw-like inner pseudoclasper, a
character which is lacking in U. collettei. Ungusurculus
collettei further differs from U. riauensis sp. nov. in the
higher D/A (25–27 vs 20–23) and the speciic shape of the
otolith (see description of U. riauensis sp. nov.), and from
Table 19. Meristic and morphometric characters of Ungusurculus
collettei sp. nov. and U. komodoensis sp. nov.
U. komodoensis
sp. nov.
holotype Paratype
holotype
uSNM uSNM
uF
374166 365821
17420
34
29
33
U. collettei sp. nov.
Standard length in mm
Meristic characters
Dorsal in rays
78
Caudal inrays
15
Anal in rays
61
Pectoral in rays
21
Precaudal vertebrae
12
Caudal vertebrae
29
Total vertebrae
41
rakers on anterior gill arch
14
Pseudobranchial ilaments
2
D/v
6
d/a
25
v/A
14
Morphometric characters in % of SL
head length
26.6
head width
13.1
head height
15.7
Snout length
5.1
upper jaw length
12.7
Diameter of pigmented eye
2.3
Diameter of pupil
1.3
Interorbital width
6.5
Posterior maxilla height
4.5
Postorbital length
18.9
Preanal length
52.4
Predorsal length
32.4
Body depth at origin of
16.4
anal in
Pectoral in length
17.3
Pectoral in base height
7.3
ventral in length
17.6
Base ventral in –
33.6
anal in origin
76
15
54
19
12
29
41
14
1
6
27
16
80
14
61
20
12
32
44
16
2
6
26
16
25.1
11.8
15.4
4.9
12.2
2.0
1.0
6.4
3.7
18.3
48.4
31.6
24.8
12.8
15.8
4.7
12.8
2.7
1.5
6.0
3.7
18.9
47.4
32.9
11.7
12.9
12.8
4.9
-
15.3
5.3
20.1
33.7
31.1
vertical rows on lower. horizontal diameter of scales on
body about 2.1 % Sl, in 20 horizontal rows. Maxillary
ending far behind eyes, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded, angular.
Anterior nostril positioned low, 1/6 the distance from
upper lip to aggregate distance to anterior margin of eye.
Posterior nostril small, about 1/6 the size of eye. Opercular
spine with free tip, moderately pointed. Anterior gill arch
with 14 rakers, thereof 3 elongated. Pseudobranchial
ilaments 1–2 (2).
Head sensory pores (Fig. 41A, B). Supraorbital
pores 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 mandibular pore with cirrus. Preopercular
pores: 3 lower, irst and second with joined or separate
opening; third non-tubular; no upper preopercular pore.
88
Dinematichthyine ishes of the Indo-west Paciic III
U. sundaensis sp. nov. in the compressed otolith (otolith
length to otolith height 1.8 vs 2.3–2.4).
The two remaining species U. philippinensis sp. nov.
and U. williamsi sp. nov. do not have the distally widened
supporter in the outer pseudoclasper with the strong
anterior hook below the ligament and the furcate tips
of the inner pseudoclaspers are directed inwards, not
outwards.
Ungusurculus collettei co-occurs with eusurculus
pristinus, from which it differs in the absence of an upper
preopercular pore (although present on one side of the
paratype specimen) (vs present), the strong claw-like inner
pseudoclasper (vs miniature sucker-disk with two small
hooks) and a compressed otolith (otolith length to otolith
height 1.8 vs 2.0–2.1) with fused colliculi (vs distinct
ostium and cauda, but partly fused colliculi).
Distribution. Ungusurculus collettei is the most
eastward occurring species of the genus, known from
south-eastern Papua New Guinea and the hermit Islands
off northern New Guinea (Fig. 39).
Etymology. Named in honour of Bruce B. Collette,
Washington D.C., uSNM, collector of the holotype, for
his many contributions to ichthyology.
Ungusurculus komodoensis sp. nov.
(Figs 39, 42, 43; Tables 2, 19)
Material examined. (1 specimen, 33 mm Sl).
hOlOTyPe – uF 17420, male, 33 mm Sl, Komodo
Island, Indonesia, coral reef at Telek Selawi, depth
0–1.2 m, coll. W. Auffenberg, January 1970.
Diagnosis. vertebrae 12+32=44, dorsal in rays 80, anal
in rays 61, D/A 26, v in D 2.1; anterior nostril positioned
1/3.5 the distance from upper lip to aggregate distance to
anterior margin of eye; two pairs of free pseudoclaspers,
outer pseudoclasper wing-shaped with broad base and
distally widened supporter with strong anterior hook
(covered by ligament), inner pseudoclasper short, with
broad base, distally with two thin, short spines positioned
oblique to axis of pseudoclasper; head with multiple cirri
on snout; cheeks with six rows of scales on upper part and
four rows on lower; upper preopercular pore absent.
Description (Figs 42, 43). The principal meristic and
morphometric characters are shown in Table 19. Body
slender; unique holotype mature at 33 mm Sl. head with
many small cirri on snout and on lower jaw, scale patch on
cheek containing six vertical scale rows on upper part and
four vertical rows on lower. horizontal diameter of scales
on body about 1.5 % Sl, in 15 horizontal rows. Maxillary
ending far behind eyes, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end slightly expanded.
Anterior nostril positioned low, 1/4 the distance from
upper lip to aggregate distance to anterior margin of eye.
Posterior nostril small, about 1/5 size of eye. Opercular
spine with free tip, pointed. Anterior gill arch with 16
rakers, thereof 3 elongate. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 43A, B). Supraorbital
pores 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 mandibular pore with cirrus. Preopercular
pores: 3 lower, irst and second with joined or separate
opening; third non-tubular; no upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with three outer
rows of granular teeth and one inner row of larger teeth.
Anteriormost teeth in inner row up to 1/2 diameter of pupil.
vomer horseshoe-shaped, with two rows of equally sized
teeth up to 1/4 diameter of pupil. Palatine with two rows;
teeth in innner row larger up to 1/3 diameter of pupil.
Dentary with three outer rows of granular teeth and one
inner row of larger teeth anteriorly, merging into one row
of larger teeth posteriorly, up to about size of 1/2 of pupil
diameter.
Otolith. Not known.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis in male holotype (no females known).
Male copulatory organ (Fig. 43 C-F). Two pairs of free
pseudoclaspers, outer pseudoclasper wing-shaped with
broad base and distally widened supporter with strong
anterior hook (covered by ligament); inner pseudoclasper
short, stout, with broad base, distally with two thin, short
spines, positioned oblique to pseudoclasper axis; outer,
Fig. 42. Ungusurculus komodoensis sp. nov. uF 17420, holotype, 33 mm Sl.
89
W. Schwarzhans and P. r. Møller
Fig. 43. Ungusurculus komodoensis sp. nov. holotype. A, lateral view of head; B, ventral view of head; C, inclined lateral view of male
copulatory organ; D, ventral view of male copulatory organ with hood not bent forward; E, view of left pseudoclasper, inner pseudoclasper
seen from ventral, outer pseudoclasper from inside; F, view of left pseudoclasper from inside.
Etymology. Named after the type locality, Komodo
between Flores and Sumbawa, Indonesia.
anterior spine longer, more flexible, inner, posterior
one very short. Penis curved, twice as long as outer
pseudoclaspers, pointed, with broad base.
Colour. live colour unknown. Preserved colour light
brown.
Comparison. Ungusurculus komodoensis is the only
species of the genus with many cirri on the head (at snout
and jaw). Together with U. philippinensis sp. nov. it is the
species with the highest vertebrae count (44 vs 40–43).
amongst the Ungusurculus species with bifurcate clawlike inner pseudoclaspers, U. komodoensis resembles most
U. collettei in lacking the knob on the inner face of the
outer pseudoclasper itting into the opening of the claw of
the inner pseudoclasper (see U. collettei for details).
Distribution. Ungusurculus komodoensis is known
from a single specimen from Komodo, Nusa Tenggara
Timur Province, Indonesia (Fig. 39).
Ungusurculus philippinensis sp. nov.
(Figs 39, 44, 45; Tables 2, 20)
Material examined. (172 specimens, 15–52 mm Sl).
hOlOTyPe – WAM P.31397-010, male, 37 mm Sl,
12°06’N, 110°51’e, off Talampetan, Busuanga Island,
Palawan, Philippines, coll. G. r. Allen, 11 Feb. 1998.
PArATyPeS – CAS 46045, 1 male, 47 mm Sl, 09°03’N,
122°59’e, southern tip of Negros, Philippines, coll. A.
W. herre, 1948; uSNM 346843, 4 males, 36–43 mm Sl,
2 females, 37–41 mm Sl, 10°37’N, 122°32’e, Guimares
Island, Philippines, coll. J. T. Williams et al., 24 Sept.
1995; uSNM 374184, 4 males, 30–45 mm Sl, 4 females,
24–44 mm Sl, 1 juvenile, 15 mm Sl, 09°03’N, 122°59’e,
southern tip of Negros, Philippine, depth 0–3 m, coll. l.
Knapp et al., 24 April 1979; uSNM 374186, 6 males, 33–41
Fig. 44. Ungusurculus philippinensis sp. nov. WAM P.31397-010, holotype, 37 mm Sl.
90
Dinematichthyine ishes of the Indo-west Paciic III
mm Sl, 10 females, 27–46 mm Sl, 1 juvenile, 25 mm Sl,
9°06’N, 122°55’e, southern tip of Negros, Philippines,
depth 0–2 m, coll. l. Knapp et al., 26 April 1979; uSNM
374190, 8 males, 30–48 mm Sl, 6 females, 34–52 mm
Sl, 9°03’N, 122°59’e, north-eastern Negros, Philippines,
depth 0.3 m, coll. h. Fehlmann et al., 18 May 1979; WAM
P.31397-020, 1 female, 40 mm Sl, same data as holotype;
ZMuC P771625–26, 1 male, 46 mm Sl, 1 female, 48 mm
Sl, same data as uSNM 374190.
Additional specimens. uSNM 160866, 1 female, 47
mm Sl, 8°00’N, 124°00’e, Opol, Mindanao, Philippines,
Albatross cruise, 04 Aug. 1909; uSNM 263698, 10
males, 37–47 mm Sl, 23 females, 26–48 mm Sl, 8°51’N,
123°24’e, Bohol Sea, north of Mindanao, Philippines;
uSNM 263693, 11 males, 34–42 mm Sl, 9 females, 32–45
mm Sl, 9°10’N, 123°26’e, Siquijor Island, Philippines;
uSNM 263696, 5 males, 29–42 mm Sl, 5 females,
29–44 mm Sl, 9°04’N, 123°08’e, southern tip of Negros,
Philippines; uSNM 263699, 22 males and 25 females,
26–50 mm Sl, 9°03’N, 122°59’e, southern tip of Negros,
Philippines; uSNM 263700, 1 male and 2 females, 36–38
mm Sl, 9°04’N, 123°16’e, Apo Island, Philippines; uSNM
344549, 3 females, 28–50 mm Sl, 10°35’N, 122°08’e, off
San Joaquin, Panay Island, Philippines; uSNM 344550, 1
female, 30 mm Sl, 10°30’N, 122°29’e, Guimares Island,
Philippines; uSNM 346178, 2 females, 38–40 mm Sl,
10°28’N, 122°28’e, Guimares Island, Philippines; uSNM
366598, 1 male, 36 mm Sl, 9°13’N, 123°28’e, Siquijor
Island, Philippines.
Diagnosis. vertebrae 12-13+29-32=42-44, dorsal in
rays 78–87, anal in rays 54–66, D/A 22–27, v in D 2.2–2.4;
anterior nostril positioned 1/4 the distance from upper lip
Fig. 45. Ungusurculus philippinensis sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of
male copulatory organ, holotype; D, inclined lateral view of male copulatory organ, CAS 46045, 47 mm Sl; E, view of left pseudoclasper
from inside, CAS 46045, 47 mm Sl; F, view of left pseudoclasper from ventral, holotype; G, median view of right otolith, holotype;
H, ventral view of right otolith, holotype.
91
W. Schwarzhans and P. r. Møller
Table 20. Meristic and morphometric characters of Ungusurculus
philippinensis sp. nov.
holotype
WaM
31397-010
Standard length in mm
37
Meristic characters
Dorsal in rays
80
Caudal inrays
?
Anal in rays
58
Pectoral in rays
20
Precaudal vertebrae
12
Caudal vertebrae
30
Total vertebrae
42
rakers on anterior gill arch
18
Pseudobranchial ilaments
2
D/v
7
d/a
23
v/A
14
Morphometric characters in % of SL
head length
27.2
head width
11.9
head height
15.1
Snout length
5.0
upper jaw length
12.9
Diameter of pigmented eye
2.3
Diameter of pupil
1.5
Interorbital width
5.6
Posterior maxilla height
3.9
Postorbital length
19.5
Preanal length
48.2
Predorsal length
32.0
Body depth at origin of anal in
14.3
Pectoral in length
12.8
Pectoral in base height
6.2
ventral in length
23.3
Base ventral in – anal in origin 31.5
holotype +
50 paratypes
n
Mean (range)
37.3 (15–52)
50
82.2 (78-87)
15.6 (15-16)
61.7 (54-66)
20.3 (19-21)
12.1 (12-13)
30.7 (29-32)
42.8 (42-44)
14.8 (12-18)
2
6.3 (6-7)
24.7 (22-27)
14.8 (14-16)
50
19
50
10
50
50
50
11
9
50
50
50
26.6 (25.8-27.4)
12.6 (11.2-16.1)
15.0 (13.7-16.5)
5.3 (4.8-5.9)
13.0 (12.4-13.7)
2.2 (1.9-2.5)
1.4 (1.2-1.6)
5.5 (4.2-6.1)
4.0 (3.6-4.3)
19.4 (18.4-20.1)
50.0 (48.2-51.2)
32.1 (31.3-33.6)
14.9 (12.5-17.1)
14.5 (12.8-16.3)
5.6 (4.9-6.2)
20.9 (19.0-23.3)
31.5 (29.7-34.1)
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
13
11
to aggregate distance to anterior margin of eye; two pairs
of free pseudoclaspers, outer pseudoclasper wing-shaped
with broad base and distally slightly widened supporter
without anterior hook, inner pseudoclasper moderately
long, with widened base, distally with two inward directed
strong spines; cheeks with 5–6 rows of scales on upper part
and 3–4 rows on lower; upper preopercular pore absent;
otolith moderately elongate, otolith length to otolith height
= 2.0–2.1, anterior and posterior tips pointed, short sulcus
with ventral indentation at joining of ostium and cauda
and partly fused colliculi, its centre anterior to centre of
otolith, inclined about 10°, otolith length to sulcus length
= 2.0–2.1.
Description (Figs 44, 45). The principal meristic and
morphometric characters are shown in Table 20. Body
slender, snout pointed; small sized, mature at about 30
mm Sl. head with scale patch on cheek containing 5–6
(5) vertical scale rows on upper part and 3–4 vertical
rows on lower. horizontal diameter of scales on body
92
about 1.2 % Sl, in 20 horizontal rows. Maxillary ending
far behind eyes, dorsal margin of maxillary covered by
upper lip dermal lobe, posterior end expanded. Anterior
nostril positioned low, 1/5 the distance from upper lip to
aggregate distance to anterior margin of eye. Posterior
nostril small, about 1/4 the size of eye. Opercular spine
with free tip, pointed. Anterior gill arch with 12–18 (18)
rakers, thereof 3–4 (3) elongated. In holotype and few
other specimens, row of long rakers interrupted by single
short raker. Pseudobranchial ilaments 2.
Head sensory pores (Fig. 45A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior): three
posterior pores about 1/2 the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior); irst
anterior mandibular pore with cirrus. Preopercular pores:
3 lower, irst and second with joined or separate opening;
third tubular; no upper preopercular pore. [See description
of Beaglichthys bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with three outer
rows of granular teeth and one inner row of larger teeth
anteriorly. Anteriormost teeth in inner row up to 1/2
diameter of pupil. vomer horseshoe-shaped, with two rows
of small teeth 1/4 diameter of pupil. Palatine with 2 rows of
teeth up to 1/4 diameter of pupil. Dentary with four outer
rows of granular teeth and one inner row of larger, teeth
anteriorly, merging into one row of larger teeth posteriorly,
up to 2/3 of pupil diameter.
Otolith (Fig. 45G, h). Moderately elongate in shape,
length to height 2.0–2.1 (37–40 mm Sl) and moderately
thin (otolith height to otolith thickness about 2.5). Anterior
and posterior tips pointed, posterior tip somewhat
expanded. Dorsal rim with broadly rounded pre- and
postdorsal angles, the latter slightly more pronounced,
small concavity above anterior tip and behind postdorsal
angle; ventral rim gently and regularly curved, deepest at
about its middle. Inner face moderately convex, outer face
slightly concave, both smooth. Otolith length to sulcus
length 2.0–2.1. Sulcus positioned slightly towards anterior,
inclined about 10° towards otolith axis, short, with ventral
indentation at joined of ostium and cauda and partly fused
colliculi. ventral furrow moderately short and distinct,
close to ventral rim of otolith, posteriorly turning upwards
towards tip of sulcus.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11 (12). First anal
in pterygophore elongated, reaching tip of last precaudal
parapophysis in males but not in females.
Male copulatory organ (Fig. 45 C-F). Two pairs of free
pseudoclaspers, outer pseudoclasper wing-shaped with
broad base and distally slightly widened supporter without
anterior hook; inner pseudoclasper moderately long, with
widened base, distally with two robust spines directed
inward and oblique to its long axis, posterior spine longer
than anterior spine. Penis curved, slightly longer than outer
pseudoclaspers, pointed, with broad base.
Dinematichthyine ishes of the Indo-west Paciic III
Colour. live colour unknown. Preserved colour light
grey to brownish grey, anteriorly darker, particularly
dorsally in front of dorsal in.
Comparison. Ungusurculus philippinensis is one of
two species of the genus which lack the anterior hook of
the outer pseudoclasper and have inwardly directed spines
of the inner pseudoclasper – the other being U. williamsi
sp. nov. They are also the only two species known from
the Philippines. Ungusurculus philippinensis differs from
U. williamsi sp. nov. in the larger inner pseudoclasper
with stronger spines (vs less than half the size of the outer
pseudoclasper and weak spines), the ratio otolith length
to otolith height (2.0–2.1 vs 1.8–2.0), the longer sulcus
(otolith length to sulcus length 2.0–2.1 vs 2.2–2.4) with
the indented ventral rim (vs undivided) and the speciic
shape of the dorsal rim of the otolith. Ungusurculus
philippinensis is unique in the genus for its incompletely
fused colliculi and the indentation of the ventral sulcus
rim at the joined of ostium and cauda.
Distribution. Ungusurculus philippinensis is fairly
common along the shores of the central Philippines from
Palawan to Panay, Negros and Mindanao (Fig. 39).
Etymology. Named after their distribution along the
Philippine Islands.
Ungusurculus riauensis sp. nov.
(Figs 39, 46, 47; Tables 2, 21)
Material examined. (22 specimens, 37–56 mm Sl).
hOlOTyPe – ZMuC P771616, male, 42 mm Sl, 1°10’N,
103°43’e, Pulau Sudong, off Singapore, coral reef, tidal
zone, Galathea expedition (1950–52), station 337, coll.
T. Wolff and party, 19 May 1951. PArATyPeS – WAM
P.31305-018, 4 male, 37–47 mm Sl, 7 females, 40–56
mm Sl, 1°N, 104°e, Bintan Island, riau Archipelago,
Indonesia, coll. G. r. Allen and r. Steene, 13 May 1997;
WAM P.31315-002, 1 male, 45 mm Sl, 8 females, 46–55
mm Sl, 1°N, 104°e, Bintan Island, riau Archipelago,
Indonesia, depth 0.2 m, coll. G. r. Allen, 20 May 1997;
ZMuC P771615, 1 female, 48 mm Sl, same data as
holotype.
Diagnosis. vertebrae 12+28-30=40-42, dorsal in rays
70–77, anal in rays 53–61, D/A 20–23, v in D 1.9–2.2;
Table 21. Meristic and morphometric characters of Ungusurculus
riauensis sp. nov.
holotype
ZMuC
P771616
Standard length in mm
42
Meristic characters
Dorsal in rays
75
Caudal inrays
16
Anal in rays
54
Pectoral in rays
19
Precaudal vertebrae
12
Caudal vertebrae
30
Total vertebrae
42
rakers on anterior gill arch
16
Pseudobranchial ilaments
2
D/v
6
d/a
21
v/A
14
Morphometric characters in % of SL
head length
26.9
head width
15.6
head height
16.1
Snout length
6.5
upper jaw length
14.4
Diameter of pigmented eye
2.0
Diameter of pupil
1.5
Interorbital width
6.6
Posterior maxilla height
4.7
Postorbital length
19.5
Preanal length
51.1
Predorsal length
33.7
Body depth at origin of anal in
18.6
Pectoral in length
15.6
Pectoral in base height
5.6
ventral in length
21.7
Base ventral in – anal in origin 29.8
holotype +
21 paratypes
n
Mean (range)
48.0 (37-56)
22
73.7 (70-77)
16
56.6 (53-61)
18.6 (17-20)
12
29.4 (28-30)
41.4 (40-42)
16.0 (15-17)
2
6.0 (6-7)
21.2 (20-23)
14.0 (13-14)
22
12
22
11
22
22
22
10
9
22
22
21
26.1 (24.8-27.4)
13.1 (11.7-15.6)
15.5 (14.7-16.3)
5.4 (4.2-6.5)
13.3 (12.4-14.4)
2.3 (1.9-2.6)
1.3 (1.1-1.5)
5.9 (5.4-6.6)
4.3 (3.9-4.7)
18.8 (17.5-19.8)
49.5 (47.4-51.1)
32.1 (29.5-34.4)
15.5 (14.2-18.6)
14.7 (12.0-19.8)
5.7 (5.5-6.1)
17.5 (12.4-21.7)
31.6 (28.4-34.1)
11
10
10
10
10
12
10
10
10
10
10
10
10
11
10
8
10
anterior nostril positioned 1/4 distance from upper lip
to aggregate distance to anterior margin of eye; two
pairs of free pseudoclaspers, outer pseudoclasper wingshaped with broad base and distally widened supporter
with massive anterior hook, prominent knob on inner
Fig. 46. Ungusurculus riauensis sp. nov. ZMuC P771616, holotype, 42 mm Sl.
93
W. Schwarzhans and P. r. Møller
Fig. 47. Ungusurculus riauensis sp. nov. A, lateral view of head, WAM P.31315-002, 45 mm Sl; B, ventral view of head, WAM P.31315002, 45 mm Sl; C, ventral view of male copulatory organ, WAM P.31305-018, 47 mm Sl; D, inclined lateral view of male copulatory
organ, holotype; E, view of left pseudoclasper from ventral, holotype; F, view of left pseudoclasper from inside, WAM P.31315-002, 45
mm Sl; G, median view of right otolith, WAM P.31305-018, female, 50 mm Sl.
Description (Fig. 46–47). The principal meristic and
morphometric characters are shown in Table 21. Body
slender with pointed snout; mature at about 35 mm Sl.
head with scale patch on cheek containing ive vertical
scale rows on upper part and three vertical rows on
lower. Maxillary ending far behind eyes, dorsal margin
of maxillary covered by upper lip dermal lobe, posterior
end expanded. Anterior nostril positioned low, 1/4.5 the
distance from upper lip to aggregate distance to anterior
margin of eye. Posterior nostril small, about 1/4 the size
of eye. Opercular spine with free tip, pointed. Anterior
face of outer pseudoclasper itting into opening of inner
pseudoclasper, inner pseudoclasper long, massive, stalked,
distally with claw-like, outwardly directed strong spines;
cheeks with ive rows of scales on upper part and three
rows on lower; upper preopercular pore absent; otolith
moderately elongate, otolith length to otolith height =
2.0–2.2, anterior tip pointed, posterior tip expanded,
postdorsal angle sharp, bent outwards, short sulcus with
small ventral indentation at joining of ostium and cauda
but completely fused colliculi, its centre anterior to centre
of otolith, inclined about 10°, otolith length to sulcus
length = 2.1.
94
Dinematichthyine ishes of the Indo-west Paciic III
gill arch with 15–17 (16) rakers, thereof 3 elongate.
Pseudobranchial ilaments 2.
Head sensory pores (Fig. 47A, B). Supraorbital
pores 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 mandibular pore with cirrus. Preopercular
pores: 3 lower, irst and second with joined or separate
opening; third non-tubular; no upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with three outer
rows of granular teeth and one inner row of larger teeth
anteriorly. Anteriormost teeth in inner row up to 2/3
diameter of pupil. vomer horseshoe-shaped, with two rows
of small teeth, 1/3 diameter of pupil. Palatine with two
rows of small teeth up to 1/2 diameter of pupil. Dentary
with three outer rows of granular teeth and one inner row of
larger teeth anteriorly, merging into one row of larger teeth
posteriorly. largest teeth up to size of pupil diameter.
Otolith (Fig. 47G). Moderately elongate in shape,
length to height 2.0–2.2 (47–56 mm Sl) and moderately
thin (otolith height to otolith thickness about 2.5).
Anterior tip pointed, posterior tip somewhat expanded,
less pointed. Dorsal rim with broadly rounded predorsal
angle and sharp, almost spiny postdorsal angle, latter
characteristically bent outwards, small concavity above
anterior tip and deep concavity behind postdorsal angle;
ventral rim gently and regularly curved, deepest at about
its middle. Inner face convex, outer face slightly concave,
both smooth. Otolith length to sulcus length 2.1. Sulcus
positioned slightly towards anterior, inclined with about
10° towards otolith axis, short, with indistinct ventral
indentation at joined of ostium and cauda and completely
fused colliculi. ventral furrow distinct, close to ventral
rim of otolith, posteriorly turning upwards, right-angled
towards tip of sulcus.
Axial skeleton. Neural spine of vertebrae 4 (-5) inclined
and (5) 6–8 depressed. Parapophyses present from
vertebrae 7 to 12. Pleural ribs on vertebrae 2 to 11 (10).
First anal in pterygophore elongated, reaching tip of last
precaudal parapophysis in males but not in females.
Male copulatory organ (Fig. 47C-F). Two pairs of free
pseudoclaspers, outer pseudoclasper wing-shaped with
broad base and distally widened supporter with massive
anterior hook, prominent knob on inner face of outer
pseudoclasper itting into opening of inner pseudoclasper,
inner pseudoclasper long, massive, stalked, distally with
claw-like, outward directed strong spines. Penis curved,
slightly longer than outer pseudoclaspers, pointed, with
broad base.
Colour. live colour unknown. Preserved colour
medium brown.
Comparison. Ungusurculus riauensis shares the
outer pseudoclasper with the distally widened supporter
carrying an anterior hook and the outwardly directed
claw-like tip of the inner pseudoclasper with U. collettei,
U. komodoensis and U. sundaensis sp. nov., and with the
latter the prominent knob on the inner face of the outer
pseudoclasper which its into the opening of the claw-like
tip of the inner pseudoclasper. Ungusurculus riauensis
differs from all other species of Ungusurculus in the low
number of dorsal in rays (mostly 70–77 vs 78–87), the low
D/A (20–23 vs mostly 24–27, 22–27 in U. philippinensis)
and the speciic shape of the otoliths, in particular the spiny
postdorsal angle, which is bent outwards.
Distribution. Ungusurculus riauensis seems to
be restricted to the Indonesian riau Archipelago and
Singapore (Fig. 39).
Etymology. Named after the riau Archipelago, from
where the majority of the investigated specimens were
obtained.
Ungusurculus sundaensis sp. nov.
(Figs 39, 48, 49; Tables 2, 22)
Material examined. (4 specimens, 41–54 mm Sl).
hOlOTyPe – WAM P.30959-018, male, 54 mm Sl,
8°46’S, 119°43’e, rinca Island near Flores, Indonesia,
coll. G. r. Allen, 4 April 1995. PArATyPe – NSMT-P
78775, 1 male, 46 mm Sl, off lombok, Indonesia, coll.
K. Matsuura and K. Shibukawa, 24 July 1996.
Additional specimens. uSNM 366475, 1 female, 41
mm Sl, 5°52’S, 112°37’e, Bawean Island, north of Java,
Indonesia; uSNM 366499, 1 female, 42 mm Sl, 5°46’S,
106°35’e, Seribu Islands, Java Sea, Indonesia.
Fig. 48. Ungusurculus sundaensis sp. nov. WAM P.30959-018, holotype, 54 mm Sl.
95
W. Schwarzhans and P. r. Møller
Fig. 49. Ungusurculus sundaensis sp. nov. holotype. A, lateral view of head; B, ventral view of head; C, ventral view of male copulatory
organ; D, inclined lateral view of male copulatory organ; E, view of left pseudoclasper from ventral, outer pseudoclasper bent away from
inner pseudoclasper; F, median view of right otolith; G, ventral view of right otolith.
anterior of centre of otolith, inclined about 5°, otolith
length to sulcus length = 2.4–2.5.
Description (Figs 48, 49). The principal meristic and
morphometric characters are shown in Table 22. Body
slender with pointed snout; mature at about 45 mm Sl.
head with scale patch on cheek containing 6 vertical scale
rows on the upper part and 4 vertical rows on the lower
part. horizontal diameter of scales on body about 1.2%
Sl, in 23 horizontal rows.
Maxillary ending far behind eyes, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior
end expanded. Anterior nostril positioned low, 1/5 the
distance from upper lip to aggregate distance to anterior
margin of eye. Posterior nostril small, about 1/5 the size
of eye. Opercular spine with free tip, moderately pointed.
Anterior gill arch with 15–16 (16) rakers, thereof three
elongate. row of elongate rakers interrupted by short
Diagnosis. vertebrae 12+30-31=42-43, dorsal fin
rays 79–86, anal in rays 60–64, D/A 26, v in D 2.2–2.3;
anterior nostril positioned 1/4 distance from upper lip to
aggregate distance to anterior margin of eye; two pairs
of free pseudoclaspers, outer pseudoclasper wing-shaped
with broad base and distally widened supporter with
anterior hook, prominent knob on inner face of outer
pseudoclasper itting into opening of inner pseudoclasper,
inner pseudoclasper long, very massive, stalked with
narrow base, distally with claw-like, outwardly directed
strong spines, inner and outer pseudoclasper matching
together in lock/key mode; cheeks with 6 rows of scales
on upper part and 4 rows on lower part; upper preopercular
pore absent; otolith elongate, otolith length to otolith
height = 2.3–2.4, anterior and posterior tips pointed, preand postdorsal angles symmetrical, section in between
straight, short sulcus with fused colliculi, its centre
96
Dinematichthyine ishes of the Indo-west Paciic III
Table 22. Meristic and morphometric characters of Ungusurculus
sundaensis sp. nov.
holotype
WaM
30959-018
54
Paratype
NSMT-P
49905
46
79
16
60
21
12
30
42
16
2
6
26
15
86
15
64
20
12
31
43
15
1
6
26
15
23.0
12.1
13.9
4.9
12.1
2.1
1.4
5.0
3.6
16.9
47.0
28.6
14.8
13.5
5.3
18.7
32.2
25.7
12.1
15.4
5.5
12.5
2.1
1.1
6.5
4.3
18.9
47.6
30.3
15.2
14.9
5.7
20.0
31.5
Standard length in mm
Meristic characters
Dorsal in rays
Caudal inrays
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 SL
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
raker in holotype but not in paratype. Pseudobranchial
ilaments 1–2 (2).
Head sensory pores (Fig. 49A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior): three
posterior pores about 1/2 the size of three anterior pores,
third posterior pore tubular. Mandibular pores 6 (3 anterior
and 3 posterior); irst anterior mandibular pore without
cirrus. Preopercular pores: 3 lower, irst and second with
joined or separate opening; third non-tubular; no upper
preopercular pore. [See description of Beaglichthys
bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with 4 outer rows of
granular teeth and one inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 2/3 diameter of
pupil. vomer horseshoe-shaped, with 3 rows of teeth up
to 1/2 diameter of pupil. Palatine with 3 rows of teeth up
to 1/2 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, up to 3/4
of pupil diameter.
Otolith (Fig. 49F, G). Moderately elongate in shape,
length to height 2.3–2.4 (46–54 mm Sl) and thin (otolith
height to otolith thickness about 2.5). Anterior and posterior
tips pointed, posterior tip slightly expanded. Dorsal rim
with marked pre- and postdorsal angles, section in between
long and straight, small concavity above anterior tip and
behind postdorsal angle; ventral rim gently and regularly
curved, deepest at about its middle. Inner face convex,
outer face slightly concave, both smooth. Otolith length to
sulcus length 2.4–2.5. Sulcus positioned slightly towards
anterior, inclined about 5° towards otolith axis, short, with
fused colliculi. ventral furrow distinct, close to ventral
rim of otolith, posteriorly turning upwards.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–9 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11 (12). First anal
in pterygophore elongated, reaching tip of last precaudal
parapophysis in males.
Male copulatory organ (Fig. 49C–e). Two pairs of
free pseudoclaspers, outer pseudoclasper wing-shaped
with broad base and distally widened supporter with
anterior hook, prominent knob on inner face of outer
pseudoclasper itting into opening of inner pseudoclasper,
inner pseudoclasper long, very massive, stalked with
narrow base, distally with claw-like, outward directed
strong spines, inner and outer pseudoclasper matching
together in lock / key mode. Penis curved, slightly longer
than outer pseudoclaspers, pointed, with broad base.
Colour. live colour unknown. Preserved colour
medium brown with dark brown back.
Comparison. Ungusurculus sundaensis most
resembles U. riauensis in the outer pseudoclasper with the
distally widened supporter carrying an anterior hook and
with a prominent knob on the inner face, which matches
with the outward directed claw-like tip of the inner
pseudoclasper. Ungusurculus sundaensis differs from all
other species of Ungusurculus in the very elongate otolith
(otolith length to otolith height 2.3–2.4 vs 1.8–2.2) and the
short sulcus (otolith length to sulcus length 2.4–2.5 vs
2.0–2.4). For further discussion see comparison of other
species above.
Distribution. Ungusurculus sundaensis is known
from few specimens from the western tip of Java to the
western tip of Flores, along the Sunda Arch of Indonesia
(Fig. 39).
Etymology. Named after the Sunda Arch of Indonesia,
from where all investigated specimens were obtained.
Ungusurculus williamsi sp. nov.
(Figs 39, 50, 51; Tables 2, 23)
Material examined. (12 specimens, 21–45 mm Sl).
hOlOTyPe – uSNM 346941, male, 45 mm Sl, 10°34’N,
122°30’e, Pulang Duta, Guimares Island, Philippines,
small cove lined with lava rock and rocky boulders,
depth 0–4 m, coll. J. T. Williams et al., 24 Sept. 1995.
PArATyPeS – uSNM 384197, 1 male, 35 mm Sl, 8
97
W. Schwarzhans and P. r. Møller
Fig. 50. Ungusurculus williamsi sp. nov. uSNM 346941, holotype, 45 mm Sl.
females, 21–45 mm Sl, 1 juvenile, 21 mm Sl, same data
as holotype; ZMuC P771627-28, 1 male, 35 mm Sl, 1
female, 42 mm Sl, same data as holotype.
Diagnosis. vertebrae 12+30-31=42-43, dorsal in rays
79–85, anal in rays 59–63, D/A 24–26, v in D 2.2–2.4;
anterior nostril positioned 1/4 the distance from upper lip
to aggregate distance to anterior margin of eye; two pairs
of free pseudoclaspers, outer pseudoclasper wing-shaped
with broad base and distally widened supporter without
anterior hook, inner pseudoclasper short, half the length of
outer pseudoclasper, with slightly broadened base, distally
with two inwardly directed weak spines; cheeks with 4–6
rows of scales on upper part and 3 rows on lower part;
upper preopercular pore absent; otolith compressed, otolith
length to otolith height = 1.9–2.1, anterior and posterior
tips pointed, latter expanded, projecting sharp postdorsal
angle, sulcus short, wide with fused colliculi, its centre
slightly anterior of centre of otolith, inclined about 10°,
otolith length to sulcus length = 2.2–2.4.
Description (Figs 50, 51). The principal meristic and
morphometric characters are shown in Table 23. Body
slender; small-sized, mature at about 35 mm Sl. head
with scale patch on cheek containing 4–6 (4) vertical scale
rows on the upper part and 3 vertical rows on the lower
part. horizontal diameter of scales on body about 1.4% Sl,
in 24 horizontal rows. Maxillary ending far behind eyes,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded. Anterior nostril positioned
low, 1/4 the distance from upper lip to aggregate distance
to anterior margin of eye. Posterior nostril small, about 1/4
the size of eye. Opercular spine with free tip, moderately
pointed. Anterior gill arch with 13–15 (13) rakers, thereof
3–4 (3) elongated. Pseudobranchial ilaments two.
Head sensory pores (Fig. 51 A-B). Supraorbital
pores 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 mandibular pore with cirrus. Preopercular
pores: 3 lower, irst and second with joined or separate
opening; third non-tubular; no upper preopercular pore.
[See description of Beaglichthys bleekeri for position of
pores.]
Dentition (of holotype). Premaxilla with 4 outer rows of
granular teeth and one inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 3/4 diameter of
pupil. vomer horseshoe-shaped, with 2 rows of teeth up
to 1/2 diameter of pupil. Palatine with outer row of small
teeth and one inner row of larger teeth up to 1/2 diameter
of pupil. Dentary with 4 outer rows of granular teeth and
1 inner row of larger, slender teeth anteriorly, merging
Table 23. Meristic and morphometric characters of Ungusurculus
williamsi sp. nov.
holotype
uSNM
346941
Standard length in mm
45
Meristic characters
Dorsal in rays
80
Caudal inrays
16
Anal in rays
60
Pectoral in rays
22
Precaudal vertebrae
12
Caudal vertebrae
30
Total vertebrae
42
rakers on anterior gill arch
13
Pseudobranchial ilaments
2
D/v
6
d/a
25
v/A
15
Morphometric characters in % of SL
head length
26.2
head width
11.9
head height
15.2
Snout length
5.0
upper jaw length
13.3
Diameter of pigmented eye
2.2
Diameter of pupil
1.6
Interorbital width
5.7
Posterior maxilla height
4.2
Postorbital length
19.2
Preanal length
49.3
Predorsal length
32.6
Body depth at origin of anal in
15.2
Pectoral in length
15.7
Pectoral in base height
6.0
ventral in length
19.5
Base ventral in – anal in
31.5
origin
98
holotype +
11 paratypes
n
Mean (range)
34.4 (21-45)
12
82.2 (79-85)
16
61.0 (59-63)
21.1 (19-22)
12
30.4 (30-31)
42.2 (42-43)
13.6 (13-15)
2
6.4 (6-7)
24.8 (24-26)
14.5 (14-15)
11
5
11
8
11
11
11
9
9
11
11
11
26.7 (25.5-28.1)
11.5 (10.5-13.0)
14.8 (13.8-15.7)
5.2 (4.6-6.0)
12.6 (11.8-13.3)
2.0 (1.8-2.2)
1.2 (0.9-1.6)
5.7 (5.0-6.2)
3.6 (3.2-4.2)
19.6 (19.0-20.7)
48.5 (45.5-52.9)
32.5 (31.3-35.0)
13.5 (12.0-15.9)
14.9 (11.6-16.7)
5.2 (4.0-6.1)
20.3 (16.5-23.3)
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
10
28.8 (25.1-32.4)
11
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 51. Ungusurculus williamsi sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, view of left pseudoclasper
from inside, holotype; D, view of left pseudoclasper from ventral, uSNM 384197, 35 mm Sl; E, inclined lateral view of male copulatory
organ, holotype; F, median view of right otolith, holotype; G, ventral view of right otolith, holotype.
into 1 row of larger teeth posteriorly, up to 2/3 of pupil
diameter.
Otolith (Fig. 51F, G). Moderately compressed in shape,
length to height 1.9–2.1 (35–45 mm Sl) and moderately
thin (otolith height to otolith thickness about 2.4). Anterior
and posterior tips pointed, posterior tip expanded. Dorsal
rim with shallow predorsal and markedly projecting
postdorsal angles, section in between long, straight,
anteriorly inclined, small concavity above anterior tip and
behind postdorsal angle; ventral rim gently and regularly
curved, deepest at about its middle. Inner face convex,
outer face slightly concave, both smooth. Otolith length to
sulcus length 2.2–2.4. Sulcus positioned slightly towards
anterior, inclined about 10° towards otolith axis, short,
wide, with fused colliculi. ventral furrow distinct, not
very close to ventral rim of otolith, posteriorly turning
upwards.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 depressed. Parapophyses present from vertebrae
(6) 7 to 12. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis in males but not in females.
Male copulatory organ (Fig. 51C–e). Two pairs of free
pseudoclaspers, outer pseudoclasper wing-shaped with
broad base and distally slightly widened supporter without
anterior hook, inner pseudoclasper short, half the length of
outer pseudoclasper, with slightly broadened base, distally
with two inwardly directed weak and short spines. Penis
curved, slightly longer than outer pseudoclaspers, pointed,
with broad base.
Colour. live colour unknown. Preserved colour light
brown.
Comparison. Ungusurculus williamsi resembles
U. philippinensis in the outer pseudoclasper with a
distally slightly widened supporter without an anterior
hook and the inward directed bifurcate tip of the inner
pseudoclasper. It differs in the short and weak inner
pseudoclasper and the otolith morphology (fused colliculi
vs partly fused with indented ventral sulcus margin and
the strongly projecting postdorsal angle, vs shallow and
rounded). The speciic shape of the dorsal rim of the otolith
and the wide sulcus distinguishes U. williamsi from all
other species of Ungusurculus.
99
W. Schwarzhans and P. r. Møller
coll. J. B. hutchins et al., 1 June 1982. PArATyPeS
– AMS I.20241-008, 1 male, 45 mm Sl, 32°01’S,
115°25’e, rottnest Island, Western Australia, depth
6 m, coll. B. C. russell, 11 April 1978; AMS I.20245012, 3 females, 45–48 mm Sl, 1 juvenile, 25 mm Sl,
32°00’S, 115°28’e, rottnest Island, Western Australia,
depth 12–15 m, coll. B. C. russell, 12 April 1978; AMS
I.20247-017, 1 male, 45 mm Sl, 31°59’S, 115°33’e,
rottnest Island, Western Australia, depth 8 m, coll. B.
C. russell, 12 April 1978; NMv A2487, 2 males, 36–43
mm Sl, 2 females, 49–50 mm Sl, 28°29’S, 113°46’e,
Wallabi Island, houtman Abrolhos, Western Australia,
coll. G. r. Allen, 19 April 1982; NMv A20829, 1 male,
49 mm Sl, 33°51’S, 121°55’e, esperance Bay, Western
Australia, coll. r. h. Kuiter, 21 March 1986; uSNM
263733, 1 male, 51 mm Sl, 2 females, 39–51 mm Sl, 1
juvenile, 23 mm Sl, rottnest Island, Western Australia,
coll. J. B. hutchins, 8 March 1977; WAM P.27616-035,
6 males, 29–47 mm Sl, 14 females, 29–58 mm Sl, 7
juveniles, 20–26 mm Sl, same data as holotype; WAM
P.27950-010, 11 males, 34–51 mm Sl, 13 females,
25–61 mm Sl, 30°18’S, 115°00’e, rottnest Island,
Western Australia, depth 4–6 m, coll. N. Sinclair et
al., 9 April 1983; WAM P.27951-008, 3 females, 33–55
mm Sl, 30°18’S, 115°00’e, rottnest Island, Western
Australia, depth 2–5 m, coll. J. B. hutchins et al., 10
April 1983; WAM P.27955-003, 4 males, 35–50 mm Sl,
4 females, 46–60 mm Sl, 29°16’S, 114°55’e, Denison,
Western Australia, depth 9–10 m, coll. J. B. hutchins,
13 April 1983; WAM P.27957-008, 5 males, 42–59 mm
Sl, 3 females, 57–67 mm Sl, 2 juveniles, 24–25 mm
Sl, 29°16’S, 114°55’e, Denison, Western Australia,
depth 7–8 m, coll. J. B. hutchins et al. 14 April 1983;
WAM P. 27959-004, 1 male, 50 mm Sl, 1 female, 53
mm Sl, 27°30’S, 114°25’e, north of Kalbarri, Western
Australia, depth 0–7 m, coll. J. B. hutchins et al., 16
April 1983; WAM P.28517-003, 1 male, 32 mm Sl,
3 females, 59–70 mm Sl, 33°35’S, 115°06’e, Cape
Naturaliste, Western Australia, depth 15–17 m, coll.
J. B. hutchins, 12 April 1985; WAM P.28522-006, 2
females, 40–58 mm Sl, 34°S, 115°e, Flinders Bay, off
Augusta, Western Australia, depth 12–13 m, coll. J. B.
hutchins, 18 April 1985; WAM P.29886-017, 2 males,
47–54 mm Sl, 10 females, 38–62 mm Sl, 3 juveniles,
21–23, 28°55’S, 114°02’e, Pelsart Island, houtman
Abrolhos, Western Australia, depth 7–9 m, coll. G. r.
Allen, 6 March 1988; WAM P.30832-004, 2 males, 38
mm Sl, 28°30’S, 113°44’e, Wallabi Island, houtman
Abrolhos, Western Australia, depth 3–4 m, coll. J. B.
hutchins et al., 27 May 1994; ZMuC P 771640-41, 1
male, 45 mm Sl, 1 female, 50 mm Sl, same data as
WAM P.27950-010.
Additional specimens. uSNM 367147, 3 males, 43–46
mm Sl, Point heron, 30 miles (48.3 km) south of Perth,
Western Australia.
Diagnosis. See generic diagnosis.
Distribution. Ungusurculus williamsi is known from
a single lot collected in a rocky cave of Guimares Island,
Philippines (Fig. 39).
Ecology. Judging from the single location from which
it was collected in reasonable numbers, U. williamsi
is probably adapted to non-reef related rocky inshore
environments.
Etymology. Named in honour of Jeffrey T. Williams,
Washington D.C., uSNM, who has collected the type
specimens, in recognition to his contribution to the
knowledge of the ishes of the Philippines.
Zephyrichthys gen. nov.
(Tables 1, 2, 24)
Type species: Zephyrichthys barryi sp. nov. Gender
masculine.
Diagnosis. Genus of Dinematichthyini with following
combination of characters: Anterior nostril placed
low on snout; male copulatory organ with two pairs of
pseudoclaspers, outer pseudoclasper wing-shaped with
broad base, inner pseudoclasper about half the size of
outer pseudoclasper, bifurcate, each branch with supporter,
anteriorly joined to outer pseudoclasper; moderate in size
reaching about 70 mm Sl; precaudal vertebrae mostly
12–13, total vertebrae 43–46; 7–8 branchiostegal rays;
head with scale patch on cheek, usually 2–3 scales on
operculum above opercular spine (rarely none); upper
preopercular pore absent; otolith moderately compressed
to elongate (otolith length to height 2.1–2.3), sulcus not
inclined, colliculi separated, caudal colliculum very short
(length of ostial colliculum to length of caudal colliculum
5–8); maxilla expanded posterio-ventrally.
Comparison. Zephyrichthys differs from all other
dinematichthyine genera in the Indo-west Pacific in
the bifurcate inner pseudoclasper with a branched
supporter, of which the anterior branch is joined to the
outer pseudoclasper. It shares with Beaglichthys and
Ungusurculus and some species of Dinematichthys the
absence of an upper preopercular pore and presence of
two pairs of pseudoclaspers. With certain species of
Dinematichthys it further shares the presence of scales on
the operculum above the opercular spine (though not in all
specimens in the case of Zephyrichthys) and the separated
colliculi in the sulcus of the otolith. The low anterior nostril
(vs high) and the small caudal colliculum (see above)
distinguish Zephyrichthys from Dinematichthys.
Species. The genus is monotypic.
Etymology. From zephyrus (latin = west wind),
referring to the distribution along the west Australian
coast. A noun in apposition.
Zephyrichthys barryi sp. nov.
(Figs 10, 52, 53; Tables 2, 24)
Material examined. (118 specimens, 20–70 mm Sl).
hOlOTyPe – WAM P.27616-010, male, 49 mm Sl,
32°00’S, 115°30’e, rottnest Island, Western Australia,
100
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 52. Zephyrichthys barryi sp. nov. A, fresh dead, WAM P.27959-004, female, 53 mm Sl; B, WAM P.27616-010, holotype, 49 mm Sl.
Description (Figs 52, 53). The principal meristic and
morphometric characters are shown in Table 24. Body
and head slender; mature at about 40 mm Sl. head
with scale patch on cheek containing 7 vertical scale
rows on the upper part and 3 vertical rows on the lower
part, up to 3 scales on operculum above opercular spine.
horizontal diameter of scales on body about 1.4 % Sl,
in 20 horizontal rows. Maxillary ending far behind eyes,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, angular. Anterior nostril
positioned moderately low, 1/3.5 to 1/4 the distance
from upper lip to aggregate distance to anterior margin
of eye. Posterior nostril small, about 1/5 the size of
eye. Opercular spine with free tip, pointed. Anterior
gill arch with 12–15 (13) rakers, thereof 3 elongated.
Pseudobranchial ilaments 2–3 (2).
Head sensory pores (Fig. 53A, B). Supraorbital
pores 2–3. Infraorbital pores 6–7 (3 anterior, 3 posterior
and occasionally 1 small pore posterior nostril): three
posterior pores about 1/3 the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior); irst
anterior mandibular pore with cirrus. Preopercular pores:
3 lower, irst and second with separate opening; third
tubular; no upper preopercular pore. [See description of
Beaglichthys bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with four outer
rows of granular teeth and one inner row of larger teeth
anteriorly. Anteriormost teeth in inner row up to 1/2
diameter of pupil. vomer horseshoe-shaped, with three
rows of small teeth 1/4 diameter of pupil. Palatine with two
outer rows of small teeth and one inner row of larger teeth
up to 1/3 diameter of pupil. Dentary with four outer rows
of granular teeth and one inner row of larger, slender teeth
anteriorly, merging into one row of larger teeth posteriorly,
up to 2/3 of pupil diameter.
Otolith (Fig. 53h, I). Moderately elongate in shape,
length to height 2.1–2.3 (23–70 mm Sl) and moderately
thick (otolith height to otolith thickness about 2.2).
Anterior tip pointed, posterior tip expanded, pointed.
Dorsal rim with obtuse predorsal and sharp postdorsal
angles, long section in between straight, marked concavity
above anterior tip and behind postdorsal angle; ventral rim
gently and regularly curved, deepest at about its middle.
Inner face convex, outer face lat to slightly concave, both
smooth. Otolith length to sulcus length 1.9–2.1. Sulcus
positioned slightly anterior to middle of inner face, not
inclined, ostium and cauda divided, colliculi separate,
caudal colliculum very short (length of ostial colliculum
to length of caudal colliculum 5–8). ventral furrow long,
distinct, close to ventral rim of otolith, slightly turning
upwards towards its tips.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–8 depressed. Parapophyses present from vertebrae
7 to 12. Pleural ribs on vertebrae 2 to 11. Basis of neural
spines 5–8 enlarged. First anal in pterygophore elongated,
101
W. Schwarzhans and P. r. Møller
Fig. 53. Zephyrichthys barryi sp. nov. A, lateral view of head, WAM P.29886-017, male, 47 mm Sl; B, ventral view of head, WAM P.
27959-004, male, 50 mm Sl; C, ventral view of male copulatory organ with hood not bent forward, holotype; D, inclined lateral view of
male copulatory organ, holotype; E, ventral view of male copulatory organ with hood bent forward, WAM P.27955-003, 50 mm Sl; F, view
of left pseudoclasper from ventral, holotype; G, lateral view of male copulatory organ with hood not bent forward, holotype; H, median
view of right otolith, WAM P.29886-017, male, 54 mm Sl; I, median view of right otolith, WAM P.29886-017, male, 47 mm Sl.
usually reaching tip of last precaudal parapophysis in
males but not in females.
Male copulatory organ (Fig. 53C–G). Two pairs of
pseudoclaspers, outer pseudoclasper large, wing-shaped
with broad base, inner pseudoclasper about half size of
outer pseudoclasper, bifurcate, each branch with supporter,
anteriorly joined to outer pseudoclasper; isthmus wide.
Penis curved, slightly longer than pseudoclasper, pointed,
with widened basis.
Colour. live colour known from three freshly caught
specimens (WAM 27951-008, WAM 27959-004, Fig. 52A,
WAM 28522-005), which all show a dark, dusky yellow
body colour, somewhat darker anteriorly and sometimes
with slightly pinkish-yellow belly. vertical fins are
102
Dinematichthyine ishes of the Indo-west Paciic III
Table 24. Meristic and morphometric characters of Zephyrichthys
barryi sp. nov.
holotype
WaM
27616010
49
Standard length in mm
Meristic characters
Dorsal in rays
86
Caudal inrays
15
Anal in rays
67
Pectoral in rays
22
Precaudal vertebrae
12
Caudal vertebrae
31
Total vertebrae
44
rakers on anterior gill arch
13
Pseudobranchial ilaments
2
D/v
7
d/a
29
v/A
16
Morphometric characters in % of SL
head length
24.9
head width
14.2
head height
15.8
Snout length
6.7
upper jaw length
13.3
Diameter of pigmented eye
2.3
Diameter of pupil
1.6
Interorbital width
6.4
Posterior maxilla height
4.3
Postorbital length
17.7
Preanal length
52.5
Predorsal length
30.8
Body depth at origin of anal in
14.2
Pectoral in length
14.2
Pectoral in base height
5.7
ventral in length
20.3
Base ventral in – anal in origin 32.3
holotype +
117 paratypes
n
Mean (range)
44.1 (20-70)
118
92.0 (86-99)
15.1 (14-16)
70.7 (67-77)
21.7 (20-23)
12.1 (12-13)
32.3 (31-34)
44.4 (43-46)
13.3 (12-15)
2.1 (2-3)
6.4 (6-7)
26.8 (24-30)
14.3 (13-16)
46
32
44
15
55
50
54
18
17
48
45
44
25.8 (24.0-27.2)
13.5 (11.9-14.6)
15.0 (13.9-15.9)
6.0 (5.1-6.7)
12.9 (12.5-13.3)
2.3 (2.0-2.5)
1.6 (1.3-1.9)
6.6 (5.8-7.7)
4.2 (3.3-4.6)
18.4 (17.2-19.7)
48.5 (45.3-52.5)
31.4 (29.9-32.7)
14.3 (13.6-15.4)
13.2 (11.6-14.3)
5.5 (4.8-6.5)
20.6 (18.2-23.6)
29.6 (26.4-32.3)
10
10
10
10
10
14
13
10
10
10
10
10
10
10
10
15
10
translucent, bright yellow at the base. Preserved colour is
medium to dark brown.
Comparison. See comparison between Zephyrichthys
and other genera.
Distribution. Zephyrichthys barryi is widely distributed
along the southern shores of Western Australia from north
of Kalbarri to esperance and including the houtman
Abrolhos and rottnest Islands (Fig. 10).
Etymology. In honour of J. Barry hutchins, Perth,
WAM, in recognition of his many contributions to the
ishes of Australia and his support in making available
the valuable material from the WAM collection.
GeOGrAPhIC DISTrIBuTION
In part I of the review of the Dinematichthyini of
the Indo-west Paciic the distribution pattern of the
very species-rich genus Diancistrus was evaluated
and discussed. It was noted that half of the 30 species
recognised were restricted to narrowly deined areas and
thus were regarded as endemic. The areas found to have
the richest endemism of the genus Diancistrus were the
NW subtropical Paciic region (ryukyus, Taiwan and
northern Philippines), and the shores of New Guinea
and certain west Paciic islands such as vanuatu, Fiji and
Tonga. Part II added to the overall notion of regionally
restricted species, this time to the south of the main reef
belt of the Indo-west Paciic. In this case the subtropical
to temperate shores of South Africa and Australia were
found to be particularly rich in such narrowly ranging
endemics.
With the present part, the overall trend of regionally
restricted distribution patterns is conirmed yet again.
All species recorded in this part are restricted to one
or two geographical areas as deined and discussed in
Schwarzhans et al.’s (2005) review of the Indo-west
Paciic Dinematichthyini. Generally, the same is true for
the genera described in this paper. The most widespread
ones are eusurculus, ranging from the Andaman Islands
along the northern shores of Australia to Ne New Guinea
and vanuatu, and Ungusurculus, which is distributed
from Singapore through Indonesia to the Philippines
and Ne New Guinea. All other genera treated herein
are restricted in distribution to three or fewer regions as
deined in Table 25 and therefore should be regarded as
endemic genera. Since endemic genera represent an older
stage of speciation in a phylogenetic sense than endemic
species (for example, Schwarzhans et al. 2005 and this
paper), their distribution patterns appear worthwhile at
this stage to analyse in more detail.
It turns out that in the Indo-west Paciic there are
only two widespread genera, namely Diancistrus and
Dinematichthys s.l. (subject, however, to the final
part of the review dealing with that genus), plus the
two subregionally distributed genera eusurculus and
Ungusurculus. All other genera are restricted to one to
three deined areas.
The most regionally restricted genera of all are
Brosmolus (NW Australia), Brotulinella (Taiwan
and the nor ther n Philippines), Dact ylosurculus
(SW Australia), Lapitaichthys (New Caledonia),
Majungaichthys (Madagascar), Monothrix (Se Australia)
and Zephyrichthys (SW Australia). Not surprisingly,
these are all monospecif ic genera. Immediately,
however, it becomes obvious that there is a particular
area emerging, which is rich in endemic genera, namely
temperate Australia (three genera).
When adding non-monospecific genera limited
to a distribution across two or three areas, this trend
receives further support for temperate Australia
(and northern New Zealand and Norfolk Island) with
Dermatopsis, Dermatopsoides and Dipulus (all three
genera from Møller and Schwarzhans (2006)), but other
areas emerge as endemic-rich as well. These are South
103
W. Schwarzhans and P. r. Møller
for the Dinematichthyini and in particular the southwestern coast with at least four endemic genera.
3. Northern Philippines and Taiwan (Brotulinella).
This area (plus the ryukyu Islands) is also very rich in
endemic species (see part I).
4. SW Paciic Islands, namely northern New Zealand
and Norfolk Island (Dermatopsis and Dipulus), New
Caledonia (Lapitaichthys and Didymothallus) and Fiji
(Dermatopsis). This area shows a strong relationship
with its composition of endemics with that of temperate
Australia (Dermatopsis and Dipulus, and Lapitaichthys,
which may be related to Monothrix). It is very likely
that they represent remnants of an originally uniform
distribution pattern that has become disrupted through
geological plate divergence, with Fiji having drifted
furthest away into tropical realms.
Africa (Dermatopsoides and Mascarenichthys), the
Mascarenes with east Africa (Mascarenichthys), NW
and Ne Australia (Beaglichthys, also extending across
the Timor Sea to Java, and Didymothallus, also extending
to New Caledonia).
As shown on Figure 54, four distinct areas are
recognised with endemic genera. They are:
1. east Africa with Madagascar and the Mascarenes
with Majungaichthys and Mascarenichthys and South
Africa south of 5° N with Mascarenichthys and south
of 30°S also with Dermatopsoides.
2. Australia with Beaglichthys, Brosmolus and
Didymothallus along its tropical shores and Dermatopsis
(also in New Zealand and Fiji), Dermatopsoides, Dipulus
(also around Norfolk Island), Dactylosurculus, Monothrix
and Zephyrichthys in southern Australia. Australia is by
far the most endemic-rich region in the Indo-west Paciic
Table 25. Geographic distribution of species of the genera of the Dinematichthyini from the Indo-west Paciic, arranged in order of treatment
(Parts I–Iv). restricted distribution patterns supposed to be endemic are shown in dark grey, other occurrences in light grey.
eastern
Indian
Ocean
Subtropical
NW Paciic
Tropical
W Paciic
Tropical
and subMicronesia
tropical
Australia
Temparate
Australia and
New Zealand
SW Paciic
Islands
east
Polynesian
Islands
red Sea, Arabia and Ne Africa
east Africa south of 05° N
Seychelles and Mascarenes
Madagascar
South Africa south of 30° S
Chagos and Maledives
Ceylon and South India
Andamans, Thailand and Sumatra
North vietnam and hainan
ryukyu Islands
Taiwan and Philippines north of 15° N
Sulu Sea and Philippines south of 15° S
Celebes Sea
Java and Indonesian Timor Seas
Banda Sea and NW New Guinea
Carolines and Palau
Marshall and Gilbert Islands
Bismarck Archip. and Solomons
NW Australia W of 142°e & N of 25° S
GBr and Se PNG
lord howe Island
SW Australia W of 145° e & S of 25° S
Se Australia e of 145° e & S of 25° S
New Zealand North Island
Norfolk Island
New Caledonia
loyalty Islands and vanuatu
Fiji
tonga
Samoa
Cook and Society Islands
Tubuai Islands
Pitcairn (incl. Ducie Atoll)
Western Indian
Ocean
Brotulinella
I Diancistrus
Paradiancistrus
Dermatopsis
II Dermatopsoides
Dipulus
Beaglichthys
Brosmolus
Dactylosurculus
Didymothallus
eusurculus
III Lapitaichthys
Majungaichthys
Mascarenichthys
Monothrix
Ungusurculus
Zephyrichthys
Iv Dinematichthys s.l.
Total number of genera
endemic genera (1-3 areas)
1 2 2 2 2 2 1 3 2 2 3 4 2 4 2 2 2 3 6 5 2 5 2 1 1 4 4 3 2 2 2 2 2
- 1 1 1 2 - - - - - 1 - - 1 - - - - 3 2 - 4 2 1 1 2 - 1 - - - - 104
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 54. Distribution of dinematichthyine genera in the Indo-west Paciic. Outline with light shading = total distribution of Dinematichthyini.
Grey shading indicates increasing number of endemic genera (see also Table 25): irst step, light grey = no endemic genera; second step
= 1 endemic genus; third step = 2 or 3 endemic genera, fourth step = more than 3 endemic genera.
From this summary it becomes obvious that the highest
degree of ‘early separated endemism’ is observed at the
periphery of the Indo-west Paciic (Fig. 54). Most of this
endemism is probably represented by primary endemics,
except possibly for that in the western Indian Ocean, which
may have been inluenced by the former tropical western
Tethyan fauna; however this idea cannot be evaluated at
present because of the sketchy fossil record.
ADDeNDuM TO PArT I OF The revIeW OF
The DINeMATIChThyINI OF The INDO-WeST
PACIFIC (SChWArZhANS eT. AL. 2005).
Paradiancistrus Schwarzhans, Møller and Nielsen,
2005
Type species: Paradiancistrus acutirostris Schwarzhans,
Møller and Nielsen, 2005 (type locality: southern coast of
epi Island, vanuatu, 16°47’S, 168°21’e).
Diagnosis. Anterior nostril placed low on snout; male
copulatory organ with two pairs of small pseudoclaspers,
the outer broad wing-shaped, inner joined anteriorly
to outer pseudoclasper forming curved feature, with
supporter; eyes large (2.0–3.0% Sl); lower preopercular
pores 1; precaudal vertebrae 11; parapophyses very
wide; head with narrow scale patch on cheek, no scales
on operculum; otolith with inclined (5–10°) and short
sulcus (otolith length to sulcus length 2.4–2.6), colliculi
fused; maxillary expanded at rear corner; anterior anal
in pterygiophore long.
Remarks. The genus Paradiancistrus was described
by Schwarzhans et al. (2005), based on two species
– P. acutirostris Schwarzhans, Møller and Nielsen, 2005,
from vanuatu and P. cuyoensis Schwarzhans, Møller and
Nielsen, 2005, from the visayan and Palawan Islands of
the Philippines. The ongoing review has revealed another
species of this rare genus, which was not recognised
previously. The description of this new species is therefore
added as an addendum at the end of the current third part
of the review of the Dinematichthyini of the Indo-west
Paciic.
Paradiancistrus lombokensis sp. nov.
(Figs 55–57; Table 26)
Material examined. (2 specimens, 20–35 mm Sl).
hOlOTyPe – SMNS 18662, male, 35 mm Sl, 8°30’S,
116°02’e, western shore of lombok, Indonesia, depth
0–2 m, 12 December 1996. PArATyPe – SMNS 26369,
1 juvenile, 20 mm Sl, same data as holotype.
Diagnosis. vertebrae 11+ 29-32 = 40-43, dorsal in rays
65 (holotype), anal in rays 52 (holotype); head length 23.8–
25.2 % Sl; head height 13.3–14.4 % Sl; predorsal length
Fig. 55. Sample sites of Paradiancistrus acutirostris, P. cuyoensis
and P. lombokensis sp. nov. One symbol may represent several
samples.
105
W. Schwarzhans and P. r. Møller
Fig. 56. Paradiancistrus lombokensis sp. nov. SMNS 18662, holotype, 35 mm Sl.
30.0–31.1; lower preopercular pores 1; outer pseudoclasper
broad wing-shaped, short; inner pseudoclasper a double
lobed broad lap, anteriorly joined to outer pseudoclasper
forming curved feature, with supporter; scale patch on
cheek with eight scale rows on upper cheek and four scale
rows on lower cheek; head with blunt snout.
Description (Figs 56, 57). The principal meristic and
morphometric characters are shown in Table 26. Body
slender; mature at about 35 mm Sl (male with fully
developed pseudoclaspers). head with patch of small scales
on cheek (8 scale rows on upper and 4 on lower cheek);
no scales on operculum. horizontal diameter of scales on
body about 1.3 % Sl, in 21 horizontal rows. Maxillary
ending far behind eyes, dorsal margin covered by dermal
lobe of upper lip, expanded posteriorly, with prominent
angle at ventral rear corner. Anterior nostril placed low
on snout, about 1/4 distance from tip of snout to anterior
margin of eye. Posterior nostril moderately small, about
1/3 the size of eye. Tip of opercular spine free, pointed.
Anterior gill arch with 15 rakers, thereof 3 elongate rakers.
Pseudobranchial ilaments 1.
Head sensory pores (Fig. 57 A–B). Supraorbital pores
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 mandibular pore without cirrus. Preopercular
pores 2 (1 lower and 1 upper), both tubular. [See description
of Beaglichthys bleekeri for position of pores.]
Dentition (of holotype). Premaxilla with three outer
rows of granular teeth and one inner row of larger teeth
anteriorly. Anteriormost teeth in inner row up to 1/3
diameter of pupil. vomer horseshoe-shaped, with two
rows of small teeth 1/4 diameter of pupil. Palatine with two
rows of small teeth up to 1/5 diameter of pupil. Dentary
with four outer rows of granular teeth and one inner row
of larger, teeth anteriorly, merging into one row of larger
teeth posteriorly, up to size of pupil diameter.
Otolith (Fig. 57e–F). elongate, length to height 2.2–2.3
(20–35 mm Sl); otolith thin (otolith height to otolith
thickness 2.5); otolith length to sulcus length 2.2–2.5;
sulcus inclined at 5°. Anterior tip of otolith pointed,
posterior tip expanded. Dorsal rim with rounded predorsal
and sharply pointed postdorsal angles, marked concavity
above anterior tip and behind postdorsal angle; ventral rim
gently and regularly curved, deepest at about its middle.
Inner face convex, outer face concave, both smooth. Sulcus
positioned slightly anterior of the middle of the inner face,
colliculi fused. ventral furrow distinct, close to ventral rim
of otolith, slightly turning upwards towards its tips.
Axial skeleton. Neural spine of vertebrae 4–5 inclined
and 6–7 depressed. Parapophyses present from vertebrae
6 to 11. Pleural ribs on vertebrae 2 to 11. First anal in
pterygophore elongated, reaching tip of last precaudal
parapophysis in males.
Table 26. Meristic and morphometric characters of Paradiancistrus
lombokensis sp. nov.
holotype
Paratype
SMNS 18662 SMNS 26369
Standard length in mm
35
20
Meristic characters
Dorsal in rays
65
Caudal inrays
Anal in rays
52
Pectoral in rays
17
Precaudal vertebrae
11
11
Caudal vertebrae
29
32
Total vertebrae
40
43
rakers on anterior gill arch
15
Pseudobranchial ilaments
1
D/v
7
d/a
20
v/A
14
Morphometric characters in % of SL
head length
23.8
25.2
head width
10.7
10.4
head height
13.3
14.4
Snout length
4.3
4.4
upper jaw length
11.7
12.9
Diameter of pigmented eye
2.0
2.7
Diameter of pupil
1.2
1.5
Interorbital width
5.0
6.1
Posterior maxilla height
3.7
3.0
Postorbital length
18.0
18.4
Preanal length
49.9
46.8
Predorsal length
31.1
30.0
Body depth at origin of anal in
14.1
16.8
Pectoral in length
14.9
12.5
Pectoral in base height
4.8
5.0
ventral in length
17.3
Base ventral in – anal in origin
32.6
29.0
106
Dinematichthyine ishes of the Indo-west Paciic III
Fig. 57. Paradiancistrus lombokensis sp. nov. holotype. A, lateral view of head; B, ventral view of head; C, view of left pseudoclasper
from inside; D, inclined lateral view of male copulatory organ; E, median view of right otolith; F, ventral view of right otolith.
Male copulatory organ (Fig. 57C–D). Two pairs
of rather small pseudoclaspers. Outer pseudoclasper
broad wing-shaped, short, with thick supporter; inner
pseudoclasper double-lobed moderately broad f lap,
anteriorly joined to outer pseudoclasper forming curved
feature, with supporter; isthmus moderately wide; penis
short, curved, with broad base.
Colouration. live colour unknown. Preserved colour
medium brownish grey, lighter at the belly and darker
on the back in front of the dorsal in.
Comparison. Paradiancistrus lombokensis is
readily recognised as a representative of the genus
Paradiancistrus because of its single lower preopercular
pore, a character which outside of the Indo-west Paciic
is only shared by Pseudogilbia Møller, Schwarzhans
and Nielsen, 2004, from America, which however
exhibits a different pseudoclasper pattern amongst other
distinguishing characters (see Møller et al. (2004) and
Schwarzhans et al. (2005)).
Paradiancistrus lombokensis differs from the two
other species of the genus – P. acutirostris and P.
cuyoensis – in the low number of dorsal in rays (65 vs
82–88 and 76–81 respectively) and anal in rays (52 vs
66–71 and 62–65 respectively), the short and narrow
head (head length <25.5 vs >27 % Sl, head height <15
vs > 20 % Sl), the short predorsal length (<31.5 vs >35
% Sl), the broad scale patch on the cheeks with 8 scale
rows on the upper cheeks (vs 2 and 5–7 respectively)
and in details of the otolith and the pseudoclasper
morphology. The high number of scales on the cheeks,
the inner pseudoclasper with two lobes and the stubby
snout resemble P. cuyoensis.
Distribution (Fig. 55). Paradiancistrus lombokensis
is known only from two specimens from the typelocation at lombok, Indonesia. The type series was found
in crevices of silty coralline rock.
Etymology. Named after the type locality, the island
of lombok.
ADDeNDuM TO PArT II OF The revIeW OF
The DINeMATIChThyINI OF The INDO-WeST
PACIFIC (Møller AND SChWArZhANS 2006).
Dermatopsis Ogilby, 1896
Species. Following the review of Møller and
Schwarzhans, (2006), the genus Dermatopsis comprises
four species – Dermatopsis greenfieldi Møller and
Schwarzhans, 2006, from Fiji, Dermatopsis hoesei Møller
107
W. Schwarzhans and P. r. Møller
Dermatopsis greenieldi Møller and
Schwarzhans, 2006
(Fig. 58)
Additional material examined. (6 specimens, 15–51
mm Sl). NhMG 1917-07-07, 1 male, 36 mm Sl, Fiji station
6; NhMG 1917-07-02, 1 male, 39 mm Sl, 3 females, 40–51
mm Sl, 1 juvenile, 15 mm Sl, Fiji station 13.
Diagnosis (updated). vertebrae 11-12 + 27-29 = 39–41,
dorsal in rays 64–70, anal in rays 44–49; scales present
on body; eye 1.8–2.6 % Sl, sharp spine on ventral maxilla
positioned behind rear tip of eye; large, broad single
pair of (outer) pseudoclaspers, inwardly connected
to isthmus with ligament, with two supporters, the
anterior inclined, short, joined at base to massive and
long posterior supporter; otolith with pointed posterior
tip and weak postdorsal angle, otolith elongate, length to
height ratio 2.1–2.2, sulcus with separated colliculi.
Description (addendum). Male copulatory organ (Fig.
58). Single pair of larger, broad (outer) pseudoclaspers
anteriorly connected to wide isthmus with ligament. Two
supporters, anterior one forwardly inclined, thin, connected
at base to posterior supporter; posterior supporter longer,
more massive, with slightly expanded and pointed tip. Penis
curved, somewhat longer than pseudoclasper.
Remarks. The presence of two supporters in the single
pseudoclasper is (so far) unique among pseudoclaspers of
the closely related genera Dermatopsis, Dermatopsoides
and Schwarzhans, 2006 and Dermatopsis macrodon
Ogilby, 1896, both from south-eastern Australia and
Dermatopsis joergennielseni Møller and Schwarzhans,
2006 from New Zealand. At the time of description, no
males were known from D. greenieldi. Such males have
now become available from the collection of NhMG in
Göteborg, Sweden. They are igured and described below
and also are the reason for the updating of the diagnosis
of the genus (new information in bold).
Diagnosis (updated). Anterior nostril placed very
close above upper lip; head without scales; tip of
opercular spine free, exposed; maxilla not vertically
expanded postventrally, ventral knob well anterior to
rear corner; lower lip with skin folds; male copulatory
organ with one pair of mostly large, but not very
diverse pseudoclaspers (probably representing the outer
pseudoclasper in terminology of Møller et al. 2004a) with
usually one supporter (two in
); penis
without hook near tip; sulcus of otolith with separated
ostium and cauda marked by strong indentation at ventral
margin of sulcus, its colliculi separated; anterior anal in
ray pterygiophore elongate; lower preopercular pores 2,
often joined in single opening and then counted as 1;
upper preopercular pore absent; posterior infraorbital
pores 2 or 3; precaudal vertebrae variable between 11
and 14, body size not exceeding 75 mm Sl.
Fig. 58. Dermatopsis greenieldi Møller & Schwarzhans, 2006. NhMG 1917-07-07, male, 36 mm Sl. A, ventral view of male copulatory
organ; B, inclined lateral view of male copulatory organ; C, view of left pseudoclasper from inside; D, view of left pseudoclasper from
outside.
108
Dinematichthyine ishes of the Indo-west Paciic III
and Dipulus (bearing in mind, however, that pseudoclaspers
are not yet known from Dermatopsoides morrisonae Møller
and Schwarzhans, 2006) and may indicate a plesiomorphic
character state. Single outer pseudoclaspers with two pairs
of supporters have otherwise only been observed from the
New World genus Gunterichthys Dawson, 1966 and the
herein newly described genus Didymothallus.
The pseudoclasper morphology of D. greenfieldi
strongly resembles that of D. macrodon except that in
D. macrodon, the equivalent of the thin anterior supporter
as seen in D. greenieldi, is developed as a leshy appendix
without a supporter.
ACKNOWleDGMeNTS
We wish to thank the following persons for helping us
with material and information: Gerald r. Allen (WAM),
M. eric Anderson (SAIAB), Dianne J. Bray (NMv), David
Catania (CAS), Daniel M. Cohen (CAS), Gavin Dally
(NTM), Guy Duhamel (MNhN), Jon Fong (CAS), ronald
Fricke (SMNS), Kiyoshi hagiwara (yCM), Karsten hartel
(MCZ), Philip C. heemstra (SAIAB), Peter A. hulley
(SAM), J. Barry hutchins (WAM) (also for the photos of
the fresh dead specimens), 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), John
e. McCosker (CAS), Mark A. McGrouther (AMS), Sue
Morrison (WAM), 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), Mark Sabaj (ANSP), Jeff Seigel (lACM), Gento
Shinohara (NSMT), Carina Sjöholm (NhMG), David G.
Smith (uSNM), William F. Smith-vaniz (uSGS), victor
G. Springer (uSNM), Andrew l. Stewart (NMNZ), Tom
Trnski (AMS), Jeffrey T. Williams (uSNM) and richard
Winterbottom (rOM).
Also thanks are due to the following colleagues
at ZMuC: 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.
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110